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OKO) ' INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION 
| RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS 


Volume XXXIV | January 20, 1989 No. 1 


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ISSN 0093-4666 MYXNAE 34(1) 1-276 (1989) 


Published quarterly by MYCOTAXON, LTD., P. O. Box 264, Ithaca, NY 14851. For 
subscription details, availability in microfilm and microfiche, and availability 
of articles as tear sheets, see inside back cover. 


This Festschrift number was generously subsidized by the authors or 
their institutions or employers, which will permit this volume 
of MYCOTAXON to include approximately 800 pages 
without additional cost to subscribers. 


SEE ERRATA 


TABLE OF CONTENTS 


Eh Pre tape aie ca at ater ELC Ue TL Stsi ars wiatalives ix) at alia! sta el 
Harry D. Thiers: Reminiscences about a teacher and friend ............... 
Remembering the morel grower: Ron Ower, 1939-1986.............+--- 
Dermocybe, subgenus Dermocybe, section Sanguineae in northern 
SRATISOLT iLiad net uses Le pace deride Sates Lay steerer we Joseph F. Ammirati 
Lichens of Mount Diablo State Park, Contra Costa County, California. 
Doris E. Baltzo 
The genus Rhodocybe: new combinations and a revised key to section Rhodo- 
phana in North America.. Timothy J. Baroni and David L. Largent 
Quality control factors for Alternaria allergens. 
Harriet A. Burge, Marion E. Hoyer, William R. Solomon, 
Emory G. Simmons, and Janet Gallup 
Thaxterogaster thiersii: a new secotioid species from California. 
Cornelia J. Calhoun 
Studies on Marasmius from eastern North America. II. New species. 


Dennis E. Desjardin and Ronald H. Petersen 
A synopsis of Colombian boletes...............---: Roy E. Halling 
Cladonia thiersii: a new lichen from California......... Samuel Hammer 


Studies in Agaricus IV: new species from Colorado.. Richard W. Kerrigan 
A new, lignicolous species of Entoloma (Entolomataceae, Agaricales) from 


BEATE OLTIGUAE LE has ocd atk ey dl ote ok tab wate a allah gay a tein es David L. Largent 
Motes on the genus Protubera -.. . oi. 5s owe ee ee es David Malloch 
Notes on Clavariadelphus. II. New and noteworthy species from North 

NCR ICOM re ee re ie ena a eae gta etait Andrew S. Methven 
A key to the species of Inocybe in California....... Florence H. Nishida 
Foliicolous fungi 8: Capnodium in California......... Don R. Reynolds 
A new species of Gymnopilus from northem California.. Michelle T. Seidl 
The lichens of serpentine rocks and soils in California... .. Lorene L. Sigal 


Lepiota sensu lato in California. III. Species with a hymeniform pileipellis. 
Walter J. Sundberg 
Examination of Stilbothamnium togoense for Aspergillus flavus group 
TEVCOLOXINS tctats ie. soy D. T. Wicklow, R. F. Vesonder, Cesaria E. 
McAlpin, R. J. Cole, and Marie-France Roquebert 
The occurrence of vesicular-arbuscular mycorrhizae in burned areas of the 
Snake River Birds of Prey Area, Idaho..... Marcia Wicklow-Howard 
Compatibility and fruiting studies of an albino form of Auricularia cornea. 
George J. Wong 
Master’s degree students of Harry D. Thiers and thesis titles............-. 
Type specimens of agarics, boletes and gasteromycetes in the San Francisco 
State University herbarium (SFSU). 
Barbara M. Thiers and Roy E. Halling 
Species named for Harry D. Thiers...........-- . Dennis E. Desjardin 


115 
119 


129 
133 


153 
181 
197 
2A 
221 


MeN, 


249 
253 
209 
267 


269 
276 


PREFACE 


January 22, 1989 marks two occasions in the life of Harry D. Thiers, his 70th 
birthday, and more importantly, his retirement from 30 years of teaching at San 
Francisco State University. A teacher often measures his success by that of his stu- 
dents, therefore we hope that this volume of 20 research papers on mycology and 
lichenology by his students will bring Harry pleasure and a sense of accomplish- 
ment. 

Festschrift projects often have the reputation of being far more time- 
consuming and difficult than the organizers envision at the outset. We admit to 
spending more time on this one than we thought we would, even with the help of 
XyWnite III + and a laser printer. However, we have found the rewards of the pro- 
ject to be far greater than the time invested; we have renewed contact with many 
old friends, chuckled over forgotten anecdotes, and expanded our mycological and 
editorial horizons. 

In the process, we learned something about Harry Thiers as well. . .he must be 
one of the world’s most inquisitive people. We resolved to keep this Festschrift 
secret from him as long as possible. During our semi-annual visits to San Francis- 
co over the last two and a half years we have tried many times to accomplish 
Festschrift-related business through a furtive poke in the herbarium or a hushed 
conference in the corner. More often than not, the Festschrift Honoree would 
catch us in the act, and some fancy footwork was needed to misrepresent our ac- 
tivities convincingly. Only Ellen Thiers proved to be cunning and cool-headed 
enough to succeed in ferreting crucial data from under Harry’s nose. 

In the introductory section to this volume, many fine and well-deserved 
tributes are paid to Harry. We know he would want us to reiterate here something 
he has said many times: he feels extraordinarily lucky to have had such a fine as- 
sortment of students. They are every bit as special to him as he is to them. 

We wish to give our warmest thanks to all of the Festschrift participants. Spe- 
cial thanks are also due to Ernst Both, Director, Buffalo Museum of Science, for 
donating the cover artwork (Leccinum manzanitae Thiers) executed by Patricia 
Eckel, and to The New York Botanical Garden for helping to manage finances of 
the project. Generous financial contributions from W. P. Jordan, B. K. Tamm, and 
an anonymous donor are very much appreciated. Richard P. Korf, Managing 
Editor of MYCOTAXON was extremely helpful and accommodating at every step 
of the project, and we are very grateful to him for making this volume possible. 


Roy E. Halling 
Barbara M. Thiers 
Guest Co-editors 
Bronx, NY 


HARRY D. THIERS - REMINISCENCES ABOUT A 
TEACHER AND FRIEND 


I served as laboratory assistant for HDT beginning in my sophomore year at 
Texas A & M in College Station, Texas. Harry supervised two beginning botany 
labs simultaneously. During the laboratory on stems, I was charged with giving the 
introduction to the class in one of the labs. I confidently showed the class where 
the cambium was located in a corn stem. HDT came into the room and very 
courteously took over the class, and relectured monocot stems from the same 
chart. I was very embarrassed, but as usual, he was very patient, and on we went. 
My first mycology course was with Harry. He was an excellent instructor, and the 
laboratories were very well planned. The lab with the water molds was amply sup- 
plied with examples of discharging zoosporangia. This was the first time I had seen 
the discharge and the zoospores swimming about. I stayed a long time looking at 
this material, and got so excited that I had to go home to bed to recover. [Don 
Reynolds]. 

x * * KK * 

As an undergraduate at San Francisco State, I was assigned an advisor in the 
Department of Biology, one Robert Sweeney. I was something of an annoyance to 
him because I continually brought mushrooms I'd collected in the environs of the 
campus to our scheduled meetings. Bob’s response, invariably was, "I don’t know 
anything about those damn things." One afternoon, Bob stopped me in the hall 
and informed me that the department had just hired a mycologist, and I should 
talk to him about those damnable mushrooms. 

Bearing a freshly collected specimen, I promptly presented myself to Harry 
Thiers. Harry, looking a bit frazzled, was busy emptying many boxes of books. 
Nevertheless, he greeted me and my mushroom with great enthusiasm. As is so 
typical of him, rather than simply identifying thé mushroom for me, Harry feigned 
ignorance, reached into his collection of books, and introduced me to the 
dichotomous key. We keyed out the mushroom together. 

Harry’s response to me and my interest was all the encouragement I needed. 
Almost daily I would bring in specimens to identify. Soon, we began to take short 
trips together looking for fungi. The very next semester, Harry set me up in the 
tiny back room to his office. I was provided with a microscope, a bottle of Mel- 
zer’s, a straight razor, and a mimeographed copy of Alex Smith’s "Preliminary key 
to the Fleshy Fungi." I remember Harry taking great delight watching me trying to 
section a minute Mycena without simultaneously removing the end of my finger. 

_ We took many trips then. . . .Land’s End, Crystal Springs reservoir, the 
Presidio, Skyline Drive. .. anywhere a mushroom might grow. Each trip would 
end with a mini lecture on morphology or taxonomy, and, in a pattern that typifies 
Harry’s lab even to this day, hours of keying out mushrooms. These were exciting 
times for me and irreversibly directed my path to a career in mycology. [Jerry 
Motta] 


* x xX kK * * 


In the later 1950’s, the science faculty of San Francisco State required all 
Biological Science students--regardless of their major interest--to experience a few 
Botany classes. As a neophyte biological science major with a presumptive interest 
in zoology, Botany would not have been one of my elective choices. The required 
first semester course did little to change my mind and convinced me to complete 
my botany requirements as quickly as possible by taking the second semester of the 
introductory course rather than the alternative--an upper division Botany elective 
at a later date. é: 

Walking into that second semester Botany class with only 14 other students in 
the room, when there had been nearly 50 in the first semester course, I quickly be- 
gan to wonder what I had gotten myself into! The first day’s lecture in the class 
consisted of an overview of what we were going to cover during the semester, with 
review, wherever it fit in, of terms and concepts we had "learned" in the previous 
course. Unusual? No. But, Dr. Thiers, the instructor, who had recently come to S. 
F. State from Texas, didn’t just lecture. He started almost immediately to involve 
the class in the process by requesting definitions, asking questions, etc. And, much 
to my surprise, he was calling on people that first day in class by name--often 
without even turning around from the blackboard!! (I learned later that many of 
the others in class had been in Professor Thiers’ laboratory section the semester 
before--I had not). Realizing that my learning of the first semester’s material was 
less than adequate for this approach, I became quickly aware that if I was to sur- 
vive this experience, something had to be done--and fast!! Extra Botany books 
were obtained from the library; notes were recopied, embellished for clarity, and 
repeatedly reviewed; vocabulary lists, constantly being upgraded, were always in 
hand. Botany between classes, Botany at meals, Botany on the bus commuting to 
and from work. Sometime early in the semester, the "chore" of studying became a 
joy. Through Dr. Thiers’ enthusiasm and excitement, Botany came alive for me in 
that class and my life was forever changed. 

In my graduate school days at San Francisco State in the mid-1960’s, the her- 
barium was a home away from home. Converted from a small storeroom in the 
original square Science building, it served as the Botany graduate student office, 
work-study area, and general meeting place. The five to six graduate students had 
work spaces sandwiched between, around, and behind the seven or eight her- 
barium cases; one had a desk looking into the unused freight elevator door! Close- 
ness created life-long friendships. Often arriving before others in the morning, 
HDT was always there to answer our questions, offer advice and support, and 
share in the excitement of our discoveries. Like the coffee always brewing, the 
love of Botany, of science, and of knowing, was rich there. [Walter J. Sundberg] 

* x eK KX * * 

I first met Dr. Harry Thiers in 1960 when I left a high school teaching job to 
further my education. It was a fortuitous meeting for me! During 1960-1961, he 
was the nucleus of my forming career, and ever since then, the impression of his 
wonderful enthusiasm for everything, especially fungi and students, has helped me 
shape my teaching and research career. 

When I came to SFSU in 1960, I hoped that I could get a masters degree, pos- 
sibly in marine algae under Dr. George Oberlander. My chagrin at learning that 
he was on sabbatical leave (in Australia) was quickly alleviated when I met Dr. 
Thiers. Although his primary interest was fungi, he was flexible and quite 
amenable when I came up with a plan on ferns. So that fall term, I began growing 
fern gametophytes on liquid media and testing their responses to various levels of 
nitrogen and other inorganic nutrients. Things finally jelled, and by the end of the 
academic year I wrote a thesis and underwent an oral exam to get my degree. Dr. 


Thiers was tremendous in the help he provided, even though he was far from an 
expert on fern gametophytes. 

In the fall, I took his mycology course, in which the labs were superbly full and 
well-organized--a multipaged guide for each lab. The spring phycology course was 
another matter. Dr. Oberlander was nowhere around, so it fell to Dr. Thiers to 
teach the phycology course that spring. It was a joy! I have never enjoyed a course 
as much. Dr. Thiers gave the standard lectures, but the labs were for us to identify 
and learn about as many freshwater algae, and then marine algae, as possible. 
And I still feel that identifying organisms is a wonderful way to learn about them. 
Well, that phycology course in the spring of 1961 convinced me, without question, 
that I was destined to become a phycologist. In fact, unless my memory fails me, I 
think that it was when Dr. Thiers took the class on a field trip to Monterey that I 
got irreversibly turned-on to marine algae. That is where I still am today, 27 years 
later. [Bill Johansen] 

xx ** kX * * 

In the fall 1961, Harry Thiers, 
two graduate students, and the 
mycology class made a weekend 
field trip to Jackson State Forest in 
Mendocino. Through the kindness 
of the California Division of 
Forestry, Harry and his students 
stayed in a wooden ramshackle 
building at the old fire station on 
Little Lake Road. The wooden fa- 
cility consisted of a large 
dormitory-like room, a rustic wash- 
room, and a large kitchen. I have 
fond memories of the kitchen for it 
was there that we would put out 
our collections on a wooden table 
and review identifications togeth- 
er. Harry would answer all of our 
questions with patience and would 
repeat the answers as often as 
necessary. It was in that room that 
he was the ultimate teacher; an 
enthusiastic and wonderfully in- 
vigorating person. 

One of the most memorable 
experiences with Harry happened 
in the dormitory. Each side of the 
dormitory was lined with metal 
bunks-- two bunks to a tier--each 
bunk with metal springs that had 
absolutely no give to them, and a 
nearly non-existent, thin, hard, 
bumpy mattress. In that room, 
Harry and all of his students would 
roll out their sleeping bags on 


Demonstration of wave action on Nereocystis. 


those bunks. One of our most favorite pastimes was to collect mushrooms after 
dinner by flashlight in the dark. After one such lovely collecting trip during which 
we had spent the evening becoming exhilarated by the sights of the gloriously 
shaped and beautifully colored mushrooms illuminated in the night by flashlight, 
we returned to the fire station in a festive mood. We had been kidding Harry the 
entire evening and he was giving as much as he was taking; however, we thought 
we were getting the upper hand. After we returned, Harry went to the kitchen, so I 
decided to take this golden opportunity to collect at least 100 Douglas Fir cones 
and put them inside Harry’s sleeping bag. All of the students knew what was hap- 
pening and as a group we decided to really badger Harry, including throwing Rus- 
sulas at him in the kitchen. Eventually all became quiet and all retired to bed; of 
course with Harry the last to arrive. In silent anticipation, we waited for Harry’s 
response as he climbed into his sleeping bag full of cones. All of us shouted glee- 
fully as Harry let out numerous loud, uncommonly used verbs after getting into his 
bag. We all laughed uncontrollably as he got up and proceeded to empty his bag 
outside. Eventually it became sufficiently calm to go to sleep. I was snuggled deep 
in my bag with the top pulled over my head when at two o’clock I was un- 
ceremoniously shoved into the bottom of my bag as someone proceeded to lift the 
bag into the air with me still inside. I struggled frantically but to no avail and felt 
myself thrown into the air and then landing with a thud on some branches. After 
about five minutes of vigorous struggling to get out of my bag, I finally extricated 
myself. All of the time there was this furious uproar from inside the dormitory. As 
I looked up into the window through which I was thrown, the first person I saw was 
Harry Thiers, the person who had picked me up and thrown me out the window. 
With a jovial laugh and a twinkle in his eye, Harry said, "See, Dave, I told you I 
would get you back!" [David L. Largent] (Now we know why HDT later chose to 
sleep in his car. Eds.) 
** * * * * 

I had just returned to San Francisco for Fall registration in 1961 and was in- 
formed that I needed an upper division elective in Botany as a graduation require- 
ment in Biological Sciences. I didn’t view this as a critical career decision because 
the main career tracks at S.F. State College were elementary and secondary educa- 
tion. Holding to my undergraduate view that nothing should interfere with 
afternoon athletic practice, I signed on for Mycology because it was offered in the 
morning and my other option wasn’t. At our first class meeting, I sensed some- 
thing was different about this group. Not only were the two graduate assistants 
(David Largent and Harriet Burge) absolutely ‘lunatic fringe’ about the fungi, but 
some of those taking the course (e.g., Jerry Motta) were similarly afflicted and had 
been preparing months in advance. Fortunately for me, HDT knew how to guide 
the rookies while continuing to challenge the ‘fringe.’ Classwork lost the dimen- 
sion of time. By November, when our Cross Country team won the Far Western 
Conference Championship at Reno, Nevada, there I was exposing Martin’s Rose 
Bengal agar plates for airborne fungal spores at the Washo County golf course. 
My teammates played a practical joke by placing rabbit dung in some of the plates, 
a sign of things to come. 

That winter, as rains came to northern California, the mushroom collecting as- 
signments began in earnest. Two women in HDT’s life, Ellen and Barbara (age 6, 
then), were present for many of the local collecting trips. Barbara could accurately 
name, to species, the different mushrooms we struggled to identify using taxonomic 
keys. Ellen collected various cryptogams of interest and worked in the Herbarium 
across the hall. How very thoughtful were Ellen’s efforts at providing refreshments 
for special occasions. This class was becoming a family. It soon became apparent 


to me that HDT’s extended family was also around evenings and weekends. This 
was not the 9-5 world of my parents and neighbors. HDT didn’t play college 
professor, he lived it. 

As other memories fade, one event remains fixed in my mind. Mendocino was 
the big December collecting trip and HDT guaranteed that we would complete our 
required collections of fleshy fungi. The drive was several hours duration and we 
arrived well after nightfall. Sleeping bags, collection baskets, drying racks, and 
kitchen supplies were unloaded and I looked around for a bunk. Then at 9:30 PM, 
the ‘fringe’ got out their flashlights and went outside to collect mushrooms. In a 
few minutes, they returned with dozens of Laccaria laccata that were found fruiting 
on the forestry station grounds. David Malloch (then a music major) and I shared 
a similar look of disbelief. Come on now, how could anyone be that hooked on the 
fungi? [Donald T. Wicklow] 

* x * * * * 

My first contact with Harry was as a very unsuccessful undergraduate student 
taking my first botany course in my senior year. As usual, I was starting the year 
off in a new major and found that as a prospective, if somewhat unenthusiastic, 
entomologist, I was required to take botany. BOTANY!! Of all the balderdash 
that I might be subjected to, I could not figure out why I needed botany. To my 
surprise, botany, as presented by Harry Thiers, was the most interesting course I 
ever took. Harry actually singled me out and invited me to go with the mycology 
class on a weekend field trip to Mendocino, which, along with other later Thier- 
sians, converted me to mycology. The field trip was especially notable for the 
quantity of fungi encountered and Harry’s unbounded enthusiasm for them. This 
field trip was also notable because it was the weekend that a well-known organiza- 
tion of mycophiles discovered the bounties of Mendocino. As the members of this 
club gleefully filled the trunks of their cars with mushrooms, Harry speculated out 
loud on a number of topics relating to this discovery. 

Later I was tied down by an evening and weekend job and could not attend 
these wonderful outings. Instead, I absorbed what I could in the lab, which of 
course, was much. I discovered that Harry could not remove a glass of water in- 
verted on the table near his manuscript on boletes without removing all of the 
papers and then spilling the water. It was a fascinating discovery until I found that 
he seemed to have no ability whatsoever for preserving marine algae, and instead, 
stored these on my desk next to my notes. 

In spite of this inability to properly dispose of algae, he was remarkably able to 
identify such things; indeed, he was able to identify algae, mosses, lichens, various 
sorts of vascular cryptogams as well as a long list of fungi. This indiscriminate in- 
terest in all sorts of cryptogams rubbed off on his students, including me, and 
sentenced most to a lifetime of teaching lower division classes that botanists usual- 
ly avoid because of their more modern specialized training. A sentence maybe, 
but with Harry as a model, something of a challenge as well....and maybe even a 
reward. I hold Harry directly responsible for the fact that I get to spend two 
wonderful weeks each August in New Brunswick teaching Marine Biology while 
my colleagues at home are attending the year’s first staff meetings. 

Staff meetings.......who of Harry’s students can forget Harry after he had been 
to a staff meeting? Red faced, breathing rapidly and muttering epithets to himself 
or muttering epithets loudly to any student who was foolish enough to confront 
him. After a staff meeting, his rate of speech increased several-fold. Lectures that 
normally lasted an hour could be delivered in 20 minutes, pictures and all, and thus 
had to be followed by the next day’s lecture to fill in the time. This could possibly 
have been avoided if we had asked a lot of questions, but who would have dared 


ask Harry a lot of questions after a staff meeting? There was at the time, a pear- 
shaped boy, who did ask questions at the wrong time and sometimes bore the brunt 
of the after-meeting effusions. This was the same kid who had his mother phone 
Harry to check out the dangers inherent in going to Mendocino and who finally up- 
set the authorities to the extent that they threatened withdrawal of cooperation. 
This kid bothered Harry more than the one who cruised rapidly up and down the 
main street of Mendocino in a flatulent Volvo with SFSC boldly written on the 
back window. 7, 

All of Harry’s students have such recollections and fond memories. From San 
Francisco they have gone out into the world and have maintained, consciously and 
unconsciously, the spirit they received from him. They remain a close-knit group 
that reassembles whenever possible to exchange the stories, experiences and 
knowledge that they possess because of their short time with this man. I am grate- 
ful and proud to be a member of this exclusive club. [David Malloch] 

xe KX Ke kK * 

In 1964, as a biology major in my junior year at SF State, I enrolled in a class 
called Cryptogamic Botany. I am pretty sure I had no idea what a "cryptogamic" 
was, but with the real and imagined demands on my time, I fit classes into avail- 
able time slots. It was there that I first experienced Harry D. Thiers, seductive 
master of the well-planned lecture. He almost never used notes. I recall one time 
when he actually apologized for their use, while extracting a single 3 by 5 notecard 
from his shirt pocket, notations on one side only. 

Harry also commanded amazingly interesting laboratories; the aura of which I 
struggle to attain in my own classes today. Components of the essential ambiance 
include somewhat untidy conditions created by stray lichen fragments, crumbled 
sheets of waxed-paper and discreet mounds of woodland debris, as well as the 
lingering fragrance of drying mushrooms; but most importantly, the drama of dis- 
covery was always present. 

For me, there are many characteristics which comprise the image of HDT. 
The most compelling, and the one that establishes him as a world class mentor, is 
the natural ease with which he makes plants alive and students important. [William 
Paul Jordan] 

x** * kk * 

It is important and valuable to reflect, from time to time, on how one began 
his professional career. Certainly for a researcher and/or teacher, or any other 
professional, there are several factors to consider and weigh, one against the other, 
in determining just how it is that the present situation came about. Typically, a 
person is influenced by one, or perhaps a few outstanding individuals, who he has 
encountered during his early years. 

For myself, I can say with certainty, that Harry D. Thiers was most influential 
in my becoming a teacher and a researcher. He was among the best teachers that I 
had as a university student, and his enthusiasm for learning, and teaching others, 
greatly influenced my decision to become a teacher. He also was my first adviser 
at San Francisco State. I am exceedingly grateful to Harry for his influence and 
guidance during my years at SFSU! 

Harry Thiers introduced me to mushrooms and his area of research, even 
though I was not "officially" one of his students. He even selected the genus 
Cortinarius for me to work on. A genus that I have been struggling with for some 
twenty years, and something for which I will never forgive him! 

It happened something like this: When I was an undergraduate student in 
Biology, I asked Dr. Thiers if I could informally attend his mushroom taxonomy 
course to learn a few basic things about fleshy fungi and absorb a few mushroom 


names. Of course, he said that would be fine. Actually, I was intimidated by the 
mushrooms, the seemingly knowledgeable graduate students, and even Professor 
Thiers himself. I learned a few names, somewhat slowly I admit, and realized very 
quickly that these organisms were indeed poorly known and difficult to identify. 

During the class, I heard something about mushroom descriptions and how im- 
portant they were in making a final and correct determination of a collection. So I 
decided to ask Professor Thiers how to prepare one. He quickly and efficiently got 
the idea of a mushroom description across to me, but it wasn’t until the following 
summer, when I was in Idaho, that I actually tried to write a description of my own. 
Looking at my notes from that period clearly shows that they are practically use- 
less. However, it did get me interested in mushroom taxonomy, and so I decided 
to try and write a few more descriptions to improve my skills. 

An approach that seemed logical to me at the time, was to work on one genus 
for awhile, and then on another, to learn something about each one. So I went to 
Professor Thiers and asked him which genus might be a good one to start with in 
terms of doing descriptions. Of course, I made it clear to him that I would do this 
as a learning experience, and that he could have the collections and notes if they 
were useful. He agreed that this was a good idea. He thought about an ap- 
propriate genus for a few seconds, and then, with that characteristic smile on his 
face and more than ample encouragement (I should have been suspicious), he 
quickly suggested the genus Cortinarius as a starting point. He said there were a 
lot of them around (a true statement) and that it would be helpful to him if I wrote 
up some of the collections. I agreed to get started at the first opportunity. Twenty 
years and some hundreds of descriptions later, I’m still working on Cortinarius and 
the closely related genus Dermocybe! Actually, I enjoy it very much and I am ex- 
ceedingly thankful to Harry Thiers for keeping me busy all these years. [Joseph F. 
Ammirati] 

x**x* xX eK X 

Unlike many college freshmen who vacillate between majors from semester to 
semester, my decision had been made several years before I stood in line on that 
first registration day at San Francisco State College. My plan was to eventually 
lead me to a career in high school teaching, that is until I met Harry Thiers. 

My first, and probably most memorable course, from Harry was Cryptogamic 
Botany. I can remember being somewhat apprehensive about being in this class, 
probably for two reasons: first, I had heard from other students that Dr. Thiers 
was a demanding teacher, and secondly, my older brother Don was also in the 
class. For the first week he wore a perpetually worried look, certain I was going to 
say or do something to embarrass the family name! As that semester went on, my 
apprehension turned to confidence and it became clear that my plan was changing 
and that I was going to do many more botany courses. Harry has a way of making 
each student feel important and unique. He is an unmerciful tease, a generous 
teacher, and a kind, thoughtful human being. 

The field trips, especially the weekend trips, have to be the most memorable 
part of any Thiers course. In particular, I remember Jackson State Forest in 
Mendocino where we collected mushrooms at spots known as "Mushroom 
Corners," "Aleuria Glen," and "Amanita Avenue," stayed in an old wooden 
dormitory affectionately called "The Villa," and shopped for food in the village of 
Mendocino at a little market called Mendoza’s. Harry was at his best on these 
trips; leading forays, reviewing his collections, "supervising" the shopping, answer- 
ing questions, "arguing" with his graduate students, helping with identifications, 
harassing the cooks, teasing us for our mistakes, appreciating our individuality, and 
encouraging our successes. 


10 


Dr. Harry Thiers nurtured our interest in the living world, shared with us his 
wonder of learning, and taught us with energy, humor, and commitment, and be- 
cause of that our lives have been forever enhanced. [Marcia Wicklow-Howard] 

x**x** * * * 

I first attended San Francisco State University in 1965. Shortly thereafter I 
took my first course from Dr. Thiers on the recommendation of several other stu- 
dents. I was overwhelmed with Dr. Thiers’ enthusiasm and knowledge of the sub- 
ject material as well as his personal interest in all of his students. We all felt that 
we were special and that our education was very important to him. My feelings 
may have been somewhat influenced by his somewhat unethical approach of brib- 
ing his students with cookies and other treats during class meetings. 

One area of concern to all students is grades. Dr. Thiers shared that concern 
with his students. His exams were fair and straightforward, no trick questions. 
This approach allowed us to be more relaxed and able to present more of our 
knowledge of the subject material. His evaluations took into account individual 
weaknesses. This special caring helped many of us develop the academic founda- 
tion to continue our education. I am sure that without his overwhelming support 
and guidance some of us would not have gone on to accomplish as much as we did. 
For this I am truly indebted to Dr. Thiers. 

The most enjoyable years of my life were spent at San Francisco State Univer- 
sity as Dr. Thiers’ student. The herbarium life was wonderful - open 24 hours a 
day. It seemed like Dr. Thiers was always there. I am sure this could be sub- 
stantiated by Ellen Thiers. The academic environment and the camaraderie in the 
herbarium that existed among a wonderful group of graduate students will always 
be with us. Thank you Dr. Thiers for a true quality education. [Bob Keller] 

* * Xe KK 

I was 21 and in my junior year at San Francisco State when I met Dr. Thiers. 
He taught a general botany course which I had taken to satisfy a college require- 
ment. Dr. Thiers taught the course many times, but when he lectured, he made me 
feel as though it were the first time he was sharing the information with a class. 
His approach was enthusiastic. When he talked of various organisms, "curious" 
was a term he often used to describe them. Curious is exactly what I became. 

His approach in lecturing was clear and scholarly, and his approach towards 
the students was friendly and encouraging. He encouraged questions and ans- 
wered even the most elementary ones without belittling the student. He knew 
when a student was struggling and made himself available for help. Except when 
teaching classes, Dr. Thiers was in his office actively working on research and 
keeping abreast of the changes in his field. His door was open, and he was never 
too busy to talk with his students. He would counsel, encourage, and listen. 

At the end of the semester of general botany, I changed my major from Ger- 
man to biology. I took several other courses from Dr. Thiers, each time finding 
them challenging and enjoyable. I have gone on to pursue the study of plants 
professionally. Without the enthusiasms and encouragement of Dr. Thiers, I might 
not have entered the field in which I have been so successful. [Barbara Keller] 

x ** * Kk * 

My first introduction to Dr. Harry Thiers was in the fall of 1972. At the time, I 
was not a student at SFSU, but my husband, Jim, was taking General Botany from 
Dr. Post. That semester, a botany retreat was held at Mendocino Woodlands and I 
tagged along. Since it was my first exposure to mushrooms, it seemed as though 
every time I turned around, there were 10 more new mushrooms. I followed Dr. 
Thiers all weekend trying to absorb all the names and characteristics. I was very 
impressed with all the information there was to learn about each mushroom, bes- 


11 


ides whether or not it was edible! The beauty of these organisms was overwhelm- 
ing - it was better than an Easter egg hunt. Eventually, I participated in countless 
more cryptogamic forays as an undergraduate, and then as a graduate student at 
SFSU under Harry Thiers. Now, long after finishing school, I still spend much of 
my free time botanizing and greatly appreciate being able to tag along with Dr. 
Thiers and his students, and all the while still learn from this wonderful man. 
[Mona Bourell] 
**e* ee * * 

As a new undergraduate at San Francisco State, I well remember HDT’s 
enthusiasm for mycology, and for the teaching of it. Harry was generally patient 
and generous with us as we struggled with esoterica which were second nature to 
him. I remember that he could also be gruff, at least in response to "whippersnap- 
perisms," my term for the awkwardly inappropriate remarks I seemed to be able to 
make all too frequently and obliviously. 

When the chips were down though, Harry was unfailingly gracious and suppor- 
tive. There were the lesser tragedies, such as our trying to clean the laboratory 
glassware after semester’s end. If we had asked how to operate the dishwasher 
properly, we probably would not have created the hurricane that pulverized a great 
quantity of HDT’s dearly-bought labware. But then it would not have been a sur- 
prise. He took it all in stride. 

A larger tragedy than an electrical fire in a herbarium drier should not even be 
contemplated. I experienced such a fire, which, in addition to seriously damaging 
the Biology building, destroyed about one third of my research collections and 
field notes, as well as material on loan from the New York Botanical Garden. For 
90 hideous minutes, we thought that all the western Agaricus holotypes might have 
vaporized. After the dust settled, Harry scraped my ego off the basement floor 
and gave me a major pep talk that helped me decide to stay in the game. 

Small goofs were usually digested relatively easily. I guess we forgot to tell 
HDT we were fixing a Mexican-style breakfast one morning at the Branscombe 
reserve. I didn’t know anyone could mistake salsa picante for strawberry jam, but I 
do now. I believe he ate the entire English muffin anyway, after that first poignant, 
pungent realization. 

There were other moments of grace under stress (his or mine), too numerous 
to mention here. If you know HDT at all you can imagine them as well as I could 
relate them. Well, I do remember one gruff response I provoked after complain- 
ing one time too many, that the Agaricus keys with which he had furnished us were 
miserable and useless testaments to human folly and ineptitude. "Well," he said, or 
rather glowered, “if you think that you can do better, then by all means be my 
guest." Or words to that effect. I like to think I spied a crafty glint behind that 
stern visage. I don’t so much mind being conned, but there are times when I 
wished I’d complained about a different genus. [Rick Kerrigan] 

* ** * * * 

As with many other students of Harry Thiers, I’m one of those who will never 
forget my first field trip with him. After my first five days in graduate school at 
SFSU, I was invited, along with two other students, to show up on Saturday morn- 
ing in front of the Biology Building at 6:00 AM ready for a collecting trip to the 
Sierra Nevada. Much to my chagrin, my alarm clock failed me, and after a five 
minute sprint (with basket on arm) from the dorm to the designated rendezvous, I 
found HDT pacing Holloway Avenue like an expectant father. After minor harass- 
ment, we were underway in HDT’s VW camper. Taking my duty as "shotgun rider" 
nonchalantly, I quickly learned not to offer hot coffee and sweet rolls to the driver 
(Harry) until after the Livermore interchange. Unfortunately, by the time we 


1s 


reached Livermore, the coffee was nearly gone and the sweet rolls were but a fond 
memory to us students. At this point, HDT initiated a pointed discussion about 
graduate student eating habits, whereupon our destination (Silver Lake Camp- 
ground) seemed several lightyears distant. Having chosen as a thesis project to 
work on Sierra Nevada boletes (a wise move I thought), I believed I now could 
redeem my tardiness and gluttony by showing my vast knowledge of Sierran 
boletes. After five minutes collecting, I soon realized that Boletus rubripes was not 
B. edulis, Gastroboletus turbinatus was not diseased, and that I should never again 
let myself be tricked into tasting B. calopus. When HDT made the suggestion to 
return to the car for lunch, I felt some relief. Finally, I thought, something I know 
how to do. On the way from the car to the picnic table, however, I dropped the jar 
of mustard (Grey Poupon no less) on the asphalt parking lot, prompting comments 
about the usefulness and qualities of today’s graduate students, and instigating in- 
stant flashbacks of all my previous blunders that day. Extraordinary caution the 
rest of the trip prevented further damage to my bruised ego. Nevertheless, I felt 
challenged. 

During the next two years, there were many more trips, especially to Silver 
Lake. These were times that I remember fondly and believe I was particularly 
lucky to have a chance to benefit from HDT’s expertise in the field. His jesting 
that demanded rebuttal, frankness, enthusiasm, and overwhelming interest in a stu- 
dent’s welfare inspired confidence to meet the challenges. Fortunately, I learned, 
among other things more academic, the appropriate times for drinking coffee and 
eating pastry in the car. [Roy E. Halling] 


* * * KK * 


Supervising the collection of hypogeous fungi near Yuba Pass. 


13 


I recently returned to visit SFSU 10 years after receiving my Master’s degree 
under Dr. Harry Thiers. As I drove to the campus, it appeared that nothing had 
changed in those intervening years. I parked my car in the same place I had always 
parked it and felt distinctly odd not carrying any books as I walked to the Botany 
building. To reach the graduate student desks in the herbarium, one must walk 
through a complex maze of herbarium cases which I found I could still negotiate 
on remote control. I have such a strong association with the aroma of naphthalene 
and dried mushrooms from my student days that it seemed as though each case I 
passed removed one of the last 10 years. 

The arrangement of the herbarium was exactly as when I'd been there, and my 
old desk was in its same place - I almost walked over and sat at it. Not only that, 
but nicknacks from a party 12 years ago were still decorating the room. Dr. Thiers 
has always been famous for the annual Christmas and end-of-the-year parties. 
One holiday season, we were required to make a gift to exchange with another 
graduate student. The only requirement was that it be made out of cryptogams. 
With the imagination typical of graduate students, we all went to work on these 
projects. The results ranged from freeze-dried mushroom arrangements to 
bracelets made out of Lactarius subdulcis to a mobile made out of lichens and 
mosses. The mobile is still tacked to the ceiling with a thumb tack! 

Dr. Thiers was also the same as ever; warm, friendly, welcoming and always 
ready with a joke or a mildly sarcastic quip. The first time Dr. Thiers teases a new 
student is like a rite of passage or the beginning of acceptance in the ranks of grad 
students. He used to tease George Wong for sleeping at his desk, Janelle Curlin 
for her diminutive stature and I got it once when we were going on a trip in north- 
ern California. Another student and I were driving together and missed the 
turnoff. We didn’t notice that we’d gone too far until we saw the "Welcome to 
Oregon" sign. This gave HDT sufficient fuel for teasing to last several days. For 
some reason, Roy Halling was labelled the "penultimate authority" during his time 
at State. Dr. Thiers was always quick to emphasize the penultimate nature of it. 
When a beginning student asked HDT for identification of some common 
mushrooms, he’d often say to take those to the penultimate authority and he might 
know what they were. 

Another major source of memories from SFSU were the numerous field trips 
and forays we took. We covered nearly every part of California, and beyond, in 
search of fungi, mosses, lichens and higher plants. As long as ’ve known Dr. 
Thiers, his expectations of students always seemed equitable, regardless of gender, 
when it came to setting up tents, cooking, building fires or hiking over any kind of 
terrain in search of the elusive mushroom. I look back very fondly on the years I 
spent studying under the exceptional guidance of Dr. Thiers. [Betty K. Tamm] 

* * * * * * 

I wonder if any one of us knew, as we luxuriated amidst the incredibly rich labs 
of Dr. Thiers’ classes, just how many levels of learning there were. I had no idea 
until I began to teach labs myself at the University. There I poured over his class 
notes, read and reread his marvelous lab handouts and slowly began to understand 
what an extraordinary education had been mine. Everything was organized around 
a deceptively simple formula: compare, contrast, synthesize. This was a formula 
for learning and it was also a formula for teaching. Now I understand why so many 
of us later taught. That is what HDT meant for us to do. He was never selfish 
with his vast knowledge and he did not mean for us to be selfish. 

The openness and generosity that HDT showed his students extended to other 
teachers and researchers as well. I look on my hours in the herbarium as the most 
intellectually stimulating time of my life to date. Dr. Thiers’ bottomless coffee urn 


14 


drew students and professors studying bees, butterflies, ecology, algae, plant 
chemistry, bryophytes and more. The list is a long one. It was in the herbarium 
that I learned about open sharing of ideas, frank admission of research problems, 
and to see the relatedness of all the natural sciences. Sharing, not secrecy, was the 
power of the herbarium. "When my students win, I win," he often said. HDT 
brought the great names in the field to our herbarium gatherings to share their 
knowledge, discuss our work and often contribute to our bulging herbarium. 

Now I understand how important it is for a teacher to respect students, for stu- 
dents to be taught to respect each other, and the power of humility in the face of 
greatness. HDT’s philosophy of life has stayed with me as surely as his beautiful 
labs in pathology, lichenology, lower vascular plants, mushrooms and soil fungi. It 
is a rich inheritance. Thank you, Sir. [Janelle M. Curlin] 

xx *e KX KK 

One morning in 1978 as a middle-aged reentry student working towards a 
masters in biology, I found myself in a large general botany lecture hall. I was 
much impressed with the energy and enthusiasm of the professor, Dr. Harry 
Thiers, and thought to myself, "Wow! This guy must have been teaching this sub- 
ject for 20 or 30 years and yet he still makes non-vascular plants sound like the best 
things that ever happened. And at eight o’clock in the morning!" 

A year later, I had a desk in the herbarium and was one of Harry Thiers’ grad- 
uate students, specializing in lichenology. It was a privilege to enter into this asso- 
ciation. The atmosphere in the herbarium was unique. There was much hard 
work, great field trips, fine parties, and Harry Thiers always present to help with 
our questions and problems. He was incredibly generous with his time, talents, 
and equipment. At the same time, he encouraged and prodded each of his stu- 
dents to accomplish as much as possible, largely by example. Thank you, Harry, 
for the "Herbarium Experience." [Janet Hoare Doell] 

* * * * KK 

A course taken from Dr. Thiers is a course never to be forgotten. Each one 
has at least one field trip; usually there are four or more. Such a wealth of in- 
formation, and presented in such an interesting and organized manner, his teach- 
ing abilities are beyond comparison. Not only in the classroom, but in the field, 
this man is full of unique knowledge as well as wonder and a yearning to discover 
more. Anyone who has had the pleasure to spend some time with Dr. Thiers, 
knows this is easily bestowed upon them. Whether it be in Marin, Sonoma, 
Mendocino, Humboldt, Del Norte, Siskiyou, Shasta, Sierra, Santa Clara, Santa 
Cruz, San Mateo, or San Francisco Counties in California, or perhaps another 
state, the experience is always greatly enhanced by his generosity and enthusiasm. 

He has such wonderful stories from years past taking place in Michigan, 
Montana, Idaho, Arizona, Texas, and the southern U.S. I wish I had taped these 
during his lectures. Though I may never get to one, I will always know how to walk 
in a Sphagnum bog after his brilliant demonstration! 

As can be seen from this volume, Harry Thiers’ contribution to mycology is 
enormous, invaluable, and something to be so proud of. And finally his contribu- 
tion to education is noteworthy, whereby many of his former students have gone on 
to be fine teachers. [Michelle Seidl] 

*x* x * * Kk X 

I first met Dr. Harry D. Thiers in the fall of 1973 while he was engaged in the 
identification of mushrooms for the Mycological Society of San Francisco’s fungus 
fair. Early the following year, I signed up for an evening class Harry offered to 
members of the mycological society who were interested in learning the technical 
ins and outs of mushroom identification. I soon discovered, via this informal class, 


15 


Harry’s gifted teaching talents, and his tremendously vast knowledge of fungi. Al- 
though our class only met for a few hours once a month, my guided discovery and 
fascination with the microscopic features of fungi quickly led to the purchase of a 
microscope and a number of technical monographs. 

It was not until the spring of 1982 that I took the first formal course from 
Harry through the San Francisco State University extension system. This course, 
"Higher Fungi of the Sierra Nevada," first offered in June 1982, was held at San 
Francisco State’s Sierra Nevada Field Station, near Yuba Pass, California. The 
course was attended by about 20 students, including Dr. Walter Sundberg, Dr. 
Andrew Methven, Dennis Desjardin, and Janelle Curlin, all of whom were either 
past or current students of Dr. Thiers. By the end of the week, our class had col- 
lected and identified some 220 species associated with the melting snowbanks and 
spring runoff in the Yuba Pass region. It was through this class that I got my first 
real taste of mycological taxonomy and encountered the tremendous pleasure of 
discovering the fascinating realm of the snowbank fungi. Harry continued to teach 
this class up through June 1987, finishing with a class size of just over 40 students. 

In January 1985, through a fortuitous change in jobs, I was able to re-enter col- 
lege, and begin a course of study under Harry’s guidance. Admitted as a junior, I 
was able to attend full time for only the first semester. That summer, after Harry’s 
class at Yuba Pass, I found another job which would allow me to continue as a part 
time student, while working full time to support my family. Aware that Harry was 
to retire in January 1989, I knew that I would need to take all of his courses quickly 
if I was to fit them in my schedule prior to his retirement. I feel fortunate indeed 
that I have been able to complete all of the classes that he offered during this time, 
and am particularly grateful to Harry for rescheduling some of these classes to 
meet during hours which I could attend. 

Harry takes a great deal of pride in each of his students, following their varied 
careers with interest and enthusiasm. His prior students often visit, write or call 
him to chat about current projects, career moves, etc. just to keep in touch. His 
correspondence desk has a transparent cover under which he keeps a large number 
of photographs of his students and close associates. He even hosts an annual 
reunion dinner for his prior students each summer at the MSA meetings. I see him 
react not unlike a father to many of his students, giving advice, guidance and direc- 
tion when called for, providing a strong foundation upon which to build a future 
career. He often displays prominently in the herbarium letters from his former 
students for the current crop of botany students and staff to read, so that they too 
feel as part of his family. I am proud to have Harry as a very respected friend, and 
feel fortunate that he has played a major role in my life and education. [Herb 
Saylor] 

xx x * * * 

Sharing the laboratory with Harry D. Thiers is an integral part of the experi- 
ence of an education in taxonomy. Dr. Thiers is a sight to behold, especially when 
he is working on taxonomic projects. I have had the privilege of observing him 
during his most recent endeavor with California Russulas. Nothing pleases Harry 
more than a drier full of specimens, spore prints already made, and preliminary 
notes written. The initial result of a successful collecting trip is an herbarium full 
of discarded wax bags, and assorted debris scattered on counter tops and the floor. 
The atmosphere in the herbarium at this point reflects Harry’s expansive mood. 
Later, when his microscopy starts in earnest, he becomes a different man. Quiet 
studiousness reigns during these weeks, while he buries himself in the wide, un- 
forgiving bosom of his oculars. "Got six done this afternoon!" indicates a moder- 
ately good session. "Why don’t you give up those Cladonias of yours?" is a playful 


16 


statement of his own frustration, and a challenge to spar. "I’ve got to go down to 
the bank," or more ominously, "Take a look at how neat everything is down at my 
end [of the herbarium]!" portends a struggle with intransigent pleurocystidia or 
spore ornamentation. Following a major cleaning session, during which the 
shelves holding Mycologia or The Bryologist are meticulously put in order and lined 
up, correspondence conquered, and reprints filed, the herbarium is slowly emptied 
of talkative students. Not until Dr. Thiers has safely returned to his microscope do 
the students filter back in. 

Despite his immense objectivity and scientific discipline, Harry has been 
known to reflect on a few of his major successes. One of these is that he convinced 
Alex Smith of the phylogenetic significance of hypogeous fungi in the Sierra Ne- 
vada. Another source of pride is his mycologist’s sense of smell and taste; the 
"green corn" [pronounced ‘carn’ until recently -Eds.] odor is known to all his stu- 
dents. I can boast of a minor accomplishment in this realm, convincing Harry that 
he can smell atranorin in certain lichen species. 

Some ways born of a harsh west-Texas boyhood die hard. Five o’clock marks 
the suspension of all work, be it in the field or laboratory, for the requisite happy 
hour. An hour or two of relaxation is inevitably balanced by a return to work. 
Nights find Dr. Thiers burning the candle until at least nine or ten PM. Even on 
damp field trips to Patrick’s Point State Park or Jackson State Forest, he finds an 
empty picnic bench after dinner to set up his specimens for spore prints. 

His late afternoon sabbatical works in tandem with another habitude, the ex- 
tension of warm and heartfelt hospitality to all who enter the herbarium. While I 
worked at SFSU with Harry Thiers, people from all parts of the world were wel- 
comed and assisted in their scientific pursuits. This marks the greatest quality of 
Harry as a taxonomist and teacher: his desire and ability to share with others. His 
sense of wonder, curiosity, and generosity continue to benefit all who cross his 
path. [Samuel Hammer] 


* * KX Ke KK 


Discussing collecting strategy at Yuba Pass. 


i? 


REMEMBERING THE MOREL GROWER: 
RON OWER, 1939-1986 


Ronald Dean Ower was born on 15 May 1939, and died on 23 March 1986, vic- 
tim of a murder-robbery. Ron was a long-time member of the San Francisco 
Mycological Society, who, after selling his sign-painting business, became a gradu- 
ate student of Harry Thiers at San Francisco State University in the fall of 1977. 
He wanted to grow morels (Morchella esculenta L.) in artificial culture, and this 
was the topic of his master’s degree research. He was not successful in inducing 
his cultures to fruit during the course of thesis work, which he concluded in the 
spring of 1980 with a thesis entitled "Cultural studies of morels". However, after 
completion of his degree, Ron was permitted to continue his culture work at the 
University. On 14 December 1980, he first detected the development of ascocarps 
that later developed to maturity. These were subsequently harvested on 11 Janu- 
ary 1981, and deposited in the San Francisco State University Herbarium (SFSU). 
After duplicating his results, he published a brief article on the achievement: Notes 
on the Development of the Morel Ascocarp: Morchella esculenta, Mycologia 74: 
142-143. 1982. 

Ron then began work on scaling up his technique, in order to develop com- 
mercially viable cultivation. He entered into an association with Neogen Corpora- 
tion, a biotechnology firm in Lansing founded by Michigan State University in 
1982. On 17 June 1986, a federal patent was granted to Neogen (Patent 
#4,594,804) for the "Cultivation of Morchella.". Ron Ower was credited with de- 
velopment of the technique. Tragically, he died three months before the patent 
was granted. The following are recollections about Ron and his work by three stu- 
dents of Harry Thiers who were contemporaries. [Barbara M. Thiers]. 

xx * kK *X * 

I remember Ron’s first spring with Dr. Thiers as a time when Ron frequently 
would be out in the field, camping in some likely but frigid spot, waiting for the 
first Morchella primordia to appear. It was apparent that Ron’s approach to un- 
derstanding reproductive initiation in Morchella was to be thoroughly familiar with 
the natural history of the phenomenon. To this end he assiduously applied him- 
self, and we saw rather little of him for several weeks. I could tell before arriving 
at the herbarium whether he had returned from such an expedition; he and I 
parked in the same peripheral sector of the SFSU environs, and one could not miss 
the large signs for the "Morchella Co." which adorned the doors of his truck. He 
would show us what, if anything, he had found, and make taxonomic predictions 
about it, before setting to work on culturing the new material. 

As his work progressed he shifted his focus to laboratory work. He explored 
the genetics of sexuality in Morchella with the help of a micromanipulator, spend- 
ing long hours on "ascus stroking,” as he called it. He explored anamorph- 
teleomorph connections in these organisms, but did not submit his findings when a 
similar report appeared in the literature. Most of his later work at SFSU focused 
on cultural techniques for inducing reproductive growth in Morchella under con- 
trolled conditions, and, as is well known, he was the first to succeed in achieving 
this. Subsequently he explored the morphology of the developing ascoma of the 
morel. 

Ron Ower was somewhat of an enigma to me. He could at times be sur- 
prisingly candid, while at others he was very reserved. From the outset of his re- 


18 


search career at SFSU he realized the commercial potential of his research objec- 
tives; as a result he became less communicative over time, to the point where I 
couldn’t say I knew his personal or professional side very well. By that time I was 
employed in the commercial mushroom industry, at one point with a firm who was 
negotiating with Ron for rights to his morel-growing process, which explains why 
Ron may have experienced a conflict as regards the nature of our communications. 

I remember Ron as a relaxed and genial individual, never the center of atten- 
tion, almost playful in his approach to social behavior and to life in general. When 
one was aware of his presence he was usually make some positive contribution to a 
conversation in the herbarium or a meal on a field trip. He accomplished his ini- 
tial scientific objective of fruiting the morel, and after years of effort he also met 
with reasonable financial success in exploiting the commercial aspects of his pro- 
cess. I last spoke with him about a year before his death when he referred to the 
continuing experiments on Morchella culture that he was conducting in his small 
private laboratory in San Francisco. He was intrigued and optimistic. [Richard W. 
Kerrigan] 

xx *¥ xk K * 

I was obsessive about cleanliness. Ron thought that was a waste of time. We 
shared a growth chamber. The combination could have been a disaster. Instead, it 
was fun and occasionally even exciting. Like the day Ron announced that wild 
morels did not fruit in sterilized soil, and that was probably why his were not fruit- 
ing. After that, my marine fungi shared their growth chamber with trays and trays 
of horse sweeping from Golden Gate Park, unautoclaved. 

My marine fungi were never contaminated. Ron’s morels fruited right on 
schedule for HDT’s farewell party as he left for Australia on sabbatical. 

Ron and I confided our fears of aging, poor memory, loneliness, and money 
troubles. He gave my various boyfriends nicknames. One I particularly remember 
was dubbed "Old Abe Lincoln." Our favorite conversation topic was HDT (of 
course). Ron said that two hours of conversation with HDT could provide him 
with weeks of good ideas. We tried to figure him out. Why was he so intellectually 
stimulating? Why did we all work so hard for him? How could we ever be like 
him? Did he really care as much as he seemed to? How could he get so much 
done on his own research when he spent so much time teaching? What was the 
secret of his encyclopedic memory? It was a topic we never tired of discussing. 

My slide collection contains many slides of Ron and his collecting finds. He 
had a sixth sense in the field and not only for morels. I treasure those slides of a 
man who really enjoyed fungi and whose big smile conveyed his delight. [Janelle 
Curlin] 

* * * KK 

Ron Ower sat facing me in the lichen lab and when I think of him I picture his 
very round eyes and open countenance. He was not enchanted with lichenology 
and especially disliked memorizing all the names, something he could not do well. 

He was a conscientious student and tried valiantly to study each collection in 
detail. He was conscientious in other ways as well and it bothered him that his 
parents had to help support him while he was devoting so much time to his morel 
growing project. That was an interesting semester. We cheered at every 1/8 mm 
of growth, grieved over the ones that did not survive, and considered it an honor to 
be invited into the inner sanctum to take a peek. 

However, he agonized in the succeeding weeks and months as he tried to turn 
his discovery into a livelihood. He was depressed more than once, but always 


19 


grabbed eagerly at hopeful developments and was happy to talk about them when- 
ever we ran into each other after graduation. He was adamant about not leaving 
San Francisco and that added to the difficulty of working out something in 
Michigan. 

Ron was something of a loner. He enjoyed an occasional meal at our home, 
including holiday dinners, and rarely seemed to have other commitments. He was 
a gentle person. It is hard to imagine him suffering such a violent death. He 
deserved something much better. [Janet Hoare Doell] 


Ronald Dean Ower: 1939-1986. 


i ) iy 
a 
et wa : 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 21-36 January 20, 1989 


DERMOCYBE, SUBGENUS DERMOCYBE, SECTION SANGUINEAE 
IN NORTHERN CALIFORNIA 


JOSEPH F. AMMIRATI 


Department of Botany, KB-15, University of Washington 
Seattle, Washington, 98195 


ABSTRACT 


Five taxa in the genus Dermocybe, subgenus Dermocybe, 
section Sanguineae are fully described. These include D. 
sanguinea and D. semisanguinea; a new species, D. sier- 
raensis; and two new combinations in Dermocybe, D. cali- 
fornica and D. phoenicea var. occidentalis. A key to these 
taxa 1S provided. 


INTRODUCTION 


In all earlier publications by the author Dermocybe was 
considered to be either a subgenus or section of Cortinar- 
jus. Here, Dermocybe is recognized as a genus with Timits 
as defined in Singer (1986). In this paper only the sec- 
tion Sanguineae is treated. Furthermore, only those 
Species recorded to date by the author from northern 
California are described below. A more extensive, but 
Still incomplete treatment of section Sanguineae for North 
America, can be found in Ammirati and Smith (1984). 


In this study all microscopical data was taken from sec- 
tions or pieces of dried basidiomata which were mounted in 
a 3% aqueous solution of potassium hydroxide. Capitil ized 
color notations are from Ridgway (1912). Uncapitilized 
color terms are regarded as useful approximations. Terms 
Such as moderate reddish orange are from Kelly and Judd 
(1965). All cited collections are deposited in the San 
Francisco State University Herbarium (SFSU) unless other- 
wise noted by use of an abbreviation from Index Herbar- 
jorum. 


22 


BRIEF SYNOPSIS OF SECTION SANGUINEAE 


Basidiomata small to medium sized; pileus surface silky to 
innately fibrillose, appressed fibril lose, or minutely 
Squamulose, dry to moist, usually not hygrophanous or 
hygrophanous to subhygrophanous, color deep to rich red, 
purple-red, reddish brown to brownish orange, or less 
commonly yellow-brown to somewhat ol ive-brown; context 
thin to moderately thick; lamellae when young dark to rich 
red, purplish red, rich brownish red, reddish orange or 
brownish orange, rarely truely orange or yellow; stipe 
usually slender, equal or the base somewhat enlarged, 
Silky to fibril lose, variously colored, often, but not 
always, concolorous with the pileus surface and lamel lae; 
interhyphal pigment deposits often lacking, when present 
usually occurring as small deposits or particles; basidio- 
mata rich in anthraquinonic pigments (see Keller and 
Ammirati, 1983); potassium hydroxide (3% aqueous solution) 
applied to pileus surface of fresh basidiomata dark red, 
purple-red, purple or blackish purple; basidiospores orna- 
mented; cheilocystidia either present, and not well dif- 
ferentiated, or absent. 


Type species: Dermocybe sanguinea (Wulf. : Fr.) WUnsche 


KEY TO SPECIES AND VARIETIES 


1. Pileus surface color usually yellow-brown to dark 
yellow-brown, at times more yellowish, orange-brown, 
or oOlive-brown; KOH (3% aqueous solution) instantly 
changing the pileus surface to purple or blackish 
purple when applied to fresh-specimens; lamellae rich 
red to purple-red; stipe typically some shade of 
yellow, pinkish or reddish tones may be present on 
the base, and similarly colored fibrils may sparsely 


COdte(ThERSUGL OC Cae rane re ae. D. semisanguinea 


1. Basidiomata not as described above; pileus lamellae, 
and often (but not always) the stipe, purple, red, 
red-brown, orange-red or more orange in color...2 


oe Pileus surface color deep red to rich red or deep 
red-brown; lamellae deep purple-red to deep red or 
bright red; stipe ground color yellow to ochraceous 
to dingy buff, the stipe base similarly colored or 


20 


pinkish to reddish, and reddish to pale ferruginous 
red or brownish fibrils may or may not coat the stipe 
SGA Cure W cirieih sta 'e Wes, g0 tate Toul ee Alto Ue te iS acu h ats het ss 3 


Pileus, lamellae and stipe basically more or less 
concolorous when fresh, usually some shade of purple- 
red, red, or orange-red to brownish orange (yel lowish 
to ochraceous colors may be present on the stipe 
base) Sees Wrst aM cht elt oir entrents Vol sikrelt snl oe oy ohh otas tet he 4 


Stipe ground color some shade of yellow, base or 
Surface fibrils may be colored reddish or pinkish 
a at D. phoenicea var. occidentalis 


Stipe ground color more dingy buff, surface sparsely 
to more heavily coated with reddish fibrils ..... 
- see discussion under D. phoenicea var. occidental is 


Pileus surface appressed silky to silky fibril lose, 
hygrophanous or not hygrophanous, color reddish 
orange, reddish brown, brownish red-orange or 
DO Wilds NeOJrdT) Cur, eae Mstte ye he oesthatis felts se aS RCE WA be) 


Pileus surface appressed fibrillose to minutely 
Squamulose, not hygrophanous, color deep red to rich 
momeey VV 10ened: OF epUrp lie =ned Wr Seen are sel etet s 6 


Pileus surface appressed silky, more or less hygro- 
phanous, color reddish brown to reddish orange, fad- 
ing to brownish orange; stipe surface some shade of 
orange to reddish orange with a more or less concol- 
orous base (often orange-red); basidiospores verru- 
eo0se, 8=-9.5'x"5-6°um:-in’Ss1ze. 2. 3. D. californica 


Pileus surface silky to fibril lose, not hygrophonous, 
color brownish orange to brownish red-orange; stipe 
deep vinaceous to almost concolorous with pileus 
Surface, base vinaceous not (yellowish or ochra- 
ceous); basidiospores verruculose, 7.5-9 (-9.5) x 5- 
SeOM LO SUM MI TacSttZGa. etleuee sie yeuretys D. sierraensis 


Pileus, stipe (except for base), and lamellae rich to 
feetaned: Ome Di LOOdmyCd Wier waist eons D. sanguinea 


Pileus, stipe (except for base), and lamellae more 
vivid red to purple-red 
: . see discussion under D. sanguinea 


24 


DESCRIPTIONS OF TAXA 


Dermocybe californica (Smith) comb nov. 


BASIONYM: Cortinarius californicus Smith, Contrib. Univ. 


Mich. Herb. 3:37-38. 1939. 


PILEUS 16-85 mm broad, obtusely conic to companulate 
becoming broadly companulate, umbonate to broadly 
umbonate; margin typical ly decurved, appressed against the 
Stipe at first; surface smooth, glabrous to appressed 
silky, more or less shiny, moist to dry, more or less 
hygrophanous, color moderate reddish brown (Bay, Hay's 
Russet) to dark reddish orange (Vinaceous Rufous) or 
strong brown (Kaiser Brown) at first, fading to brownish 
orange (Cinnamon Rufous), with the disc often remaining 
darker. Context moderately thick on the disc, watery at 
first, moist to dry, concolorous with surface of moist or 
faded pilei, often pale reddish ferruginous; odor and 
taste not distinctive or slightly fungoid-raphanoid. 


LAMELLAE adnate becoming slightly adnexed, narrow to 
moderately broad (5-7 mm wide), often more or less ventri- 
cose, subdistant, color dark reddish orange to moderate 
reddish orange (English Red, Mars Orange) or deep orange 
(Orange Rufous) at first, becoming deep orange (Burnt 
Sienna) or ferruginous brown; edges long retaining the 
original color, fimbriate or slightly serrulate. 


STIPE 50-150 (-200) mm long, 5-15 (-22) mm thick, the 
base often 10-20 mm thick, terete, equal or more frequent- 
ly the base slightly enlarged to clavate, occasional ly 
narrowed below; surface dry appressed silky or sparsely 
fibrillose, subshiny, color dull orange, dark reddish 
orange to moderate reddish orange (Ferruginous) or 
moderate orange (Flesh-Ochre), sometimes with orange 
fibrils from remains of cortina, basal mycelium often 
orange-red. Context solid becoming hollow, color pale 
moderate orange (pale Flesh-Ochre) to rusty red near the 
surface, unchanging. CORTINA profuse, fairly persistent, 
color pale moderate orange (pale Flesh-Ochre). 


BASIDIOSPORES (7.4-) 8-9.5(-11) x (4.8-)5-5.85(=720) 
um, in profile view elliptical to broadly elliptical, at 
times somewhat fusiform or more or less amygdaliform, in 
face view elliptical to broadly elliptical or fusiform, 


25 


more or less verrucose, ornamentation coarser toward 
distal end, yellow-brown with darker brown ornamentation. 
BASIDIA 4-spored, 24.1-32.9 x 5.8-8.0 um, elongate-clavate 
to clavate or broadly clavate, hyaline to faintly purple 
or pinkish, often containing deep purple to red-purple 
granules, some containing brownish to yellow-brown pig- 
ment. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 21.9-38 x 6.6- 
13.1 um, more or less cylindrical to cylindrical-clavate, 
clavate, broadly clavate or somewhat ventricose, thin- 
walled, color similar to basidia, usually mixed with 
basidia but sometimes in groups. TRAMAL HYPHAE OF LAMELLAE 
Subparallel, somewhat interwoven, 3.3-29.2 um wide, cylin- 
drical to more or less inflated, hyaline to slightly 
pinkish or purplish, some containing purple, red-purple or 
pinkish purple granules. CUTICULAR HYPHAE OF PILEUS inter- 
woven, more or less radially arranged, cylindrical to 
inflated, 3./-26.5 um wide, in surface layer mostly 3.7- 
8.8 um, the broader, inflated hyphae more common in the 
Subcuticular region (this often appears as a distinct 
hypodermium between the surface layer and trama), thin- 
walled, some encrusted, hyaline, yel lowish, light brownish 
or yellowish brown in the surface layer, in the subcuticu- 
lar region similar to tramal hyphae of pileus; no pileo- 
cystidia seen. TRAMAL HYPHAE OF PILEUS interwoven, more or 
less radially arranged, especially in upper trama, cy] in- 
drical to inflated, 5.1-29.2 um wide, color similar to 
tramal hyphae of lamellae. CLAMP CONNECTIONS of the normal 
or medallion type, present throughout the basidiomata. 
INTERHYPHAL PIGMENT DEPOSITS absent (some small interhy- 
phal granules may be present but no large deposits). 


Solitary or gregarious to caespitose in mixed hard- 
wood-conifer and conifer forests; October into December. 


COLLECTIONS EXAMINED. California. Del Norte Co.: B.F. 
Isaacs 506 (WTU); J.F. Ammirati 140, 141; A.H Smith 8957 
(holotype, MICH); H.D. Thiers 14308, 14496. Humbo 1dt Coz: 
J.F. Ammirati 148. Mendocino Co.: J.Fe Ammirati rye Ole 
and 6186, (both MICH); J. Motta ID 328; H.D. Thiers 9628, 
18460, 21751, 30464, 30825, 3317/7, 35602, 40601; G. Wong 
Bi. Siskiyou COs aan Us Thiers 46798. Tuolumne Co.: Hel. 
Thiers 46934. Yuba Co.: H.D. Thiers 47143. 


Dermocybe californica is one of the most common and 
widespread species of section Sanguineae in Northern 
California. In general it occurs along the west coast from 
California into Canada and east as far as northern Idaho. 


26 


Its overall] coloration (reddish brown, reddish orange, 
brownish orange), more or less silky, usually hygrophanous 
pileus, and coarsely ornamented basidiospores, easily 
separate it from all other Sanguineae. It is very closely 
related to Dermocybe cinnabarina (Fr.) Wlunsche, differing 
primarily in its preference for coniferous forests. D. 
cinnabarina occurs in Europe in deciduous forests with 


Fagus, Quercus or Carpinus. 


Dermocybe phoenicea var. occidentalis (Smith) comb. 
nov. 


BASIONYM: Cortinarius phoeniceus var. occidentalis 
Smith, Contr. Univ. Mich. Herb. 2:30-31. 
1939. 


PILEUS 30-80 mm broad, broadly convex, becoming ex- 
panded, more or less umbonate; margin usual ly decurved; 
surface moist to dry, innately silky becoming appressed 
fibrillose at maturity, often cracking radially in age, 
color evenly dark red to rich red (Ox-Blood Red to Garnet 
Brown) or dark mahogany red. Context rather thin, firm, 
color buff, with reddish tints near the cuticle and olive- 
brown coloration near the stipe; odor and taste none or 
not distinctive. 


LAMELLAE adnate to adnexed, subdistant to distant or 
appearing close in small pilei, broad, equal, color deep 
purplish red (Bordeaux) to deep red, sometimes with 
brighter red (almost Carmine) edges, with a changeable or 
metallic sheen when viewed at different angles, becoming 
more or less rusty in age. | 


STIPE 40-110 mm long,. 6-15(-30) mm thick, equal or 
Sometimes clavate to ventricose, surface moist to dry, 
color evenly yellow (Mustard Yellow) to dull yellow 
(Chamois, Honey Yellow), or dull ochraceous, with a coat- 
ing of yellowish fibrils, mycelium around base and in soil 
ochraceous to ocher-yellow (Yellow Ocher) or tinged red- 
dish to vinaceous red. Context light yellowish olive (Old 
Gold), becoming sordid brownish in age. 


CORTINA scanty, dull yellow (Chamois), light 
ochraceous or yel low-buff. 


27 


BASIDIOSPORES (6-)6.5-8(-9) x 4-5(-5.5) um, in pro- 
file view elliptical, in face view elliptical, verrucu- 
lose, pale brownish with darker brown ornamentation. 
BASIDIA 4-spored, 24-31 x 6.5-7.5 um, clavate, thin-wall- 
ed, hyaline or reddish to pinkish, some containing hyaline 
or reddish granules. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 
apparently absent. TRAMAL HYPHAE OF LAMELLAE subparal lel 
to slightly interwoven, cylindrical to inflated, 5-18 (- 
23) um wide, hyaline to pinkish or light reddish vina- 
ceous. CUTICULAR HYPHAE OF PILEUS interwoven, more or less 
radially arranged, cylindrical to inflated, 5-18 um wide, 
thin-walled, faintly colored or light brownish, light 
reddish to light vinaceous, or containing a concentrated 
reddish purple to bluish purple pigment and often purplish 
granules; pileocystidia absent (apex of end cells rounded 
to tapered). TRAMAL HYPHAE OF PILEUS interwoven, more or 
less radially arranged, cylindrical to inflated, 3-31 um 
wide, hyaline, yellowish, yellowish brown, or pale vina- 
ceous. CLAMP CONNECTIONS of the normal type, present 
throughout the basidiomata. INTERHYPHAL PIGMENT DEPOSITS 
present in the pileus trama, the stipe cortex, and among 
the hyphae of the universal veil, in KOH yellowish, brown- 
ish orange, orange, or reddish orange. 


Caespitose to gregarious or scattered in conifer or 
mixed woods (Picea, Pinus, Pseudotsuga, Tsuga). October to 
mid-December or sometimes as early as August in the higher 
mountains. 


Collections examined. California. Del Norte Co.: A.H. 
Smith 8339, 8591, 8997, 55937 (all MICH). Humboldt Co.: 
J.F. Ammirati 8550 (WTU); H. Lamphere 34 (MICH); A.H. 
Smith 8679, 56057, 56228, 56282, 56517 (all MICH). Mendo- 
eynom0o0,:) J.-F. Ammirati 6213, 6215 (both ;MICH)3) H.D. 
Thiers 30399, 30429, 30862, 33278, 38386, 40483, 41649, 
#3935. Monterey Co.: H.D.) Thiers 32169, 39598. YubaiCo.: 
H.D. Thiers 44130, 47142. 


Dermocybe phoenicea var. occidentalis is the most 
frequently encountered member of section Sanguineae on the 
Pacific Coast, extending from California into Alaska. It 
also occurs in the mountains of the interior, where it can 
be found in some years as early as August. It prefers 
coniferous woods, but also occurs in mixed forests of 
conifers and deciduous trees. 


28 


As described by Smith (1989), D. phoenicea var. occi- 
dentalis has a dark red to rich red, silky to appressed 
fibril Tose pileus, deep purplish red to deep red lamellae, 
and a yellowish to ochraceous stipe with a coating of 
yellowish fibrils. The stipe base being either ochraceous 
or tinted reddish. This is the most common form of this 
variety. On the Pacific Coast there is a less frequent 
form which has a dingy buff (Cinnamon-Buff) stipe coated 
with pale ferruginous red fibrils (for examples, A. H. 
Smith 56282 and 56517). This is not recognized here for- 
mally for two reasons. Firstly, these collections may 
Simply represent one end of a color spectrum where the 
Stipe ground color has lost the yellow coloration and the 
surface fibrils are more reddish. A similar pattern of 
variation can be seen in a close relative D. semisanguinea 
in both North America and Europe. Secondly, D. phoenicea 
var. occidentalis needs to be more carefully compared we 
European material of D. phoenicea var. phoenicea (Bull. 
Maire) Moser, to determine if they are really parts of the 
Same species. Once this is accomplished variation in D. 
phoenicea var. occidentalis can be better evaluated and 
additional infraspecific taxa established it this is 
appropriate. D. phoenicea var. phoenicea, which has a more 
brownish to brownish red pileus, is not particularly 
common in North America and may occur ROOKIES in eastern 
North America. 


Dermocybe sanguinea (Wulf. : Fr.) Wlnsche, Die Pilze. 125. 
1877. 


BASIONYM: Agaricus sanguineus Fr, Systema Mycologicum. I. 
22001821". 


SYNONYMS: Cortinarius sanguineus (Fr.) S.F. Grayoeer 
Natural Arrangement of British Plants. I. 
OAS 


Cortinarius sanguineus (Fr.) "Fro Eptouasae 


Systematis Mycologici. 288.1838. 


PILEUS 18-45 mm broad, somewhat obtuse to convex or 
plane, disc more or less depressed; margin incurved to 
decurved; surface appressed fibril lose to minutely squamu- 
lose, sometimes radially rimose on the margin, color on 
disc rich red (Garnet Brown) to deep red (Maroon to deep 


29 


Garnet Brown) or tinted with colors of the margin, on 
margin rich red (Garnet Brown to Ox-Blood Red) or duller 
red (Coral Red), sometimes streaked deep red to fuscous- 
red (Maroon to Victoria Lake). Context watery deep red 
(Maroon to Garnet Brown), where faded or beneath cuticle 
lighter red (near Coral Red); odor somewhat fragrant, mild 
or raphanoid; taste mild to raphanoid. 


LAMELLAE sinuate to broadly depressed with a more or 
less decurrent tooth, close, more or less ventricose when 
mature, at first rich red (0x-Blood Red, dull Carmine or 
Garnet Brown), becoming shaded rusty brown to dusky brown 
from the basidiospores; edges uneven. 


STIPE 45-85 mm long, apex 3-9 mm thick, equal or the 
base slightly enlarged; surface fibrillose, shiny, rich 
red (Garnet Brown, O0x-Blood Red) or duller red (Coral 
Red), sometimes with watery deep red (Maroon) streaks, 
base or lower third dull ochraceous (Ochraceous-Buff) or 
the ochraceous coloration slightly tinted orange to red- 
dish. Context stuffed to narrowly hollowed, rich red 
(Garnet Brown to 0x-Blood Red) throughout or deeper red 
(Maroon) in the lower portion, where faded paler red. 


BASIDIOSPORES’ 6:5~9(-10)x) (3.5-)4.5-5(-5.5) um, in 
profile view elliptical, in face view elliptical to ovate, 
verruculose, color brownish to fulvous. BASIDIA 4-spored, 
15-25 x 6-7.5 um, broadly clavate, clavate, or more or 
less ventricose, hyaline to pinkish. PLEUROCYSTIDIA ab- 
sent. CHEILOCYSTIDIA 12-23 x 7-10 um, clavate to broadly 
clavate, thin-walled, color similar to basidia. TRAMAL 
HYPHAE OF LAMELLAE subparallel to slightly interwoven, 
cylindrical to more or less inflated, 5-25 um wide, 
hyaline, pale pinkish or light vinaceous. CUTICULAR HYPHAE 
OF PILEUS interwoven, more or less radially arranged, 
cylindrical to inflated, 8-30 um wide, thin-walled, pale 
pinkish to pale vinaceous or containing amore or less 
concentrated reddish to reddish vinaceous pigment; pileo- 
cystidia absent (hyphal end-cells with rounded to tapered 
apices). TRAMAL HYPAHE OF PILEUS interwoven, more or less 
radially arranged, cylindrical to inflated, 7-30 um wide, 
color as for the tramal hyphae of the lamellae or contain- 
ing vivid red pigment and granules. CLAMP CONNECTIONS of 
the normal type, present throughout the basidiomata. 
INTERHYPHAL PIGMENT DEPOSITS present as small yellow to 
orange particles in H50, in KOH none observed. 


Gregarious in mixed woods, mixed conifers, and 
Douglas fir stands. Mainly November through mid-January. 


COLLECTIONS EXAMINED. California, Humboldt Co.: J.F. 
Ammirati 6187 (MICH); H. Lamphere 56 (MICH); A.H. Smith 
3747, 9156 (both MICH); H.D. Thiers 14358, 14427. Mendoci- 
no Co.: H.D. Thiers 9944, 14684. 


Dermocybe sanguinea is widely distributed in North 
America and almost always occurs in coniferous woods. It 
is characterized by rich to deep red basidiomata that have 
ochraceous to orange-ochraceous mycelium over the stipe 
base. The pileus surface is typically appressed fibril lose 
with minute squamules or scales, especially on the margin. 
Along: the Pacific,Coast a mone vivid red (Roseskegiaro 
purplish red (Bordeaux) variant occasionally has been 
collected, for example, A.H. Smith 79922 (MICH), Til 1amook 
Co., Oregon. Since its coloration integrades with that of 
typical D. sanguinea, and it does not differ significantly 
from the Tatter in other features, it is not recognized 
here as a separate taxon. Species in section Sanguineae, 
which might be confused with D. sanguinea, are D. sier- 
raensis and D. california. Check descriptions of these 
taxa before making a final determination. 


Dermocybe semisanguinea (Fr.) Moser, Schweiz. Z. Pilk. 


SPAS ROLE 


BASIONYM: Agaricus cinnamomeus L. : Fr. var. semisanguin- 
eus Fr. 


Cortinarius semisanguineus (Fr.) Gillet, Les 


Hymenomycetes. 486. 1874. 


PILEUS 15-50(-70) mm broad, conic-campanulate to 
rounded-conic becoming convex to plane, umbonate to subum- 
bonate; margin slightly inrolled at first becoming incurv- 
ed to decurved; surface appressed fibrillose to fibril- 
lose-scaly, sometimes more or less rimose on the disc, 
moist to dry, not hygrophanous, color on disc yel low-brown 
(Ochraceous-Tawny) or orange-brown (Amber Brown) or dark 
yellow-brown (Sudan Brown), on margin typically yel low- 
Cinnamon to ochraceous brown or ochraceous-buff, sometimes 
when young tinted orange-buff (Apricot Buff), in age tint- 
ed or streaked with colors of disc, occasionally the 


31 


overall] coloration tinted with light olive-brown (01d 
Gold, Tawny Olive). Context solid, firm, dull whitish to 
light dull yellowish or slightly brownish, often becoming 
more or less sordid; odor and taste raphanoid or indis- 
tinctive. 


LAMELLAE adnate to adnexed or emarginate, seceding in 
age, close to subcrowded, up to 9 mm broad, more or less 
ventricose mature, dull red (Dragon's-Blood Red to Brick 
Red) to deep red (near Morocco Red) or purple red (Bor- 
deaux) becoming brownish to brownish orange in age; edges 
even to wavy. 


STIPE 25-80 mm long, 4.5-15 mm thick, equal to more 
or less clavate or ventricose; surface appressed fibril- 
lose, sometimes with a slight fibrillose annular zone near 
the apex, color pale yellow (pale Empire Yel low) to dul] 
yellow, more or less concolorous with the pileus margin, 
or sometimes rusty-brown from the basidiospores in age, 
the base colored as above or with light reddish (Peach 
Red) to dull pinkish tones, becoming olivaceous in age in 
Some specimens, some surface fibrils may be tinted brown- 
ish to reddish or pinkish. Context stuffed becoming hol- 
lowed, yellowish white to dull yellow or more or less 
concolorous with the surface, developing a watery oliva- 
ceous cast in age, in some the cortex of the stipe base 
has a reddish to reddish orange cast. CORTINA yellowish, 
evanescent. 


BASIDIOSPORES in deposit moderate yellow-brown (near 
Snuff Brown), (5.5-)6-8.4(-11.7) x 3.8-4.8(-5.8) um, in 
profile view elliptical, in face view elliptical to broad- 
ly elliptical, verruculose, light brownish with darker 
brown ornamentation. BASIDIA 4-spored, 20-31 x 5.5-7 um, 
clavate to more or less ventricose, hyaline or reddish 
purple to purple. PLEUROCYSTIDIA absent. CHEILOCYSTIDIA 
apparently absent. TRAMAL HYPHAE OF LAMELLAE subparallel 
to more or less interwoven, cylindrical to inflated, 5- 
15(-20) um wide, hyaline or more commonly light purple, 
vinaceous purple, or reddish purple, some containing pur- 
plish granules. CUTICULAR HYPHAE OF PILEUS interwoven, 
more or less radially arranged, cylindrical to more or 
less inflated, 5-16 um wide, thin-walled, some encrusted, 
brownish to faintly colored or frequently containing more 
or less concentrated purple, bluish purple, or light vina- 
ceous purple to bluish pigment and granules; pileocystidia 
absent (apex of end-cells rounded to tapered). TRAMAL 


32 


HYPHAE OF PILEUS interwoven, more or less radially arrang- 
ed, cylindrical to inflated, 6-25 um wide, some encrusted, 
hyaline to pale yellowish or dingy yellow or in upper 
portion some similar to cuticular hyphae. CLAMP CONNEC- 
TIONS of the normal type, present throughout the basidio- 
mata. INTERHYPHAL PIGMENT DEPOSITS present as small de- 
posits or particles in the pileus, lamellae, and stipe, 
and among the cortinal hyphae, yellowish to orange or 
reddish. 


Solitary, scattered, gregarious or caespitose in 
conifer and mixed woods, occasionally on rotten conifer 
wood. Rarely found in California, look for it in the 
autumn season, October thru December. 


Collections examined. California. Humboldt Co.: H. 
Lamphere 33 (MICH). 


Dermocybe semiSanguinea appears to be a rare species 
in California; only one collection seen by the author to 
date. It is found across North America, typically in 
coniferous forests, but also in mixed forests. The distri- 
butions of D. semisanguinea and D. phoenicea sensu lato 
overlap in North America. However, the frequency with 
which one encounters the two species in western and east- 
ern North America is striking different. D. seminanguinea 
is much more common in eastern North America than D. 
phoenicea. The latter rarely occurs there. D. semisan- 
guinea 71S much less frequently found in the west, particu- 
larTy along the Pacific Coast. Instead one finds D. 
phoenicea var. occidentalis as the common representative 
of this group. As one goes down the Pacific Coast into 
southern Oregon and Northern California D. semisanguinea 
is so infrequent that it is truely a rare find! 


Dermocybe semisanguinea is a highly variable species 
in North America and is still under investigation. The 
description here represents a fairly broad concept of the 
Species based on a number of collections. The closest 
relative of D. semisanguinea, D. phoenicea var. occiden- 
talis, differs from the former primarily in its pileus 
color. In the former the pileus is usually some shade of 
yel low~brown while in the latter it is dark to rich red. 


Dermocybe sierraensis sp. nov. 


SYNONYMS: Cortinarius sanguineus (Wulf. : Fr.) S.F. Gray 
var. sierraensis G. Keller & J.F. Ammirati nom. 
prov., Mycotaxon 18(2):364. 1983. 


Cortinartusssanguineus (NUIT. Fr.) S.Fse Gray 
var. Sierraensis J.F. Ammirati & A.H. Smith nom. 
prov., McIlvainea 6(2):62. 1984. 


PILEUS 20-40 mm latus, novus convexus, maturior 
plano-convexus, denique DTanus; margo incurvus, demum 
decurvatus vel planus; Superficies $1 Sicca, glabra vel 
adpresse eal ae s nitida, da, brun brunneo- 
aurantiaca. Contextus roseolus, -2 mm crassus, sapor 
odorque haud proprit. propria. ne Pa 


LAMELLAE adnatae vel adnexae, angustae, atrorubrae, 
margines integri, concolores. 


STIPES 20-40 mm longus, 2-4 mm crassus, aequalis, 
siccus; Superficies pannis veli exceptis glabra, obscure 
Vinacea ai pili superficie colorem abeuns, mycelio basim 
versus vinaceo; panni veli cum St Superfici Lev concosioness: 

contextus solidus, cum superficie concolor. 

Brom WOSPORAEIE/ 6 / <9. Lint Ie) G0 4eO nO sob ioe lu UIs 
facie obliqua elliptica, nonnumquam plus minusve reniform? 
vel enormi, (Cte aren ae BASIDIA quadrispora, 24.1-35.6 x 
6.6- 6.6-8.8 um, Clavata vel aliquantum enormiter Clavata, 
‘Succum diffusum pallide purpureo-cari =carneum continentia vel 
paulum carnescentia vel hyalina. CHEILOCYSTIDIA specie 
nulla. PILET HYPHAE CUTICULARES plus minusve intertextae, 

aliquid per radios pret tae, stratum distinctum 
antes pferumque 2.9-13.1 um “1 atae, cylitndricae vel 
aliquatenus infTatae, succum diffusum pal Tide vel dilute 
purpureo-carneum vel carneo-rubrum continentes, denique in 
hyalinum pallescentes, tenuiter Eunicatae, part partim 
St liter incrustatae; pileocystidia nulla — PIGMENTUM 


AS nul lum in di rate KOH observatum. vatum. 


0 EE SE Es 0 ee Eee ere 


In solo sub Pino contorta gregarii, altitudine 6500 


SRE Se cite 9 a eaten | te Bid ache Hae Oh ale Pe red Detect tai Ree Rast > Della baa 


OER EEE ERS amare artes 


HOPULYPUS Saha De iniers 32671 su comicatuv Alpine, 
respublicae Californiae, in herb. SFSU conservatus. onservatus. 


PILEUS 20-40 mm broad when fully expanded, convex 
when young becoming plano-convex to more commonly plane 
when fully mature; margin incurved becoming decurved to 
plane, entire, no veil fragments attached to edge and no 
apparent veil fragments on surface; surface dry, glabrous 
to appressed silky-fibril lose, usually shiny with age, 
color brownish orange to brownish red-orange (Cinnamon 
Rufous to Tawny) during all stages of development. Context 
pink, unchanging when exposed or bruised, 1-2 mm thick; 
taste and odor not distinctive. 


LAMELLAE adnate to adnexed, narrow, somewhat wavy 
when mature, close (based on dried specimens), thin, 
several tiers of lamellulae, color dark red (Hays Russet 
to Grenadine Red) during all stages of development; edges 
entire, concolorous with faces. 


STIPE 20-40 mm long, 2-4 mm thick, equal, dry; sur- 
face glabrous except for patches of veil tissue, color 
deep vinaceous to almost concolorous with the surface of 
the pileus, vinaceous mycelium at base; veil fragments 
concolorous with the surface. Context solid and concolor- 
ous with the surface. 


BASIDIOSPORES: °7.7-9.1(-9.5) x 4.8-5.5(-5.8 lune 
profile view elliptical, some more or less reinform or 
somewhat irregularly shaped, in face view elliptical to 
broadly elliptical, verruculose with coarser ornamentation 
towards distal end, light Ochraceous Tawny to light 
yellow-brown with darker brown ornamentation. BASIDIA 4- 
Spored, 24.1-36.5 x 6.6-8.8 um, clavate to somewhat irreg- 
ularly clavate, containing a light purplish pink diffuse 
pigment or slightly pinkish to hyaline. PLEUROCRYSTIDIA 
absent. CHEILOCYSTIDIA apparently absent. SUBHYMENIAL 
HYPAHE compactly interwoven, cylindrical, mostly 2.9-6 um 
wide, color similar to tramal hyphae of lamellae. TRAMAL 
HYPHAE OF LAMELLAE subparallel, more or less interwoven, 
cylindrical to inflated, mostly 3.3-25.6 um wide, at first 
pale to light pinkish purple or pinkish red, often fading 
to paler or hyaline, some with purplish red granules. 
CUTICULAR HYPHAE OF PILEUS more or less interwoven, some- 
what radially arranged, forming a distinct layer, mostly 
2.9-13.1 um wide, cylindrical to somewhat inflated; con- 
taining diffuse pale to light dull purplish pineeoo 
pinkish red pigment, eventually fading to hyaline, thin- 
walled, some finely encrusted; scattered hyphal end cells 
present but no true pileocystidia seen. TRAMAL HYPHAE OF 


35 


PILEUS interwoven, more or less radially arranged, 2.9- 
29.9 um wide, cylindrical to inflated, reddish pink to 
purplish red-pink or pale pinkish purple at first, fading 
to hyaline or nearly so, scattered hyphae with light 
yellow-brown pigment. CORTICAL HYPHAE OF STIPE longitudi- 
nally arranged, subparallel to somewhat interwoven, 2.9- 
18.3 um wide, cylindrical to inflated, purplish red to 
purplish pink fading to hyaline; hymenial elements decur- 
rent on stipe apex, some more or less differentiated as 
caulocystidia, 16.1-42.3 x 10.2-13.1 um (some additional 
hyphal end-cells scattered over surface), clavate to 
broadly clavate more or less cylindrical, or more spheri- 
cal, occasionally catenulate, pale pinkish purple to 
hyaline. CORTINAL HYPHAE cylindrical, mostly 3-5 um wide, 
color similar to cortical hyphae of stipe (few observed). 
CLAMP CONNECTIONS of the normal type, present throughout 
the basidiomata. INTERHYPHAL PIGMENT DEPOSITS not observed 
in KOH. 


Gregarious in soil under lodgepole pine, 6,500'," 
August. 


Collections examined. California. Alpine Co.: H.D. 
fee seo70) 19(holotype, SFSU). Mariposa Co.:,H.D. Thiers 
21106. 


Dermocybe sierraensis appears to be a rare species of 
the western mountains. So far it is only known from the 
higher elevations of the Sierra Nevada Mountains. Studies 
of its pigmentation (Keller and Ammirati, 1983) show a 
close relationship to D. sanguinea, except that the former 
contains much less emodin and dermocybin in the material 
Studied to date. D. sierraensis is distinguished from D. 
Sanguinea by its silky fibril lose, brownish orange to 
brownish red-orange pileus, the lack of ochraceous myceli- 
um on the stipe base, and somewhat larger basidiospores. 
Dermocybe sanguinea var. vitiosa Moser has some character- 
istics (more red-brown color of the pileus, rose colored 
basal mycelium and absence of emodin) in common with D. 
Sierraensis, but an overall comparison indicates that they 


are not the same taxon. 


ACKNOWLEDGEMENTS 


Support for this study from the Graduate School Research 
Fund, University of Washington, and the National Science 
Foundation, Division of Environmental Biology, Systematics 
Biology Program (Grant number DEB-8118972) is greatly 
appreaciated. Loans of specimens from the San Francisco 
State University Herbarium, H. D. Thiers, Curator, and the 
University of Michigan Herbarium, R. L. Shaffer, Curator, 
made this project possible. I wish to thank both institu- 
tions for their cooperation and patience during this 
study. The author expresses thanks to D. P. Rogers for 
preparing the Latin description. 


REFERENCES 


Ammirati, J.F. and A.H. Smith. 1984.) Cort inar 1ussaiieee 
preliminary treatment of species in the subgenus 
Dermocybe, section Sanguinei, in North America, north 
of Mexico. McIlvainea 6(2):54-64. 

Keller, G. and J.F. Ammirati. 1983. Chemotaxonomic signif- 
icance of anthraquinone derivatives in North American 
species of Dermocybe, section Sanguineae. Mycotaxon 
132507 7ONis 

Kelly, K.L. and D.B. Judd. 1965. The ISCC-NBS method of 
designating colors and a dictionary of color names. 
U.S. Nat.. Bur,. Stand, Circ: 553, Washing tong 

Ridgway, R. 1912. Color standards and color nomenclature. 
Published by the author, Washington, D.C. 

Singer, R. 1986. The Agaricales in modern taxonomy. Koeltz 
Scientific Books, Koenigstein, Federal Republic of 
Germany. 


Vol. XXXIV, No. 1, pp. 37-46 January 20, 1989 


LICHENS OF MOUNT DIABLO STATE PARK, 
CONTRA COSTA COUNTY, CALIFORNIA 


Doris E. BALTZO 


2092 Ahneita Drive 
Pleasant Hill, California 94523 


SUMMARY 


A preliminary catalog of the lichens of Mount Diablo State 
Park is presented with updated nomenclature and com- 
ments. Fifty-five genera are reported, including 140 
species (nine are new additions). Most abundant are 
Parmeliaceae, which include nine genera and 24 species. 


INTRODUCTION 


Mount Diablo rises to 3849 ft in Contra Costa County, California, about 40 mi 
ENE of San Francisco. Because of its height and location along the western edge of 
California’s large inland valley, United States surveyors selected Mount Diablo as a 
base meridian (37°53’N) used today in legal descriptions and maps. 

In 1965, when my study began, Mount Diablo State Park did not include North 
Peak or Eagle Peak, which were added during the acquisition of additional parcels of 
land (lichens from these areas are included in the list). The state park now covers 
18,000 acres. 

According to Pampeyan (1963), most of Mount Diablo is underlain by a plug of 
broken and jumbled Upper Jurassic sedimentary, igneous, and metamorphic rock of the 
Franciscan formation, which was thrust upward through surrounding rocks and lubri- 
cated by serpentine veins present on the north side. Exposures of greenstone, chert, 
graywacke, shale, limestone, schist, and conglomerate comprise most of the northern 
end of the mountain, including the summit. Three ridges on the northern side of the 
mountain are North Peak to the northeast (3563 ft), of greenstone, pillow basalt and 
fine-grained basalt; Eagle Peak to the northwest (2369 ft), of diabase; and Deer Ridge, 
just south of Eagle Peak, a grassy area supporting lichens on soil. The southwest side 
of the mountain consists mainly of fossiliferous clastic marine beds ranging from late 
Jurassic to late Miocene. Sandstone is abundant, consisting of 33-50% feldspar, of 
granitic origin. 

On the north-facing slopes of the summit, Quercus chrysolepis Liebm. grows with 
occasional Umbellularia californica Nutt. On lower north-facing slopes on the southern 
side of the mountain, there is an association consisting of Q. agrifolia Nee and Aes- 
culus californica (Spach) Nutt. with some Arbutus menziesii Pursh. Umbellularia 
californica and Acer macrophyllum Pursh are found in canyons. An association of Q. 
douglasii H. & A. and Pinus sabiniana Dougl. occurs on dry slopes. A chaparral cover 
on the south side of the mountain consists chiefly of Adenostema fasciculatum H. & A. 
and Salvia mellifera Greene, with Arctostaphylos glauca Lindl., A. auriculata Eastw., 
and A. manzanita Parry in areas more protected from the fires that occasionally sweep 
through the area. (An extensive fire occurred in 1968). A chaparral association of Q. 
durata Jeps. and A. glauca occurs mostly at low elevations along canyons (Bowerman, 
1944). Juniperus californica Carr. is frequent on rock outcrops, especially at ca. 2990 
ft, and along a series of chert outcrops from the summit to the margin of the chaparral 


38 


1000 ft lower (Bowerman, 1944). Other chaparral plants include Q. wislizenii A.DC. 
var. frutescens Engelm. and Ceanothus cuneatus (Hook.) Nutt. Lichens were also found 
on Salix sp., OQ. dumosa Nutt., and Pinus coulteri D. Don. 

Mount Diablo has a Mediterranean climate. Annual temperature averages 59.3°F, 
with average annual extremes between 26.1°F and 103°F, normally 43-46°F (rainy 
winter, with occasional snow) to 73-82°F (dry summer). Fog from the west, which 
may occur 0.2 of the year on the westerly slope, also encompasses a mixture of smog 
that may limit lichen growth. However, the influence of smog on Mount Diablo 
lichens has not been studied. Many Usnea thalli exhibit excessive formation of fibrils, 
an apparent indicator of unfavorable growth conditions, many of which probably in- 
volve environmental factors detrimental to growth (Tavares, pers. com.). 

A comparison of the following list with Herre’s (1910) enumeration of the lichens 
of the Santa Cruz peninsula reveals the absence of many coastal taxa as well as taxa 
characteristic of the North Coast Ranges [e.g., Nephroma laevigatum Ach. and 
Sphaerophorus globosus (Huds.) Vainio]. A comparison with Hebert and Meyer’s list 
(1984) of lichens of the San Joaquin Experimental Range reveals the presence of some 
taxa found on the eastern slopes bordering the Central Valley of California [e.g., 
Lecidea atrobrunnea (Ramond in Lam. & DC.) Schaerer]. Other taxa from the Sierra 
Nevada foothills have not been found on Mount Diablo [e.g., Peltula zahlbruckneri 
(Hasse) Wetm., Rhizoplaca glaucophana (Nyl. ex Hasse) W. Weber, and R. marginalis 
(Hasse) W. Weber. On the other hand, Bryoria and Usnea were not reported from the 
San Joaquin Experimental Range. 


LIST OF LICHEN SPECIES ON MOUNT DIABLO 


In the following list, an * indicates a new record not included in Baltzo (1970). 
All lichens on the list were found within the present park boundaries except Lecidea 
fuscoatra from Marsh Creek Springs. For nomenclature see Egan (1987) and Tucker 
and Jordan (1978). The name used by Baltzo (1970) or in a published record is indi- 
cated in parentheses when it differs from the lichen name currently used. Voucher 
specimens are in the author’s herbarium. 


Acarospora chlorophana (Wahlenb. ex Ach.) Massal. Literature report (as Lecanora 
chlorophana, Tuckerman (1882). 

*4. fuscata (Nyl.) Arnold. Medium to dark brown, shiny to dull, one to several apo- 
thecia in each areole-squamule. C* fleeting red (section, on bruised cells just un- 
der brown cortical cells). On rocks in sun. (Baltzo 4992c-82FF). 

A. schleicheri (Ach.) Massal. Pale sulphur yellow (as contrasted to lemon yellow of A. 
chlorophana). Apothecia darker than thallus. On sunny rocks. 

Actinogyra see Umbilicaria. 

Alectoria see Bryoria. 

*Aspicilia calcarea (L.) Mudd. Flat to convex, medium gray areoles; dark fimbriate 
hypothallus, cortex K~. On red Franciscan chert. (Baltzo 4990-82FF). 

A. cinerea (L.) Koerber. On shale (as Lecanora cinerea). On rock (as Lecidea tesselata). 

A. gibbosa (Ach.) Koerber (as Lecanora gibbosula). On sandstone, unspecified rock. 

A. laevata (Ach.) Arnold (as Lecanora laevata). On jasper-like rock, sandstone. 

Bryoria oregana (Tuck. ex Willey) Brodo & Hawksw. (as Alectoria oregana). Poor 
specimen, on Adenostoma (collected before 1968 burn), 2500 ft. 

Buellia punctata (Hoffm.) Massal. On Pinus sabiniana. 

Caloplaca bolacina (Tuck.) Herre. Thallus dark lemon yellow, apothecia orange, spores 
12-16 x 6-8 um. On sandstone. 1400 ft. 

C. chrysophthaima Degel. On bark. 

C. decipiens (Arnold) Blomb. & Forss. On sunny siliceous shale, serpentine, jasper; 
widespread. 

C. ferruginea (Huds.) Th. Fr. On bark of deciduous Quercus, chaparral. 

C. laeta Magnusson. On sandstone. 

C. stanfordensis Magnusson. On old Salix. 


She 


C. sp. (as C. elegans), sect. Gasparrinia. Dark red-orange, long narrow lobes to 6 mm 
long and 0.3 mm wide, not spreading at tips; spores ellipsoid to almost lemon 
shaped, 4.8-6.4 x 10-12.8 um. On jasper-like rock in sun, serpentine. 

C. spp. (as C. murorum). Thalli vary from orange with orange pruina to dark orange or 
two-toned red to orange, the lobes 0.5-1 mm wide, 2-3 mm long. On serpentine, 
non-calcareous rock. 

Candelaria concolor (Dickson) B. Stein. On bark of most trees, dead and down wood, 
sandstone, widespread. 

Candelariella vitellina (Hoffm.) Mill. Arg. On soil, moss, rock, sandstone. 

Catapyrenium lachneum (Ach.) R. Sant. (as Lecidea lurida). In noncalcareous soil 
crevice of North Peak saddle. 

Cetraria see Tuckermannopsis. 

Cladonia cervicornis (Ach.) Flowtow. ssp. verticillata (Hoffm.) Ahti (as C. sub 
cervicornis). syn. C. verticillata (Hoffm.) Schaerer. P* yellow to red, K~. On sandy 
soil. +1500 ft. 

C. chlorophaea (Floerke ex Sommerf.) Sprengel. Variously P* dark red, orange; K’; 
KC; C. (chemical species not separated). On sandy soil, sandstone, soil and moss, 

1400-1600 (-3000) ft, North Peak area. 

C. fimbriata (L.) Fr. On soil, 1500 ft. 

C. macilenta Hoffm. On Pinus, +2100 ft. 

C. ramulosa (With.) Laundon (as C. pityrea). On shaded sandstone, 1500 ft. 

Collema furfuraceum (Arnold) Du Rietz. Pointed isidia. On bark, base of Quercus 
agrifolia, on Juniperus californica, on Sambucus, +1500-2900 ft. 

C. nigrescens (Huds.) DC. Isidia granular to globular when present; many apothecia. 
On Quercus bark, +1500-2100 ft. 

Dermatocarpon miniatum (L.) Mann. On rock, near waterfalls and moist vertical sur- 
faces, +2200 ft. 

D. reticulatum Magnusson. On rock near summit, +3849 ft. 

*Dimelaena oreina (Ach.) Norman. On rock, between Muir Area and North Peak, 
2970-3563 ft (Baltzo 777-69R). 

Diploschistes muscorum (Scop.) R. Sant. (as D. actinostomus). On sandstone, Cladonia, 
and on moss. 

*D. scruposus (Schreber) Norman. On rock, Boundary Area (Baltzo 1605-78CC). 

Evernia prunastri (L.) Ach. (as E. prunastri var. sorediifera). Esorediate to sorediate or 
isidiate; juvenile with aspect of Pseudevernia, white below, lobes flat, expanded, 
curled when sorediate; widespread on dead wood; chaparral (Quercus, Adenostoma). 

Flavoparmelia caperata (L.) Hale (as Parmelia caperata). With pustulate or powdery 
lamina! soralia, no pseudocyphellae, rare apothecia with sorediate margin. On 
Quercus bark, sandstone, +1500-1600 ft, widespread. 

Flavopunctelia flaventior (Stirton) Hale (as Parmelia flaventior). Pseudocyphellate, dif- 
fering from Punctelia subrudecta by more yellow-green color and dark brown un- 
derside. On bark, dead wood, sandstone, Acer, chaparral (Arctostaphylos, Quercus), 
widespread. 

Hypocenomyce scalaris (Ach.) ex Liljeblad) M. Choisy (as Lecidea scalaris). On burnt 
wood of Pinus coulteri, +1500 ft. . 

Hypogymnia imshaugii Krog (as H. enteromorpha). Variable in form: small thalli with 
lobes 1-2 mm wide, slightly flattened, regularly dichotomous with pointed tips, in- 
ternodes 1-2 mm long, cortex P* yellow to orange, K* yellow, on twigs of 
Adenostoma; more typical elongate form, gray, lobes 1-3 mm wide, well devel- 
oped, black to dark brown below, almost channeled but hollow, elongating tips 
pointed but not flattened, internodes 4-6 mm long, apothecia large, on Quercus 
wislizenii var. frutescens, Umbellularia californica; narrow, dark gray or brown, 
lobes 1-1.5 mm wide, regular, not flattened, with perforations below near tips, in- 
ternodes to 5 mm long; or compact, light gray-brown, lobes to 2 mm wide, be- 
coming wrinkled-inflated or pustulate centrally, internodes to 2 mm long, on 
Quercus, Pinus bark. 

H. tubulosa (Schaerer) Havaas. Tubes with tips of soredia around edge. On bark, 
Quercus. 


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42 


Koerberia biformis Massal. On Aesculus, 2150 ft. 

Lecanora see also Aspicilia. 

L. caesiorubella Ach. ssp. merrillii Imsh. & Brodo (as L. pallida group). On bark, 
twigs. Also literature report (Imshaug & Brodo, 1966). 

L. hagenii (Ach.) Ach. On sandstone, +1500 ft. 

L. mellea W. Weber (as L. bolcana, L. muralis var. diffracta, L. garovaglii). Rich, 
dark honey color, the lobate margins with thin black edge, the lobes breaking up 
into several thinner lobe tips, thallus very variable, apothecial disc and margin 
same color as thallus, inconspicuous. In sun on jasper-like rock, quartz, igneous 
rock, shale, often in association with Aspicilia sp. Also literature report (Weber, 
1975). 

L. muralis (Schreber) Rabenh. (in part as L. muralis var. versicolor). On sandstone, 

jasper and other rock. 

L. pacifica Tuck. or similar to it in appearance. No crystals in the apothecia. Thallus 
K?* yellow. On twigs, 2500 ft. 

L. rupicola (L.) Zahlbr. Pruinose, pale. On rock, 2380 ft. 

Lecidea see also Aspicilia, Hypocenomyce, Lecidella, Pannaria, Psora. 

L. atrobrunnea (Raymond in Lam. & DC.) Schaerer. Shiny red brown thallus, areole- 
squamules thin to thick and convex; waxy epicortex is thin; cortex UV* yellow- 
cream; medulla I*. Apothecial discs smooth in appearance. Similar to Rhizocarpon 
bolanderi, which is UV" and has much smaller areoles and rough apothecial discs. 
On shale. 

*7.. fuscoatra (L.) Ach. Semi-shiny to dull dark brown thallus, areole-squamules thin- 
ner than in L. atrobrunnea, with more intervening space showing black 
hypothallus; squamules becoming irregularly crenulate and turned up marginally, 
UV; apothecia may be pruinose. Between Sunset Area and Rocky Point, 2200 ft 
(Baltzo 90-65C); collected outside the State Park, on volcanic plug at Marsh Creek 
Springs (Baltzo 9048-87JJ) (several volcanic plugs were mentioned by Pampeyan, 
1963, northeast of the base of Mount Diablo along Marsh Creek Rd.). 

L. mannii Tuck. Literature reports (Tuckerman, 1888; Herre, 1910). Thalli on "vol- 
canic rock," "Diablo" (Bolander 208a,b: see Tuckerman, 1888) (FH) are dull me- 
dium brown to pale brownish cream color, the areoles crowded, their edges turning 
downward; hypothallus not clearly visible between areoles. Cortex UV; medulia 
thick, white, UV pale greenish, I. Precise collection locality not known and rocks 
vary. 

Lecidea sp. (as L. mannii). Much darker brown than L. mannii of Bolander; shinier, 
UV’, medulla I (Baltzo 110-65C). 

*TLecidea sp. Pale cream, dull; large slightly convex, crowded areoles, paler than 
Bolander specimens of L. mannii and having larger areoles. With Rhizocarpon 
bolanderi, near summit. Dull rusty orange under UV, perhaps because of lack of a 
dark brown pigment. On rock. 

*Tecidea sp. Medium gray brown granular eyed broken into areolate segments; 
apothecia black. On sandstone. 

Lecidella euphora (Floerke) Hertel (as Lecidea glomerulosa). C°, K°. On Juniperus 
twigs, Pinus, dead or down wood. 

Lepraria neglecta (Nyl.) Lettau. On sandstone, 1500 ft. 

Leptochidium albociliatum (Desmaz.) M. Choisy (as Polychidium albociliatum). On moss 
and soil, among grasses, mossy rock. 

Leptogium californicum Tuck. On shaded sandstone. 

L. corniculatum (Hoffm.) Minks (as L. palmatum). On moss, soil, rock. 

L. furfuraceum (Harm.) Sierk. On Acer, Aesculus. 

L. lichenoides (L.) Zahlbr. On moss, soil. 

L. minutissimum (Floerke) Fr. On soil. 

Letharia columbiana (Nutt.) Thomson. On burnt bark, Pinus sabiniana; immature, 
isidiate, nonsorediate. 

Lichinella stipatula Nyl. Literature report (Henssen, 1963; W. A. Weber collection). 

Melanelia fuliginosa (Fr. ex Duby) Essl. (as Parmelia glabratula). Medulla Ct red, 
shiny cylindrical isidia. On bark of Quercus, Pinus. 


43 


M. glabra (Schaerer) Essl. (as Parmelia glabra). Medulla C* red, thallus minutely 
pubescent, lowerside dark brown to black. On bark of Quercus, Pinus, Sambucus, 
Adenostoma, Juniperus, chaparral, widespread. A form referred to as "subglabra" 
differed by its paler olive-green color perhaps due to shady conditions, its lower 
side pale in a broad band on the margins and medium brown centrally. Loosely 
adnate on mosses on bark. 

M. glabroides (Essl.) Essl. (as Parmelia "pseudoglabra"). Medulla C* red, KC? red; 
lobes shiny scalloped-crenulate, reticulate-ridged, lacunose, in cushions. On sand- 
stone, soft shale, moss, easily removed from substrate. 

M. incolorata (Parr.) Essl. (as Parmelia elegantula). Medulla C°, warts not prominent; 
isidia tround at first, not pinched at base, becoming cylindrical, sometimes in 
clusters, infrequently branched. Thallus sometimes pruinose. On bark, twigs, sand- 
stone. 

M. multispora (A. Schneider) Essl. (as Parmelia multispora). Medulla C’, thallus like 
M. subolivacea but thinner, asci with more than 8 globose to subglobose spores. On 
bark, Quercus. 

M. subargentifera (Nyl.) Essl. (as Parmelia subargentifera). Medulla C* red; marginal 
labriform soralia, minutely pubescent upper side. On sandstone, moss. 

M. subaurifera (Nyl.) Essl. (as Parmelia subaurifera). Medulla C* red; soralia laminal, 
punctiform, tending to yellowish to greenish; isidia cylindrical, short. On bark, 
Quercus; frequent. 

M. subelegantula (Essl.) Essl. (as Parmelia "pseudoaspera"). Medulla C’; warts with 
white area a top that is sometimes depressed; starting at thallus margins, warts be 
come transformed into dull cylindrical isidia in center; isidia may becom coralloid 
or lobulate and drooping. On bark, Quercus sp., Q. wislizenii, Pinus, Ceanothus 
cuneatus, Umbellularia, to 3849 ft. 

M. subolivacea (Nyl. in Hasse) Essl. (as Parmelia subolivacea). Medulla C’; warts on 
mature thalli irregular in shape, never isidiate-coralloid or lobulate. On bark, 
Quercus. 

Neofuscelia loxodes (Nyl) Essl. (as Parmelia isidiotyla). Medulla C; cortex C* dark 
blue gray; isidia coarse, dull, globular clusters that may break to show white 
medulla or resemble soredia. On rock, shale, sandstone. 

Normandina pulchella (Borrer) Nyl. Minute pale green ear-like squamules that become 
sorediate on the edges. On bark of Acer and sandstone. 

Ochrolechia subpallescens Vers. Thallus and thick apothecial margin C* red. On bark, 
Acer and Quercus. 

O. upsaliensis (L.) Massal. Literature report (Herre, 1910). Thallus C.. On sandstone 
and over mosses. 

Pannaria leucophaea (Vahl.) P. Jorg. (as Lecidea demissa). Minute, dull, medium 
brown. On sandstone. 

*P. leucostictoides Ohllson. Pale tan, becoming bluish pruinose; isidia blue-gray. On 
Quercus (Baltzo 47b-65A). 

P. praetermissa Nyl. in Chyd. & Furuhj. (as Parmeliella praetermissa). Dark brown; 
isidia dark gray. On moss on rock. 

Parmelia see also Flavoparmelia, Flavopunctelia, Melanelia, Neofuscelia, Parmelina, 
Parmotrema, Punctelia, and Xanthoparmelia. 

P. saxatilis (L.) Ach. Isidiate. On bark of Quercus, moss, sandstone. 

P. sulcata Taylor. Sorediate on ridges. On bark, Quercus, Pinus, Juniperus, on shaded 
sandstone, widespread. 

Parmeliella see Pannaria. 

Parmelina quercina (Willd.) Hale (as Parmelia quercina). On bark, Quercus sp., Q. 
wislizenii, Pinus, Juniperus, Umbellularia, widespread. 

Parmotrema arnoldii (Du Rietz) Hale (as Parmelia arnoldii). Medulla K’; soralia on 
small lobes, submarginal. On bark of Quercus agrifolia. 

P. chinense (Osbeck) Hale & Ahti (as Parmelia perlata). Medulla K* yellow; soralia on 
revolute lobe tips. On Quercus. 

P. stuppeum (Taylor) Hale (as Parmelia stuppea). Medulla K*; soralia linear on raised 
margins; cilia long; lobes wide. On sandstone. 


44 


Peltigera collina (Ach.) Schrader. On moss on sandstone in shade. 

Peltula euploca (Ach.) Ozenda & Clauz. On rock, 3700 ft. 

Pertusaria albescens (Huds.) M. Coisy & Werner. On Quercus. 

P. amara (Ach.) Nyl. Bitter taste. On bark, Quercus. 

P. chiodectonoides Bagl. ex Massal. On rock. 

P. lecanina Tuck. On Quercus, bark. 

Phaeophyscia orbicularis (Necker) Moberg (as Physcia orbicularis). On bark, Quercus, 
Juniperus, Aesculus. 

Phlyctis argena (Sprengel) Flotow. On bark. 

Physcia adscendens (Fr.) H. Olivier. Hooded soralia. On Quercus, Juniperus, 
widespread. 

P. aipolia (Ehrh. ex Humb.) Fuernr. White-spotted, cortex K* yellow. On Acer, Aes- 
culus, Juniperus. 

P. alba (Fee) Mill. Arg. var. obsessa (Mont.) Lynge (as Physcia alba). On dead wood 

(Juniperus?). 

. albinea (Ach.) Nyl. On red jasper-like rock. 

. callosa Nyl. On shale, shaded sandstone, moss on rock. 

. cascadensis Magnusson. On Quercus. 

. mexicana B. de Lesd. On Quercus agrifolia. 

. millegrana Degel. On heavy rock with quartz. 

. Dhaea (Tuck.) Thomson. On Quercus agrifolia. Also literature report (Thomson, 

1963). 

. semipinnata (J.F. Gmelin) Moberg (as P. leptalea). On Adenostoma. 

. Stellaris (L.) Nyl. On Adenostoma, Quercus, Juniperus, chaparral. Four forms. 

. tenella (Scop.) DC. in Lam. & DC. On Quercus, Juniperus, over other lichens. 

Physconia detersa (Nyl.) Poelt (as Physcia grisea f. detersa). On sandstone. 

P. distorta (With.) Laundon (as Physcia pulverulenta). On bark of Quercus, Acer, 
Juniperus, Umbellularia, chaparral. Three forms. 

P. enteroxantha (Nyl.) Poelt (as Physcia grisea f. enteroxanthella). On Quercus, Sam- 
bucus, shaded sandstone. 

P. grisea (Lam.) Poelt. On Juniperus, Quercus, Umbellularia, sandstone. 

P. muscigena (Ach.) Poelt. On dead and down wood, soil, moss. 

Polychidium see Leptochidium. 

Pseudocyphellaria anomala Brodo & Ahti. On moss at base of Quercus agrifolia. 

P. anthraspis (Ach.) Magnusson. On base of Quercus agrifolia. 

Psora californica Timdal (as Lecidea globifera). On soil in rock crevice. 

P. globifera (Ach.) Massal. Medulla K~. See Schneider (1979), Timdal (1986). On soil, 
sandstone. 

P. nipponica (Zahlbr.) Schneider (as Lecidea novomexicana). In rock crevice. 

Punctelia subrudecta (Nyl.) Krog (as Parmelia subrudecta). Differs from Flavopunctelia 
flaventior by mineral blue-gray-green color and pale beige underside. On bark of 
Quercus, Pinus, Adenostoma, chaparral. 

Ramalina farinacea (L.) Ach. Typical form with narrow laciniae 1-2 mm wide, bran- 
ching at 4-5 mm intervals, marginal soralia P* rusty orange, medulla P* rusty 
orange, cortex with two layers, K* yellow, on bark of Quercus agrifolia and Acer. 
Shorter tufted form, branching at 3-4 mm intervals, soralia only P* rusty orange, 
cortex without inner layer, on bark. Short form to 4.5 cm long, non-fistulose, ex- 
panding finger-like soraliate apices with marginal isidioid outgrowths, no inner 
cortex, on Quercus sp., Q. agrifolia. Broad form, laciniae to 2 mm wide, soralia on 
laciniae and margins, medulla P’, soralia P* pale orange, no inner cortex. On Q. 
agrifolia. 

R. leptocarpha Tuck. On bark. 

R. menziesii Tayl. Coarse, rigid nets. On bark or draped on branches, infrequent. 

*Rhizocarpon bolanderi (Tuck.) Herre. Thallus shiny, dark red-brown, similar to 
Lecidea atrobrunnea. Apothecial disc appears rough. On rock near summit, 3800 ft. 
(Baltzo 892b-69T). 

R. ferax Magnusson. On rock, 2200-3849 ft. 


me) ast as} as) sis, 


as) ash as 


45 


Rinodina hallii Tuck. On bark. 

Sticta fuliginosa (Hoffm.) Ach. On shaded sandstone, Quercus, Pseudocyphellaria 
anomala. 

Thelomma mammosum (Hepp in Hartung) Tibell (as Cypheliopsis bolanderi). On 
sunny rocks. 

Tuckermannopsis chlorophylla (Willd. in Humb.) Hale (as Cetraria chlorophylla). On 
Adenostoma, shaded sandstone. 

T. merrillii (Du Rietz) Hale (as Cetraria merrillii). On Adenostoma, Quercus bark. 

T. orbata (Nyl.) Lai (as Cetraria orbata). On bark, on Quercus. 

Umbilicaria phaea Tuck. On sandstone, metamorphic rock, shale, widespread. 

U. polyphylla (L.) Baumg. On rock. 

U. polyrrhiza (L.) Fr. (as Actinogyra polyrrhiza). On shaded rocks, 3000 ft. 

Usnea arizonica Mot. s. lat. Medulla K* red. On bark of Acer, Quercus agrifolia. 

U. cavernosa Tuck. (as U. trichodea). On Quercus. 

Usnea spp. (tufted and pendent taxa). On Quercus, chaparral, 1500-2500 ft. 

Xanthoparmelia cumberlandia (Gyelnik) Hale (as Parmelia cumberlandia). On sandstone, 
shale, in sun, 2100 ft. 

X. lineola (Berry) Hale (as P. lineola). On sandstone, quartz, jasper. 

X. mexicana (Gyelnik) Hale (as P. mexicana). On sunny rock, sandstone, quartz, or 
jasper. 

X. novomexicana (Gyelnik) Hale (as P. novomexicana). On shale. 

X. taractica (Kremplh.) Hale (as P. ioannis-simae, P. taractica). Loosely attached to soil 
on rock, on crumbly shale, on quartz. 

Xanthoria candelaria (L.) Th. Fr. On Quercus, Umbellularia. 

X. fallax (Hepp in Arnold) Arnold. On moss and rock, sandstone. 

X. lobulata (Floerke) B. de Lesd. On calcareous sandstone. 

X. parietina (L.) Th. Fr. On bark (Quercus?), 2700 ft; on Aesculus, Donner Canyon. 

X. polycarpa (Hoffm.) Rieber. On Quercus agrifolia, other Quercus spp, Pinus 
sabiniana. 


ACKNOWLEDGEMENTS 


I would especially like to thank Dr. Harry D. Thiers for setting me on this course 
many years ago. I remember well his energetic, cheerful attitude and am grateful to 
him for his encouragement and patience. 

I wish to express my appreciation to Dr. Isabelle Tavares for her review of the 
manuscript; to Dr. T. Elliot Weier for his patience, comments and suggestions, and also 
for reviewing the manuscript; and to Ethel Cole for her generosity in preparing the 
camera-ready copy. 

Grateful thanks go to all those who have assisted me, particularly with identifica- 
tion. I would especially like to thank Dr. Mason E. Hale, Dr. W. L. Culberson, Dr. J. 
W. Thomson, Dr. T. Ahti, Dr. J. R. Laundon, Dr. Per Jorgensen, Dr. Isabelle Tavares, 
and Dr, T. Elliot Weier. I am indebted to Dr. Donald Pfister, Farlow Herbarium, for 
the opportunity to study the Bolander specimens of Lecidea mannii. 


LITERATURE CITED 


Baltzo, D. E. 1970. A study of the lichens of Mount Diablo State Park. M.A. thesis, San 
Francisco State Univ. 

Bowerman, M. L. 1944. The flowering plants and ferns of Mount Diablo, California. 
Their distribution and association into plant communities. Gillick Press, Berkeley. 

Egan, R. S. 1987. A fifth checklist of the lichen-forming, lichenicolous and allied 
fungi of the Continental United States and Canada. The Bryologist 90: 77-173. 

- Hebert, J. R. & R. W. Meyer. 1984. Lichens of the San Joaquin Experimental Range, 
California. The Bryologist 87: 251-254. 

Henssen, A. 1963. Eine Revision der Flechtenfamilien Lichinaceae und Ephebaceae. 
Symb. Bot. Ups. 18(1): 1-123. 

Herre, A. W. C. T. 1910. The lichen flora of the Santa Cruz Peninsula, California. Proc. 
Washington Acad. Sci. 12: 27-269. 


46 


Imshaug, H. A. & I. M. Brodo. 1966. Biosystematic studies on Lecanora pallida and 
some related lichens in the Americas. Nova Hedwigia 12: 1-59. 

Pampeyan, E. H. 1963. Geology and mineral deposits of Mount Diablo. Special Report 
80, Calif. Div. Mines & Geology, San Francisco. 

Schneider, G. 1979. Die Flechtengattung Psora sensu Zahlbruckner. Versuch einer 
Gliederung. Bibl. Lichenologica 13: 1-291. 

Thomson, J. W. 1963. The lichen genus Physcia in North America. Beih. Nova Hed- 
wigia 7: 1-172. 

Timdal, E. 1986. A revision of Psora (Lecidaceae) in North America. The Bryologist 
89: 253-275. 

Tucker, S. C. & W. P. Jordan. 1979. A catalog of California lichens. Wasmann J. Biol. 
36: 1-105. 

Tuckerman, E. 1882. A synopsis of the North American Lichens: Part I., Comprising 
the Parmeliacei, Cladonei, and Coenogoniei. S. E. Cassino, Boston. 

. 1888. A synopsis of the North American Lichens: Part II., Comprising the 

Lecideacei, and (in part) the Graphidacei. H. Willey, New Bedford. 

Weber, W. A. 1975. Two new species of Lecanora, section Petrasterion, with a key to 
North American species. The Bryologist 78: 206-210. 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 47-53 January 20, 1989 


THE GENUS RHODOCYBE: NEW COMBINATIONS AND A REVISED 
KEY TO SECTION RHODOPHANA IN NORTH AMERICA 


TIMOTHY J. BARONI 


Department of Biological Sciences 
State University of New York 
College at Cortland, Cortland, NY 13045 


AND 


DAVID L. LARGENT 


Department of Biology, Humboldt State University 
Arcata, CA 95521 


ABSTRACT 


Entoloma trachyosporum and its varieties are placed in 
Rhodocybe. Rhodocybe carlottae var. carlottae is consider- 
ed a synonym of R. trachyospora var. trachyospora, while 
R. carlottae var. vinacea is placed in R. trachyospora as a 
new variety. A key to Rhodocybe section Rhodophana in 
North America is presented. 


KEY WORDS: Rhodocybe, section Rhodophana, nomencla- 
ture, key to species for North America. 


INTRODUCTION 


It is appropriate that this report concerns a furthering of our 
knowledge of fungi of the Pacific Northwest, since the scientist honored 
by the collection of papers in this festschrift has spent most of his dis- 
tinguished professional career working towards that end. We hope this 
latest addition to our knowledge of the California mycota will please 
him. 


The genus Rhodocybe Maire has recently received careful attention 
in North and Central America (Lennox, 1979; Baroni, 1981; Halling & 
Baroni, 1985; Redhead & Baroni, 1986; Ovrebo & Baroni, 1988) because 
the species of Rhodocybe can occasionally be a distinctive, though usual- 
ly not a prominent, part of the mycota of a given area. Although a 


48 


revision of Rhodocybe (Baroni, 1981) has helped clarify the unique fea- 
tures that circumscribe this member of the Entolomataceae Kotlaba & 
Pouzar, there still remains much to be learned about the number, dis- 
tribution and phylogeny of species in Rhodocybe for North America, 
and on a world wide scale. 


In this past collecting season, during the months of November and 
December in Humboldt and Mendocino Counties of California, we had 
the opportunity to collect all of the varieties of Entoloma trachyosporum 
Largent, i.e. E. trachyosporum var. trachyosporum, E. trachyosporum var. 
griseoviolaceum Largent and E. trachyosporum var. purpureoviolaceum 
Largent. An examination of the unusual basidiospores of each of these 
collections revealed to the senior author that these taxa clearly belong in 
Rhodocybe. 


TAXONOMY 


In the sense of Singer (1986), Rhodocybe is one of three genera 
placed in the Entolomataceae (Clitopilus and Entoloma sensu lato are the 
other two, but see Largent & Benedict, 1971; Baroni, 1981; and Baroni 
& Petersen, 1987). All taxa placed in the Entolomataceae possess the 
following suite of characters: spore deposit pinkish, flesh, vinaceous or 
rarely grayish; spores always distinctly angular or rounded angular in 
polar view, or the spores are angular in all views; spore walls evenly 
cyanophilic. Rhodocybe can only be accurately identified and separated 
from Clitopilus and Entoloma sensu lato by the unique, mostly isolated 
pustulate or tuberculate-warty ornamentation of its basidiospores. The 
basidiospores of Clitopilus are never angular in profile view, but possess 
distinct to obscure longitudinal ridges running the length of the spores, 
while the basidiospores of Entoloma sensu lato are provided with short 
interconnected ridges, which make the basidiospores distinctly angular in 
all views. The basidiospores of Entoloma sensu lato never have isolated 
pustulate or tuberculate-warty ornamentation. 


After a study of the macroscopic and microscopic characters of all 
the collections of E. trachyosporum available to us, it became apparent 
that the recently described Rhodocybe carlottae var. carlottae Redhead & 
Baroni (Redhead & Baroni, 1986) from the Queen Charlotte Islands of 
British Columbia is conspecific with E. trachyosporum var. 
trachyosporum. However, in the case of R. carlottae var. vinacea Red- 
head & Baroni, the colors of the stipe, the lamellae and the pileus con- 
text indicate that this taxon is yet another distinct color variant of E. 
trachyosporum from the Pacific Northwest of North America. 


The following new combinations and synonymy are necessary: 


Rhodocybe trachyospora (Largent) Baroni & Largent comb. nov. 


= Entoloma trachyosporum Largent, Madrojfio 22: 369. 1974. 
= Rhodocybe carlottae Redhead & Baroni, Canad. J. Bot. 64: 1451. 
1986. 


49 


Rhodocybe trachyospora var. griseoviolacea (Largent) Baroni & Largent 
comb, nov. 


= Entoloma trachyosporum var. griseoviolaceum Largent, Madrono 
2201041974; 


Rhodocybe trachyospora var. purpureoviolacea (Largent) Baroni & 
Largent comb. nov. 


= Entoloma trachyosporum var. purpureoviolaceum Largent, Madrono 
22: 371. 1974. 


Rhodocybe trachyospora var. vinacea (Redhead & Baroni) Baroni & 
Largent comb. noy. 


= Rhodocybe carlottae var. vinacea Redhead & Baroni, Canad. J. 
Bot. 64: 1451. 1986. 


After referring to our field data, and after a reexamination of the 
microscopic characters of the type collections of FE. trachyosporum and 
its varieties, the following information can be added to the original des- 
criptions (Largent, 1974). 


Rhodocybe trachyosporum var. trachyosporum Figs. 1 & 2 


Basidiospores 6-8(-9) (without apiculus), 7-10 (including apiculus) 
x 6-7(-8) wm, subglobose to ovoid or short-broad-ellipsoid, often 
obscurely to distinctly angular in profile view, rounded angular (6-9 
facets) in polar view, weakly to moderately undulate-pustulate in all 
views, walls evenly cyanophilic but weakly so on many older (larger) 
spores. Caulocystidia present in inconspicuous clusters or scattered at 
apex of stipe, 22-45 x 8-10 um, often capitate or subcapitate and 
pedicillate, or clavate or broadly ventricose, thin-walled, hyaline, with 
clamp connections. 


Rhodocybe trachyosporum vars. griseoviolaceum and purpureoviolaceum 


The microscopic features for both of these varieties are in- 
distinguishable from those of var. trachyosporum. 

These taxa were originally reported from a number of different 
counties around the greater Seattle, Washington area (Largent, 1974) as 
occurring under various conifer species. For example, R. trachyosporum 
var. trachyosporum was collected from beneath Pseudotsuga menziesii 
(Mirb.) Franco, R. trachyosporum var. griseoviolaceum was found 
beneath P. menziesii, Thuja plicata D. Don. and Tsuga heterophylla 
(Raf.) Sarg., while R. trachyosporum var. purpureoviolaceum was col- 
lected beneath TJ. plicata. The collections of R. trachyosporum var. 
trachyosporum (Baroni 5794, 5801 and 5802) and var. griseoviolaceum 
(Baroni 5803) from California were typically found under Picea 
sitchensis (Bong.) Carr. or P. sitchensis mixed with Sequoia sempervirens 


50 


Endl., while R. trachyosporum var. purpureoviolaceum (Baroni 5856) was 
found under a mixture of P. menziesii, T. heterophylla, S. sempervirens 
and Lithocarpus densiflora (H. & A.) Rehd. It appears then that these 
Rhodocybes may be found under various kinds of conifers throughout 
the Northern Coastal Coniferous Forest and the Redwood Forest (Munz, 
1970) of California and the Pacific Northwest. 


Using the infrageneric classification scheme of Baroni (1981), this 
group of taxa would be placed in Rhodocybe section Rhodophana 
(Kiihner) Singer based on the presence of clamp connections and the 
lack of hymenial pseudocystidia. Rhodocybe trachyospora and its variet- 
ies would be closely aligned with R. mycenoides, R. speciosa and R. 
priscua due to their subglobose basidiospores, which are often angular in 
profile view. Rhodocybe mycenoides is known only from South Amer- 
ica, while these other taxa with subglobose basidiospores are found in 
North America. 


Key to North American taxa of Rhodocybe section Rhodophana 


1. Basidiospores ellipsoid or amygdaliform and not angular in profile 


VIEW... ccccscocgescecncsseescostecavcoysleuntescecbecsiseeccenteustseratns cenit este iat ta 2 
1. Basidiospores subglobose to short-broad-ellipsoid and often angular 
IM PrOf tle’ VIEW.) loco.csecc-cueesacssatcecascesecacesoeteyencocunch st bento site tatnmmm nn 3 


2. Lignicolous; stipe eccentric; pileus light buff but soon developing 
pinkish hues; cheilocystidia thin-walled and clavate to 
sphaeropedunculate; laterostratum of lamellar trama gela- 
{INIZEd Sue ee ee eee: R. eccentrica Baroni & Ovrebo 

2. Terrestrial; stipe central; pileus ochraceous-tawny to orange cin- 
namon but fading to pinkish cinnamon to cinnamon buff; 
cheilocystidia not differentiated; lamellar trama not gela- 
tiNlZed ee el erat rete neem R. nitellina (Fr.) Singer 


3. On decayed wood in coniferous forests; pileus tan or yellowish 
brown; stipe yellow or pale orange; end cells in pileipellis inflated 
as variously shaped pilocystidia; hyphae of lamellar trama and pileal 
context dextrinoid see ee R. speciosa Lennox ex Baroni 

3. Terrestrial in coniferous forests; pileus differently colored; 
pilocystidia lacking; trama and context not dextrinoid....................66 4 


4. Under pines on needle beds; pileus and stipe umber or hazel, 
Stipe eventually paler than pileus, becoming vinaceous buff or 
buff, stipe dry, finely striate and with a patchy hoary covering; 
odor of plastic; caulocystidia mostly clavate to cylindric............. 
eee SOREN Meroe tvs Deon anim ln Aenea: MeN SRA R. priscua Baroni 

4. Under sitka spruce, western hemlock, Douglas fir, western red 
cedar, redwood, etc. (but not under pine) on needle beds or 


51 


Figs. 1-2: Rhodocybe trachyospora var. trachyospora. 1. Basidiomes, x] 
(Baroni 5794). 2. Basidiospores, scale bar = 10 wm (Largent 2169, 
HOLOTYPE). 


52 


bare soil; pileus mostly with grayish hues mixed in with 
browns, yellow-browns, vinaceous, purplish or fuscous colors, 
stipe differently colored than above, lubricous in wet or humid 
weather, glabrous or finely fibrillose-appressed striate or 
rimulose; odor not distinctive or somewhat fruity fragrant; 
caulocystidia variously shaped from cylindric to broadly 
ventricose, but most often sphaeropedunculate or pedicellate- 
Capitate si AR A) a, eyes. 


5. Stipe off-white to pale grayish to pale grayish brown, context con- 


COIOTOUS 1. i287scc secs ea) Cla car tae tens ca ceed ven emeRR Etuenee ct set oe R. trachyospora 
5. Stipe dark blue to bluish gray, violaceous gray, or pale vinaceous 
gray, context concolorous: or’slightly paler.....:1.......,,...slesneneeneeee ae 6 


6. Pileus dark grayish brown, becoming paler yellowish-brown over 
the margin with loss of moisture; lamellae pallid to grayish 
brown at first; pileus context sordid flesh ‘buff......................000000 
TN EU MUS Meso ah AR aa LEM AO) Rage, R. trachyospora var. griseoviolacea 

6. Pileus dark purplish brown or reddish brown or fuscous purplish 
with vinaceous to reddish gray margin at first; lamellae bluish 
gray or livid to pale vinaceous at first; pileus context deep blue 
to grayish blue: or pale vinaceous s...2.......2...100:s4-s-0eaenee fi 


7. Stipe dark blue to bluish gray; lamellae bluish gray at first; pileus 
context deep blue to bluish: gray.....20)........-cscels casusedossecseen eee 
pO ea en Re rene Pp ASTER MAN KIER BR ait R. trachyospora var. pur pureoviolacea 

7. Stipe pale vinaceous gray; lamellae livid to pale vinaceous at first; 
pileus context pale vinaceous................... R. trachyospora var. vinacea 


For complete descriptions of these taxa refer to Baroni (1981), Ov- 
rebo & Baroni (1988), Redhead & Baroni (1986) and Largent (1974). 
All color terms used in the keys are adapted from Kornerup and Wans- 
cher (1978) and Rayner (1970). 


ACKNOWLEDGMENTS 


A grant from the Buffalo Society of Natural Sciences to the senior 
author (grant no. 223-0205A, The Research Foundation of the State 
University of New York) is directly responsible for the research which 
led to this report. Ms. Dawn Van Hall, photographic technician for the 
State University College at Cortland, is gratefully acknowledged for her 
assistance in producing the black and white negatives from Kodachrome 
slides of the basidiomes of R. trachyospora var. trachyospora. Dr. Roy 
Halling performed the presubmission review, and we are sincerely grate- 
ful for his time and expertise. 


53 


LITERATURE CITED 
BARONI, T. J. 1981. A revision of the genus Rhodocybe Maire 
(Agaricales). Beih. Nova Hedwigia. 67: 1-194. 
BARONI, T.J. & R. H. PETERSEN. 1987. Rhodocybella: a new genus 
in the Entolomataceae. Mycologia 79: 358-361. 


HALLING, R. E. & T. J. BARONI. 1985. Rhodocybe pulchrisperma 
(Entolomataceae): a new species from North America. Brittonia 37: 
182-185. 


KORNERUP, A. & J. H. WANSCHER. 1978. Methuen handbook of 
colour. 3rd ed. Eyre Methuen, London. 

LARGENT, D. L. 1974. New or interesting species of Claudopus and 
Entoloma from the Pacific coast. Madrofio 22: 363-373. 

PARGENT, D..-L, & R. G. BENEDICT. 1971. Studies in the 
rhodophylloid fungi. I. Generic concepts. Madrofio 21: 32-39. 

LENNOX, J. W. 1979. Collybioid genera in the Pacific Northwest. 
Mycotaxon 9: 117-231. 

MUNZ, P. A. 1970. A California flora. Univ. of California Press, 
Berkeley. 

OVREBO, C. L. & T. J. BARONI. 1988. Three new species of 
Rhodocybe from Costa Rica. Mycologia 80:(in press) 

RAYNER, R. W. 1970. A mycological colour chart. Commonwealth 
Mycological Institute, Kew. 


REDHEAD, S. A. & T. J. BARONI. 1986. Clitopilus fuscogelatinosus 
and Rhodocybe carlottae, new species in the Entolomataceae 
(Agaricales) from Canada. Canad. J. Bot. 64: 1450-1452. 


SINGER, R. 1986. The Agaricales in modern taxonomy. 4th ed. Koeltz 
Scientific Books, Koenigstein. 


Vol. XXXIV, No. 1, pp. 55-63 January 20, 1989 


QUALITY CONTROL FACTORS FOR 
ALTERNARIA ALLERGENS 


Harriet A. Burge, Marion E. Hoyer, William R. Solomon 
Department of Internal Medicine 
University of Michigan 
Ann Arbor, MI 48109-0529 


Emory G. Simmons 
Mycclogical Services 
717 Thornwood Road 
Crawfordsville, IN 47933 


Janet Gallup 
Specialty Labs of Orange County 
725 W. La Veta 
Orange, California 92640 


Alternaria species are among the best known and most 
clinically important sources of fungal allergens (Lehrer, et 
al. 1983). To assess the role of environmental exposures to 
these allergens in human disease, fungus materials are grown 
in culture and extracted in aqueous media to produce 
reagents used for skin testing or in-vitro tests for the 
presence of allergen specific antibodies. The same 
materials are often used for desensitization therapy 
(immunotherapy). However, methods used in the production of 
Alternaria allergen materials for diagnosis and treatment as 
well as research utilize single isolates often without 
expert identification and improperly named (Vijay, et al. 
1986) in spite of the fact that allergen content is known to 
vary with species and strain used. (Burge et al 19xx) Most 
methods are based on mycelial growth and metabolic products 
rather than spores (Solomon et al, 1980) and fail to take 
into account the inherent variability that is the bane of 
mycologists interested in Alternaria taxonomy (Norman, 
190.25)5, 


We have studied spore-derived allergens from airborne 
strains of Alternaria alternata and morphologically similar 
taxa as well as a series of verified isolates of A. 
alternata known to be stable in culture. 


56 


METHODS 


Four airborne isolates of Alternaria (entries 1-4 in 
Table I]: two A. alternata, and two unidentified species 
that were assumed to be A. alternata by mycologists not 
trained in Alternaria identification were maintained in 
culture on Sabouraud’s broth (Hoffman, et al. 1981). On 
five occasions (12/84, 2/85, 5/85, 11/85, 5/86) spores and 
mycelium were harvested separately. Spore preparations, 90- 
95% pure by microscopic examination, were obtained by the 
method of Kozak and Gallup (U.S. patent #4,280,000). 
Briefly, a suspension of spores in distilled water was 
inoculated onto the surface of Sabouraud’s Dextrose Agar 
(DIFCO, Detroit, MI) and spread evenly over the surface 
with a flame-sterilized bent glass rod. An overlay of 
Whatman filter paper #54 and a nondigestible material type 
TX 1040 (Pallflex Co.,Putnam, CN) were smoothed onto the 
inoculated agar surface. After incubation under room lights 
at room temperature for three weeks, the overlay growth mat 
was removed, culture medium discarded, and the overlay 
growth mat placed back in the petri dish and allowed to dry 
for 3-4 days. Spores were harvested by scraping the overlay 
surface with a dull knife. The resulting spore/mycelium 
mixture was separated in a 45 micron sieve (USA Standard 
Testing Sieve, Dual Manuf. Co., Chicago, ILL.) on a shaker 
at 2,000rpm. When the appearance of material in the sieve 
changed from the black dust of spores to the lighter color 
of broken mycelium, sieving was stopped. 


Five A. alternata strains that had remained 
morphologically stable in culture for at least one year 
(Simmons 34-016, 34-039, 35-056, 35-193, 38-066) were 
maintained on 20% V-8 juice agar slants. On three occasions 
(9/29/86, 10/6/86, 10/28/86) spore material was harvested 
following the procedure of Kozak and Gallup, except that 20% 
V-8 juice agar was used instead of Sabouraud’s medium, and 
growth time before harvesting spores was 5 days instead of 
three weeks. Material scraped from the overlay material was 
not separated by sieving, as microscopic examination 
revealed the presence of very few mycelial fragments. 


Spores of the random air isolates (#1-4) were extracted 
at 1:10 w/v in 0.02% sodium azide in distilled water for 48 
hours at 4°C, centrifuged at 10,800g, filtered through a 
0.45um nitrocellulose filter, dialyzed for 48 hours against 
distilled water at 4°C, concentrated x10 and frozen. 


ay 


Simmons’ spore materials were extracted in the same manner, 
but each resulting spore extract was lyophilized. The 
possible differential effects of culture medium and 
lyophilizing vs concentrating and freezing on the allergen 
profile of extracts were assessed. Alternaria isolate A1005 
was grown on 20 V-8 juice agar and spores were harvested and 
extracted on two separate occasions, both times dividing the 
extract into two portions; one concentrated and frozen, the 
other, lyophilized. Extracts were compared using the 
following isoelectric focusing (IEF) and Western blot 
techniques, and allergens detected as described. 


For IEF, extracts of isolates 1-4 were thawed and used 
immediately, while lyophilized material was reconstituted to 
10 mg/ml with distilled water. Each extract was focused 
twice on 0.5mm polyacrylamide gels, pH 3-10, using Pharmacia 
Pharmalytes (Pharmacia Inc. Uppsala, Sweden). Gels were 
prefocused for 30 minutes at constant current of 3 milliamps 
using a Pharmacia Flat Bed Apparatus FBE-3000, Pharmacia 
3000/150 power supply and a Sargent water bath cooler (E.H. 
Sargent & Co., Chicago, ILL). Samples were loaded onto gel 
surface using Pharmacia applicator strips, 20ul per extract, 
and focused for 1.5-2 hours at 3 milliamps. 


Focused extracts were transferred to 0.45 
nitrocellulose paper (Schleicher and Schuell, Keene, NH) by 
the method of Towbin, et al. (1979) using a Bio-Rad Trans- 
Blot cell, 250/2.5 power supply (Bio-Rad Laboratories, 
Rockville Centre NY) and Sargent Cooling Bath. Blotting was 
carried out in 0.7% acetic acid at 100 volts for 3 hours. 


To detect allergens, a human serum pool was collected 
from 118 patients with 3 or 4+ prick test reactions to a 
commercial Alternaria extract. Alternaria allergens were 
detected on the immunoblots using methods described by 
Turner, et al. (1983) and Kroutil, et al (1987). Briefly, 
focused extracts on nitrocellulose were washed for 30 
minutes in buffer containing 0.1% gelatin to block remaining 
protein binding sites, and incubated overnight with the 
specific IgE-containing human serum pool. After washing, 
the blots were incubated for 4 hours with monoclonal mouse 
anti-human IgE, followed by 4 hours in alkaline phosphatase- 
labelled goat anti-mouse IgG (Atlantic Antibody, Charles 
River Co., Scarborough ME) and developed in fast blue and 
naphthol magnesium sulfate. 


58 


TABLE I. Alternaria isolates: 
1. Al1000 A. alternata-like, but not a perfect match to 
Simmons’ reference strains 


2. Al002 not A. alternata; 

3. Al004 A. alternata. 

4. Al1005 not A. alternata 

5. 34-016 A. alternata (E. G. Simmons) 
6. 34-039 A. alternata . 

7. 35-056 A. alternata . 

8. 35-193 A. alternata iY 

9. 38-066 A. alternata s 
RESULTS 


Initial comparisons between culture media and between 
concentrated, frozen vs lyophilized extract preparations of 
Alternaria isolate A1005 revealed essentially identical 
allergen profiles by the methods described above encouraging 
comparison among concentrated, frozen extracts and 
lyophilized preparations of other strains. 


A total of 32 different allergens were visualized in 
spore extracts from the four random isolates with all 
batches and strains combined (figure 1): 23 in the range pH 
3.00-6.57 and 9 in the 6.57-10.00 range. In the five stable 
A. alternata strains studied, 18 allergens were detected, 15 
between pH 3.00-6.57 and three between pH 6.57-10.00 (figure 
22 


pH A1000 AI1002 A1004 A1005 


8.15- 


6.55- 
5.85- 


3.75-- 


Figure 1. Comparison of IgE Immunoblots from four random Alternaria strains. 


a) 


Batch to batch variability is displayed in figure 3 in 
immunoblots from random isolate A1002 (the most allergen 
rich and least variable random isolate). Figure 4 displays 
immunoblots of 3 extraction batches each for the most 
allergen rich (35-193) and for the least variable (34-016) 
of the stable Simmons isolates. 


38-066 


Figure 2. Comparison of IgE Immunoblots from five stable A. alternata 


strains. 
pH 
8.15-- 
6.55- 
5.85- 
3.75- 


Figure 3. IgE Immunoblot variability in strain Al002. 


Considering all extractions, five allergens were 
demonstrated in all four random strains, but no single 
allergen was present in all batches. No single strain of 
these 4 was more allergenically stable than the others 
during the less than 18 month period in which extractions 
were made. With all spore batches combined, A1002, the most 
allergenically complete isolate, still lacked 19% (6/32) of 
qualitatively identified allergens. A1004 had the most 
incomplete allergen profile, missing 53% (17/32) of the 
demonstrated spore allergens. 


34-016 35-193 
wo ve) 
wo wo co wo wo (oe) 
Sc vit een Pehl) He = 
= o er a So N 
hi = = N —— — 


folgl tspeatd 


6.55 — 


5.85 — 


3. 7 Sam 


Figure 4. IgE Immunoblot variability in two stable Altermaria strains. 


The two strains that correctly represent A. alternata 
(A1000 and Al1004) differed from each other as much as from 
the two Alternaria strains misidentified as A. alternata. 


Five allergens were also present in all five of the 
stable strains, four of which corresponded to four of the 5 
allergens consistently present in the random isolates. 
Except for the 10/6/86 batch of strain 35-193, these 


61 


allergen bands were conserved in every batch of each strain. 
All batches of 34-016 were allergenically identical. Each 
of the other Simmons strains had at least one batch which 
was deficient in three or more allergens other than the five 
above. 


Of the five allergens conserved for each group of 
strains studied, four of these were probably present in all 
nine strains, and all of these had migrated in the pI range 
below 6.57. 


DISCUSSION 


These findings emphasize: 1) the danger that isolates 
of Alternaria may be incorrectly identified to form species 
by an observer inadequately trained in Alternaria taxonomy, 
2) that a single randomly isolated strain of A. alternata 
may show limited allergen expression, 3) that batch to batch 
differences for single strains may be extremely prominent 
and, 4) that the use of culturally stable strains can 
significantly increase the reliability of recovering of 
expected Alternaria allergens. 


A1002, an isolate clearly not A. alternata, was richest 
in allergens recognized by "Alternaria-sensitive" sera, and, 
it appears from figures 1 and 3 that the random isolates 
produced more allergen bands than the culturally stable 
strains. Preliminary work in our laboratory comparing spore 
and mycelial preparations indicates that these differences 
may be due to the higher percentage of mycelium present in 
the preparations from the random isolates. While every 
possible attempt was made to exclude mycelium from the 
preparations, many of the random isolate spores had begun 
germination either before or during harvest, and the random 
isolate cultures were more mycelial than the stable strains. 
It is worth noting that, although spores are the intuitively 
obvious unit of fungus exposure for most of the population, 
significant levels of hyphal fragments do occur in air and 
that allergen activity not shared by spores does appear to 
be present in myceliun. 


The diagnosis and treatment of clinical mold 
sensitivities has been difficult and, it seems, often better 
approached by environmental control rather than by 
immunologic approaches. Methods of allergen preparation 


62 


based on experience with field collected, easily recognized, 
and relatively stable plant materials (e.g. ragweed pollen) 
have not transferred well to fungus materials which can 
seldom be field collected, are identifiable only by a few 
specialists, and tend to be pleomorphic and biochemically 
variable in time (Burge, 1985). Until recently, potential 
interference resulting from these confounding factors has 
not been recognized (Helm, et al. 1987). The intrusion of 
non-medical mycologists into the field of allergy has at 
least served to alert those doing research on fungus 
allergens to possible dangers. It is apparent that before 
the true role of the fungi in human allergic disease can be 
accurately assessed, methods based on a solid grounding in 
fungus taxonomy, physiology and biochemistry will have to be 
developed. This will only be possible when medical 
researchers become willing to rely on mycological experts, 
and when students of mycology are willing to become involved 
in this important and exciting field. 


ACKNOWLEDGEMENTS 


This work was supported in part by Grant #AI-10181 from the 
National Institutes of Health. 


LITERATURE CITED 


Burge, H. A., Simmons, E. G., Muilenberg M., Hoyer, M., 
Gallup, J., Solomon, W. 1987. Intrinsic variability in 
airborne fungi: implications for allergen standardization. 
Advances in Aerobiology, Birkhauser Verlag Basel. 


Burge, H. A. 1985. Fungus Allergens. Clin Rev Allergy 3:319- 
329. 


Helm, R. M., Squellace, D. L., Aukrust, L., Borch, S. M., 
Baer, H., Bush, R. K., Lowenstein, H., Znamirowski, R., 
Nitchuk, W., Yunginger, J. W. 1987. Production of an 
international reference standard Alternaria extract. Int 
Archs Allergy Appl Immunol. 82:178-189. 


Hoffman, D. R., Kozak, P. P., Gillman, S. A., Cummins, L. 
H., Gallup, J. 1981. Isolation of spore specific allergens 
from Alternaria. Annal Allergy 46:310-16. 


Kroutil, L. A. & Bush, R. K. 1987. Detection of Alternaria 
allergens by Western blotting. J Allergy Clin Immunol 
80:170-6. 


63 


Lehrer, S. B., Aukrust, L., Salvaggio, J. E. 1983. 
Respiratory allergy induced by fungi. Clin Chest Med. 4:23. 


Norman, P. S. 1982. Quality control of allergen extracts. J 
Allergy Clin Immunol. 69:1. 


Solomon, W. R., Burge, H. A., Muilenberg, M. L. 1980. 
Allergenic properties of Alternaria spore, mycelial and 
"metabolic" extracts. J Allergy Clin Immunol. 65:229. 


Towbin, H., Staehelin, T., Gordon, J. 1979. Electrophoretic 
transfer of proteins form polyacrylamide gels to 
nitrocellulose sheets: procedure and some applications. 
Proc Natl Acad Sci USA 76:4350. 


RO ae he ih 


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Ny ae . 
aie me Sei . 


Vol. XXXIV, No. 1, pp. 65-70 January 20, 1989 


THAXTEROGASTER THIERSIT: A NEW SECOTIOID SPECIES 
FROM CALIFORNIA 


CORNELIA J. CALHOUN 


Life Sciences Division, SRI International 
Menlo Park, California 94025 


SUMMARY 


Thaxterogaster thiersii sp. nov is described from the 
coast of Northern California. It is compared to two similar 
species: T. porphyreum (Cunningham) Singer, which is 
known only from New Zealand, and JT. conicum (Hesler) 
Singer & Smith of the eastern U.S. 


INTRODUCTION 


This new secotioid species was discovered during a six year survey of the mac- 
roscopic fungi of Audubon Canyon Ranch, a 1000 acre nature preserve located in 
Marin County at the edge of the Bolinas Lagoon. It was collected in a successional 
mixed evergreen forest habitat at an altitude of approximately 100 feet, and it is 
distinguished from related taxa by significant macroscopic and microscopic fea- 
tures as well as by distribution. 


THAXTEROGASTER THIERSII sp. nov. FIGS: 132A. 2033 


Gasterocarpi Stipitato-pileati. Pileus 2.8-3.5 cm altus, subglobosus vel pyriformis, 
undulatus, margine incurvato et stipitem affixo, superficie viscida pallida, cristis partem 
violaceo-tinctis, sulcis pallido-griseo-aurantiaco-tinctis. Gleba loculata, brunneo- 
aurantiaca, loculis 0.2-1.5 mm longis. Stipito-columella nonnunquam percurrens, parte 
excurrenti brevi, apicem versus ramificans, laminas tramae radiantes formans, superficie 
sicco, violaceo. Contextus pallido-albus. Hyphae peridii 2-6 wm latae, tramae (4)6- 
10(12) pm latae. Basidia (32)40-50(65) x 10-15 um, tetrasterigmata, sterigmatibus 
3.5-6 x 2-3 um, symmetricis vel semifalcatis, apicalibus vel subapicalibus. 
Basidiosporae (12.5)14-16.5(18) x 10-12(13) pm, late ellipsodeae, subsymmetricae vel 
symmetricae, grosse verrucosae, cinnamomeo-brunneae, appendice hili brevi, hyalina. 
Fibulae nullae. Subhypogeus sub Querco agrifolia et Pseudotsuga menziesii. Holotypus: 
C. J. Calhoun 80-1475, 20 Jan 1980, Audubon Canyon Ranch, Marin Co., California 
(SFSU). 


Gasterocarps stipitate-pileate. Pileus 2.8-3.5 cm tall, 3-4 cm broad, subglobose 

to pear-shaped, undulate, margin curving in toward and remaining attached to 
_ stipe; surface viscid, glabrous, pallid, ridges partly violaceous-tinged, furrows with 
pale grayish-orange tints. Gleba loculate, chambers irregular, 0.2-1.5 mm in the 
longest dimension, colored brownish-orange (SC4, Kornerup & Wanscher, 1978), 
unchanging on exposure or injury, maturing first near stipe-columella apex, drying 


66 


cinnamon-brown. Stipe-columella well-developed, widest below (8-15 mm), taper- 
ing upwards, percurrent or not, tramifying toward apex into radiating tramal 
plates; context solid, white-pallid, unchanging on exposure, violaceous-tinged 
where damaged by insect larvae, stipe portion relatively short, narrowing some- 
what toward base; surface dry, glabrous, violaceous. Odor not distinctive, taste not 
recorded. 

Peridium about 250 um thick, composed of hyaline, loosely organized, 
filamentous hyphae, 2-6 um wide, lying + parallel to the surface within and below a 
gelatinous matrix; hyphae of the peridium trama up to 12 wm wide, some with 
gold-colored encrustations; hyphae of tramal plates (4)6-10(12) um wide, sub- 
parallel, smooth, hyaline. Oleiferous hyphae present in trama, common in mature 
gasterocarps, 2.5-6 4m wide, even or gnarled, hyaline to cinnamon-brown when 
revived in 3% KOH. Basidia (32)40-50(65) x 10-15 4m, mostly clavate, 4-spored, 
with sterigmata 3.5-6 x 2-3 wm, symmetric to semifalcate, apical to subapical. 
Pleurocystidia and cheilocystidia absent. Basidioles present, 20 um wide. 
Basidiospores (12.5)14-16.5(18) x 10-12(13) wm, broadly ellipsoid, axially sub- 
symmetric to symmetric, thick-walled, coarsely verrucose, cinnamon-brown, with a 
short, hyaline hilar appendage. Stipe hyphae subparallel, smooth, hyaline. Clamp 
connections absent in all tissues. 

Subhypogeous. Collected in mixed woods under Quercus agrifolia Neé and 
Pseudotsuga menziesii (Mirb.) Franco. 


TYPE: U.S.A. California. MARIN CO.: Audubon Canyon Ranch, 20 Jan 1980, 
C. J. Calhoun 1475 (HOLOTYPE: SFSU). 


Additional material examined: U.S.A. California. MARIN CO.: Audubon Canyon 
Ranch, C. J. Calhoun 80-1447, 80-1603 (SFSU). 


DISCUSSION 


Thaxterogaster thiersii belongs in section Aporpogaster Singer & Smith (Singer 
& Smith, 1963). It is distinguished macroscopically by the following combination 
of characters: a subglobose to pear-shaped pileus with a gently undulating surface 
and a margin that remains attached to the stipe-columella throughout develop- 
ment; a moderately viscid, pallid-colored, glabrous peridium with light violaceous 
and pale grayish-orange tints; a finely loculate, brownish-orange gleba which ma- 
tures first near the stipe-columella apex; a dry, light violaceous-colored, relatively 
short stipe proper. Distinctive microscopic characters include: the broadly ellip- 
soid, often axially subsymmetric, coarsely verrucose, cinnamon-brown-colored 
spores with thick exosporium; basidia up to 65 um long, with prominent, apically to 
subapically attached, straight to +curved sterigmata that are often 6 um long; the 
absence of clamp connections from all tissues. 


In combination, the macroscopic and microscopic characters of T. thiersii dis- 
tinguish it from the other species in section Aporpogaster, namely T. porphyreum 
and T. conicum. The following discussion includes observations published by 
Singer & Smith (1958). Thaxterogaster porphyreum (illustrated in Cunningham, 
1942) occurs in association with Nothofagus species in New Zealand and has a 


67 


Fig. 1. Carpophores of Thaxterogaster thiersii (HOLOTYPE), approx. x 1. 


globose to globose-depressed pileus with a viscid violaceous-colored peridium. 
Macroscopically, 7. porphyreum differs from T. thiersii in several other respects: 
the peridium is more highly pigmented; the stipe is comparatively long; the 
columella, which is often partly free from the gleba, always extends to the apex of 
the pileus; the context of the peridium, stipe, and columella is light lilac-colored. 
Some microscopic differences are: the spores of T. porphyreum are narrower 
[(7.5)9-11 um (Figs. 2A, 2B)], coarsely verruculose (Cunningham, 1924) with thin 
epispore, and are mostly axially symmetric; the basidia are 22-32 x (6)10-11 um; 
clamp connections are present in 7. porphyreum (Horak & Moser, 1965). Thax- 
terogaster conicum occurs from Ohio to Tennessee, U.S.A., and is found on the 
ground under Quercus sp. and Pinus echinata Mill. Gasterocarps of T. conicum are 
"apparently" always fully epigeous at maturity, and the species is considered to be 
the most agaricoid of the genus. Furthermore, they are generally much larger than 
those of T. thiersii, and typically have an elongate, conic pileus. Illustrations that 
show the distinctive stature of T. conicum appear in Hesler (1933) and Singer & 
Smith (1958). In contrast to T. thiersii, the peridium of T. conicum is innately 
white-silky-fibrillose, and colored grayish-pallid to grayish-ochraceous with a 
darker disk that becomes tinged violaceous on exposure; a white arachnoid veil is 
present; the pileus margin separates early along the entire length of the stipe- 


68 


Fig. 2. Spores from Thaxterogaster thiersii (HOLOTYPE), (A), photographed at 
Same magnification as B. (B). Thaxterogaster porphyreum, Cunningham 923 
(SYNTYPE). (C). Cross section of peridium from T. thiersii (HOLOTYPE), showing 
gelatinous matrix and simple septate hyphae. Scale bars = 10 pm. 


69 


Fig. 3. Scanning electron micrographs, basidiospores of Thaxterogaster thiersit 
(HOLOTYPE). In (A) note hilar appendages. Prominent sterigmata are seen at lower 


left in (B). Scale bars = 10 pm. 


70 


columella, exposing the gleba; the gleba is coarsely loculate (up to 10 mm long) 
with a lamellate development. The verrucose to warty spores of T. conicum reach 
20 um and are ellipsoid, while mature spores of T. thiersii rarely reach 18 um and 
are broadly ellipsoid. The basidia of T. conicum are 32-40 x 6-16.5 um, while the 
majority of basidia are 42-50 x 12-14 wm in T. thiersii. Hyphae in the trama of the 
peridium of T. conicum measure up to 22 um wide, but only reach 12 um in T. 
thiersii. 


ACKNOWLEDGEMENTS 


I would like to thank Barbara Thiers and Roy Halling for their very generous 
contributions to the preparation of this manuscript. I am also indebted to Rupert 
Barneby of the New York Botanical Garden for rendering the Latin description. 
Finally, I would like to express my sincere gratitude to and great respect for Dr. 
Harry Thiers whose outstanding scholarship and generosity are a continuing in- 
spiration. 


LITERATURE CITED 
Cunningham, G. H. 1924. A critical revision of the Australian and New Zealand 
species of the genus Secotium. Proc. Linn. Soc. New South Wales 49: 97-119. 1924. 
. 1942. The Gasteromycetes of Australia and New Zealand. J. McIndoe, Dunedin. 


Hesler, L. R. 1933. Pleurotus tremulus and Secotium conicum. J. Elisha Mitchell Sci. 
Soc. 49: 153-155. 


Horak, E. & M. Moser. 1965. Fungi Austroamericani XII, Studien zur Gattung Thax- 
terogaster Singer. Nova Hedwigia 10: 211-241. 


Kornerup, A., & J. H. Wanscher. 1978. Methuen Handbook of Colour. 3rd ed., Methuen 
Ltd., London. 


Singer, R. & A. H. Smith. 1958. Studies on secotiaceous fungi -I: A monograph of the 
genus Thaxterogaster. Brittonia 10: 201-216. 


. 1963. A revision of the genus Thaxterogaster. Madrofo 17: 22-26. 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 71-92 January 20, 1989 


STUDIES ON MARASMIUS FROM EASTERN 
NORTH AMERICA. II. NEW SPECIES 


Dennis E. Desjardin 
and 
Ronald H. Petersen 


Department of Botany 
University of Tennessee 
Knoxville, Tennessee 37996-1100 


ABSTRACT 


Four new taxa in Marasmius are described as new: M. 
ilicicola (sect. Marasmius), M. ciliatomarginatus, M. 
paludigenus and M. falcatipes (sect. Sicci). All taxa are 
illustrated and compared to phenetically similar taxa. 


Key Words. Marasmius ilicicola, Marasmius 
ciliatomarginatus, Marasmius paludigenus, Marasmius 
falcatipes. 


Although significant contributions have been made 
toward understanding of the genus Marasmius as it occurs 
in the northeastern United States (Morgan, 1905, 1906; 
Pennington, 1915; Gilliam, 1975a, 1975b, 1976; Halling, 
1983) and western United States (Desjardin, 1985, 1987a, 
1987b), no comprehensive taxonomic treatment of the genus 
from the southeastern United States is available. During 
the course of preparing a floristic monograph of Marasmius 
from southeastern North America, extensive herbarium 
studies and field work have uncovered several new taxa. 

Color terms and notations are from Kornerup and 
Wanscher (1978). In the sections on material examined, 
commonly used abbreviations are as follows: GSMNP, Great 
Smoky Mountains National Park; DED, D. E. Desjardin. 


72 


Where five or more specimens of each taxon were available 
for a comparison of intraspecific variation, we have 
followed the suggestion of Parmasto and Parmasto (1987: 
111) and calculated the 90Z%-expected tolerance limits of 
the mean spore size at 90% probability level [TL90(90Z%)]. 
Other spore factors determined include: x, the arithmetic 
means of the spore length (L) and spore width (W) in a 
sample (specimen); x, the mean of means where more than 
one specimen is available; s, standard deviation of a 
sample (indicated as sL X sW, and sQ), or standard 
deviation of the mean values where more than one specimen 
is available; E, the quotient of L and W in any one spore 
(indicated as a range of variation in n spores measured); 
Q, the mean of E-values in a sample; Q, the mean of Q- 
values where more than one specimen is available. 


MARASMIUS ILICICOLA Desjardin sp. nov. Figs Silane 


Pileus 2-4 mm latus, convexus, sulcatus, umbilicatus, 
subvelutinus, disco atrobrunneus, margine pallidior. 
Lamellae adnatae, distantes, latae, collariatae, pallidae. 
Stipes 10-22 mm longus, 0.1-0.2 mm crassus, filiformus, 
glaber, insititius, atrobrunneus; rhizomorphae atrae. 

Odor et sapor nullus. Basidiosporae 7.5-9.5 X 3.5-4.8 um, 
amygdaliformes, laeves, hyalinae, inamyloideae. Basidia 
tetraspora. Pleurocystidia nulla. Cheilocystidia 
elementis pileipellis similia. Pileipellis hymeniformis 
ex elementis M. rotali similibus; diverticuli pallido- 
brunnei. Trama ex hyphis inamyloideis, fibulatis. In 
foliis Ilicis opacae. Holotypus: Mississippi, De Soto 
National Forest, Perry Co., Black Creek Wilderness Area, 
16 Jul 1987, D...E. Desjardin, no. 4355) (no: 4762570 ;enean 


Basidiomata (Fig. 1) marcescent, reviving. Pileus 2- 
4 mm broad, convex, sulcate, centrally depressed, with or 
without a broad, low papilla; surface dull, dry, opaque, 
subvelutinous; central depression or papilla dark brown 
(6-7F4-8) throughout maturation; margin evenly light brown 
(6D4-5) when young, remaining so in age or fading to 
greyish orange (5B4), lacking a whitish zone surrounding 
the dark central dot; context thin, concolorous with 
adjacent cuticle. Lamellae distant (9-11 reach the 
collarium), broad, adnate to a complete, free collariun, 
pale orange white (5A2) when young, becoming pale brownish 
orange (5B3-5C4) in age, collarium and lamellar edges 
concolorous with lamellar faces or darkening slightly when 


74 


dried, lamellulae absent. Stipe 10-22 X 0.1-0.2 mn, 
terete, equal, filiform, hollow, shining, glabrous, 
insititious; apex concolorous with lamellae, base dark 
brown (6F4-8), darkening overall in age. Rhizomorphs 
scattered, wiry, black, basidiomata not arising from the 
rhizomorphs. Odor and taste not distinctive. 

Basidiospores (Fig. 2) 7.5-9.5 X 3.5-4.8 um [x = 8. 
X 4.3) ‘um, si= 0.435% 082630 Ei= >). 8-2 .1370 = 3) soe D: 
n = 30], ellipsoid or amygdaliform, hyaline, inamyloid. 
Basidia (Fig. 3) 22-26 X 5.5-6.5 um, cylindric or 
subclavate, 4-spored. Basidioles (Fig. 3) fusoid. 
Pleurocystidia absent. Cheilocystidia (Fig. 4) numerous, 
similar to pileipellis elements, 16-20 X 8.5-12 um, 
hyaline overall or apically pale ochraceous, inamyloid. 
Pileipellis a hymeniform layer of Rotalis-type elements 
(Fig. 5), 17-25.5 X 9-16(-20) um, broadly clavate or 
sphaeropedunculate, apically pale brown and thick-walled, 
basally hyaline and thin-walled, diverticula -1.6 X -1.2 
um, numerous, apical and subapical, knob-like, solid, 
pigmented portions inamyloid or weakly dextrinoid, 
unpigmented portions inamyloid; some scattered elements 
slightly thicker-walled and more deeply pigmented than 
average giving the pileipellis a weakly mottled appearance 
under low magnification. Pileal and lamellar tramal 
hyphae interwoven, 2.5-8 um diam, hyaline, smooth or 
weakly granular-incrusted, inamyloid. Stipe cortical 
hyphae parallel, 2.5-6.5 um diam, hyaline to pale 
ochraceous at the stipe apex, deep olivaceous brown in KOH 
at the stipe base, with dextrinoid walls up to 1 um thick. 
Stipe medullary hyphae parallel, 2-8 um diam, hyaline, 
inamyloid, thin-walled. Caulocystidia absent. Clamp 
connections common on tramal hyphae and hymenial elements. 

HABIT, HABITAT AND DISTRIBUTION. Scattered on 
senescent leaves of Ilex opaca Ait. in mixed bottomland 
hardwoods with scattered pines. Mississippi. July. 
Locally abundant. 

MATERTAL EXAMINED. UNITED STATES. MISSISSIPPI: De 
Soto National Forest, Perry Co., Black Creek Wilderness 
Area, \16.vii.87. DED sno. 4355 (no. 4/7625. 6 LENNee 
Holotype) . 


3 
1: 


> 


OBSERVATIONS. Marasmius ilicicola belongs in sect. 
Marasmius, subsect. Pararotulae (Sing.) Sing. because of 
the presence of collariate lamellae, insititious stipe and 
Rotalis-type pileipellis elements. It is microscopically 
Similar to several neotropical taxa, viz. Marasmius 
scototephrodes Sing. from Mexico, and M. tetrachrous 


75 


Sing. from Bolivia. Marasmius scototephrodes differs in 
having an ash grey pileus with a whitish zone surrounding 
a grey central dot, dark greyish-marginate lamellae and 
fruiting habit on sticks and leaves of various 
dicotyledonous plants but not on Ilex [Holotype: Singer 
M8299 (F!)]. Marasmius tetrachrous differs by forming 
smaller (1.5-2 mm diam), bullet-shaped, ferruginous pilei 
with a pallid zone surrounding a dark central dot, in 
having shorter and broader basidia and in lacking 
rhizomorphs (fide Singer, 1976). In eastern North 
America, M. ilicicola is superficially similar to M. 
capillaris Morg., in that both species form basidiomes 
with brownish, sulcate pilei, collariate lamellae and dark 
filiform stipes accompanied by rhizomorphs. Marasmius 
capillaris differs, however, in forming pilei with a buff 
or pale yellowish zone surrounding the central depression, 
in having more numerous and more closely spaced lamellae 
and by growing on leaves of Quercus species or rarely on 
coniferous needles [Lectotype: Morgan, Oct. 1890 (ISC!)]. 
Of the nearly 100 herbarium collections of M. capillaris 
we have studied, no basidiomes were observed growing from 
leaves of Ilex. Field studies also support these data. In 
areas where Ilex opaca and Quercus spp. are sympatric, 
extensive searches for M. capillaris on Ilex and M. 
ilicicola on substrates other than Ilex have proved 
unsuccessful. See Gilliam (1976) for a description of M. 
capillaris. 

If the collarium is overlooked, M. ilicicola might be 
confused with another macroscopically similar Ilex leaf- 
degrading taxon, viz. M. ilicis Singer. The latter 
differs microscopically in having ventricose or lageniform 
pleurocystidia, thick-walled dermatocystidia similar in 
shape to the pleurocystidia and interspersed among 
Rotalis-type pileipellis elements, and in having 
diverticulate stipe cortical hyphae [Isotype: Singer B88 
(MICH! )]. See Singer (1953) for a description of M. 
er LS « 

Although Ilex opaca (American Holly) is common 
throughout the southeastern United States and ranges 
northward into coastal Massachusetts (Elias, 1980), M. 
ilicicola is known at present only from coastal 
Mississippi. It is presumed that the high summer 
temperatures and humidity plus mild winter temperatures 
and abundant rainfall characteristic of the gulf coast 
region are important determining factors in the 
Hieeripucion of MS ilicicola. 


76 


MARASMIUS CILIATOMARGINATUS Desjardin sp. nov. Figs. 6-11. 


Pileus 5-20 mm latus, campanulatus vel plano- 
convexus, ruguloso-striatus, subvelutinus, badius vel 
ferrugineus. Lamellae adnatae, subdistantes, angustae, 
alboluteae, ferrugineus-marginatae. Stipes 18-45 mm 
longus, 0.5-1 mm crassus, teres, aequalis, pubescens, non 
insititius, apice bubalinus, base brunneus, aetate atrans. 
Odor et sapor nullus. Basidiosporae 13.5-18 X 3.2-4.7 um, 
clavatae, laeves, hyalinae, inamyloideae. Basidia 
tetraspora. Pleurocystidia nulla. Cheilocystidia 
cylindrica vel flexuosa, gleocystidiodea, contentibus 
cinnabarinis, rariter elementis pileipellis similia. 
Pileipellis hymeniformis ex elementis M. sicco similibus; 
diverticuli ferruginei. Trama ex hyphis dextrinoideis, 
fibulatis. Caulocystidia cheilocystidiis gleocystoideis 
similia. Ad folia dejecta dicotyledonum. Holotypus: 
North Carolina, Macon Co., Highlands, Horsecove, 10 Aug 
1987, D. E. Desjardin no. 4414 (no. 47626, TENN). 


Basidiomata (Fig. 6) marcescent, reviving. Pileus 5- 
20 mm broad, conic or campanulate when young, expanding in 
age to broadly campanulate or plano-convex, often with a 
low umbo and upturned margin; surface dull, dry, opaque, 
subvelutinous; disc rugulose, margin rugulose-striate; 
context thin, buff; color reddish brown (8D6-8), brown 
(7E7-8) or deep orange brown (7D8) overall when young, 
disc remaining so in age or fading to brownish orange 
(7C7-8), margin soon becoming brownish orange and fading 
in age to light brownish orange (6C5-7) or rarely when old 
and wet becoming light orange (5A3-4). Lamellae adnate, 
subdistant to nearly close (20-25 reach the stipe), narrow 
(<1.5 mm), yellowish white or cream (4A2-3), seldom forked 
near the margin, interlamellar spaces sometimes venose and 
assuming pileus tints at maturity; edges granular- 
-erystalline, pale-concolorous with the pileus; lamellulae 
in 0-2 series. Stipe 18-45 X 0.5-1 mm, terete, equal or 
seldom with a small subbulbose base, pruinose to pubescent 
overall, arising from a small ring of buff or cream- 
colored mycelium; when young, upper half yellowish white 
(4A2) to dingy buff, lower half brown (6E5-6), 
hysterochroic, in age upper few mm pallid, base brown (6- 
7E4-6), reddish brown (8E5-6) or dark brown (7F5-6). Odor 
and taste not distinctive. 

Basidiospores (Fig. 7) 13.5-18 X 3.2-4.7 um [x = 15.7 

X'4 um, s°=)0595 X)0.2630E = 3-4.6; 9 = 3.9)..s0 =10pgGee 
= 30/3 collections], clavate or fusiform-elliptical, often 


77 


Figs. 6-9. Features of Marasmius ciliatomarginatus 
[Desjardin 4414, Holotype]. 6. Basidiomata Gale 

7. Basidiospores. 8. Basidia and basidioles. 

9a. Gloeocystidioid cheilocystidia. DeoLccus. Lye 
cheilocystidia. Scale bar = 10 um. 


78 


Figs. 10-11. Features of Marasmius ciliatomarginatus 
[Desjardin 4414, Holotype]. 10. Pileipellis elements. 
lla. Caulocystidia from stipe apex. 11b. Caulocystidia 
from stipe base. Scale bar = 10 um. 


79 


curved in profile, hyaline, inamyloid. Basidia (Fig. 8) 
22-30 X 6.5-8.5 wm, subcylindric or clavate, 4-spored. 
Basidioles (Fig. 8) clavate, ventricose or fusoid. 
Pleurocystidia not differentiated. Cheilocystidia of two 
types: 1) numerous gloeocystidioid elements (Fig. 9a), 38- 
56 X 3-5 um, cylindric, flexuous or strangulate, obtuse, 
thin-walled, with or without tawny to reddish orange 
globular cytoplasmic contents typically congregated 
nearest the apices of the cells, or with reddish orange 
globular masses adherent to the external cellular 
surfaces, walls inamyloid; 2) rare (or absent), scattered 
Siccus-type elements (Fig. 9b) similar to pileipellis 
elements, with hyaline or pale orange diverticula. 
Pileipellis a hymeniform layer of Siccus-type elements 
(Fig. 10), with rare, inconspicuous, clavate, thin- 
walled, non-diverticulate cells interspersed; Siccus-type 
elements 8-16 X 3.2-8 um, cylindric, clavate or irregular 
in outline, sometimes lobed, thin- to thick-walled, 
hyaline or tawny-colored; diverticula 1.5-8 X 0.8-2 um, 
apical, irregularly conic or cylindric, subnodulose or 
not, thick-walled or solid, ochraceous or reddish brown, 
pigmented portions weakly dextrinoid; some elements with 
tawny, globular contents; some elements thicker-walled and 
more deeply pigmented than average giving the pileipellis 
a mottled appearance under low magnification. Pileal and 
lamellar tramal hyphae interwoven, 1.5-9 um diam, 
cylindric or inflated, branched, hyaline, non-incrusted, 
inamyloid or weakly dextrinoid, thin-walled. Stipe 
cortical hyphae parallel, 2.5-6 um diam, dextrinoid, 
hyaline or pale ochraceous and thin-walled apically, 
ochraceous or brown and thick-walled below. Stipe 
medullary hyphae parallel or subparallel, 2.5-11 um dian, 
hyaline or pale yellowish, inamyloid or weakly dextrinoid. 
Highly refractive oleiferous hyphae common. Stipe vesture 
of abundant caulocystidia: at stipe apex (Fig. lla) 13-32 
X 3-5 um, irregularly cylindric or strangulate-contorted, 
rarely lobed, obtuse, hyaline, inamyloid, thin-walled, 
with or without reddish orange globular contents or 
adherent exudates; at stipe base (Fig. 11b) -40 X 4-8 un, 
cylindric to ventricose or acuminate, pale ochraceous, 
inamyloid, walls -1.2 um thick. Clamp connections common 
throughout basidiomata. 

HABIT, HABITAT AND DISTRIBUTION. Scattered to 
gregarious on senescent hardwood leaves or stems (rarely 
on Rubus stems or hickory nuts) in mixed woods containing 
Liriodendron, Quercus, Alnus, Cornus, Carya and Ilex with 
scattered Pinus and Tsuga. Aug. - Sept. Uncommon. NC, TN. 


MATERIAL EXAMINED. UNITED STATES. NORTH CAROLINA: 
Macon Co.: Highlands, Horsecove, 10.viii.87, DED no. 4414 
(no. 47626, TENN - Holotype). TENNESSEE: Blount Co.: 
GSMNP, Cades Cove, 31.viii.86, DED no. 4078 (no. 47628, 
TENN); Knox Co.: Knoxville, 7.ix.86, DED no. 4154 (no. 
47627, TENN). 


OBSERVATIONS. Features which in combination are 
diagnostic for M. ciliatomarginatus include: a) reddish 
brown or brownish orange, rugulose-striate pileus; 

b) subdistant, narrow, cream-colored, orange-marginate 
lamellae; c) pubescent, apically pallid and basally 
brownish, non-insititious stipe; d) clavate spores 
averaging 15.6 X 4 um; e) absence of pleurocystidia; 

f£) flexuous cheilocystidia with reddish orange contents 

or adherent exudates; and g) cylindric-contorted, non- 
diverticulate caulocystidia. Collectively, this diagnosis 
suggests placement in sect. Sicci, subsect. Siccini, ser. 
Actinopodes Singer (1976). The most distinctive feature 
of this species is the abundant gloeocystidioid 
cheilocystidia and relative absence of Siccus-type 
cheilocystidia, resulting in a lamellar edge morphology 
unique in section Sicci. Consequently, M. 
ciliatomarginatus is not likely to be confused with any 
other members of the section. There are, however, several 
North American taxa that are phenetically similar in many 
other respects, viz. M. sullivantii Mont. (widespread in 
eastern North America), and M. corrugatus var. aurantiacus 
(Murr.) Sing. (known from Florida and the neotropics). 

Marasmius sullivantii differs in having much smaller 
spores*, abundant pleurocystidia and broom-cell-type 
caulocystidia [Representative material: DED no. 4342 (no. 
47648, TENN)]. Marasmius corrugatus var. aurantiacus 
differs in having a glabrescent stipe with scattered 
broom-cell-type caulocystidia, and smaller spores* 
[Holotype of Gymnopus aurantiacus: Murrill, F17904 
(FLAS!)]. See Gilliam (1976) and Singer (1976), 
respectively, for descriptions of the latter two taxa. 


*Spores of M. sullivantii: 6.4-8.8 X 3.2-4 um, x = 7.6 X 
3.6 um,. su 04264X%- 00123 0B =41.8-276;'01=0122175)sOe=ome ee 
TL90( 90%) = 7.1-8.1,/X 3.4-3.9 um,.Q)= 2-2.2; n = 30/8 
specimens. Spores of Gymnopus aurantiacus: 7.2-9 X 3.4- 
4.2) ums X= 2 Bale xwavOr umanso=— 0. 4s Xv 01005 ee” Oe 


Ov=, 251 5) sOse0, 2s) nye 30... 


81 


MARASMIUS PALUDIGENUS Desjardin sp. nov. Figs. l2-16. 


Misapplied epithet: Marasmius glabellus Peck sensu 
Ellis, in de Thumen, Mycotheca Universalis Exsiccati Cent. 
6, no. 505, 1876. Marasmius glabellus Peck sensu Ellis, 
in North American Fungi Exsiccati Ser. I, Cent. 10, no. 
910, 1883. 


Pileus 7-18 mm latus, convexus, sulcatus, 
subvelutinus, disco ochraceus, margine griseolo-aurantius 
vel alutaceus. Lamellae adnexae, remotae, latae, 
griseolo-aurantiae vel fusciochraceae. Stipes 20-65 mm 
longus, 0.5-1 mm crassus, teres, aequalis, glaber, non 
insititius, apice bubalinus, base brunneus, aetate atrans. 
Odor spermaticus. Basidiosporae 16-24 XK 4.4-6 um, 
clavatae, laeves, hyalinae, inamyloideae. Basidia 
tetraspora. Pleurocystidia nulla. Cheilocystidia 
elementis pileipellis similia. Pileipellis hymeniformis 
ex elementis latis, M. sicco similibus, etiam ex elementis 
laevibus, vesiculosis. Trama ex hyphis dextrinoideis, 
fibulatis. Ad folia dejecta in palustria. Holotypus: 
New Jersey, Salem Co., Camp Edge Boy Scout Camp, 18 Aug. 
Pos /eene. Halling (no, 47622,. TENN). 


Basidiomata (Fig. 12) marcescent, reviving. Pileus 
7-18 mm broad, obtusely conic or parabolic when young, 
expanding and becoming convex in age; disc rugulose, 
margin sulcate or plicate; surface dull, dry, opaque, 
subvelutinous; context thin, buff; color yellowish brown 
(5D5) overall when young, disc remaining so in age or 
fading slightly, margin fading to pale brownish orange 
(5C4-5) or greyish orange (5B3-4), eventually becoming tan 
or cream-colored (4A3). Lamellae adnexed or nearly free, 
remote [11-13 (rarely 16) reach the stipe], broad (2-4 
mm), greyish orange (5B4-5) when young, soon becoming 
brownish orange (5C4-5), darkening in age to dark 
yellowish brown (5E-F5-6), neither forked nor intervenose; 
edges concolorous with the faces; lamellulae typically 
absent. Stipe 20-65 XK 0.5-1 mm, terete, equal, shiny, 
glabrous, hollow, arising from a small pad of yellowish 
mycelium; white to buff above when young, brown below 
(6F4-8), darkening overall in age. Odor of crushed pilei 
spermatic. Taste not observed. Ne 

Basidiospores (Fig. 13) 16-24 X 4.4-6 um [x = 19.4 X 
Seem, s1= 0245 X°OsL53 B= 9371-4553 = Ou sO'="0.1 38 
TL90(90Z) 18.5-20.2 X 4.8-5.4 um, Q = 3.5-4; n = 30/9 
specimens], clavate or fusiform-elliptical, sometimes 


Il 1 


82 


Bigs 2 16, 
U2 eBasidiomatayCx1)s( Fiynne {Snel 
14. Basidium and basidioles. 15. 
16. Pileipellis elements. [Figs. 13-16: 


18.viii.84, Holotype]. Scale bar = 10 um. 


Features of Marasmius paludigenus. 
Basidiospores. 
Cheilocystidia. 
Halling, 


83 


curved in profile, hyaline, inamyloid; readily collapsing 
and reviving poorly. Basidia (Fig. 14) 25-38 X 7-8.8 um, 
clavate, 4-spored. Basidioles (Fig. 14) clavate or 
ventricose. Pleurocystidia absent. Cheilocystidia (Fig. 
15) abundant, 7.5-18(-25) X 4-8.8 um, similar to the 
diverticulate pileipellis elements, hyaline. Pileipellis a 
hymeniform layer of versiform cells (Fig. 16), 6.5-25.5 X 
4.5-12(-16) um, irregularly cylindric, clavate, 

obclavate, pyriform or vesiculose, many lobed, inamyloid, 
with or entirely without apical diverticula; immature 
pilei composed mainly of diverticulate elements, basal 
portion of cells hyaline, thin-walled, diverticula 1-4.5 
Meus 4.) im, hyaline or ochraceous; thin-" or thick- 
walled; in mature pilei diverticulate elements abundant on 
the pileal disc, less frequent or rare elsewhere, pileus 
margin regions composed mainly of smooth, often lobed, 
non-diverticulate elements interspersed among tramal 
hyphae. Pileal and lamellar tramal hyphae interwoven, 3- 
OC=12) um diam, often inflated, branched, dextrinoid, 
hyaline to brownish yellow; pigment soluble in 3% KOH. 
Stipe cortical hyphae parallel, 2.5-6.5 um diam, hyaline, 
dark ochraceous or brown, dextrinoid, walls -1l um thick. 
Stipe medullary hyphae subparallel, 2.5-9 um diam, hyaline 
or pale yellowish, weakly dextrinoid, thin-walled. 
Refractive oleiferous hyphae common. Caulocystidia absent. 
Clamp connections common throughout basidiomata. 

HABIT, HABITAT AND DISTRIBUTION. Scattered on 
senescent leaves or among mosses in sandy and swampy areas 
with Quercus, Ilex and Pinus. Aug. - Oct. Locally 
abundant. Dit, NJ; NY, VA. 

MATERIAL EXAMINED. UNITED STATES. DELAWARE: no 
location data, 19.viii.84, Flynn, Vilgalys & Cotter, US 
900515 (BPI). NEW JERSEY: Salem Co.: Camp Edge Boy Scout 
Campemie.viii.o4, R.°E. Halling s.n. (no. 47622), “TENN - 
Holotype). Gloucester Co.: Newfield, Ellis s.n. (BPI); 
Newfield, Ellis 22.vii.1875 (BPI); Newfield, Ellis, 
viii.1875 in de Thtimen, Myco. Univ. no. 505 (as Marasmius 
glabellus) (BPI, FH, presumably other distributions); 
Newfield, Ellis, viii.1882, in N. Amer. Fungi no. 910 (as 
M. glabelius) (BPI, FH, MICH, NY, NYS, PENN at PH, PH, 
presumably other distributions). NEW YORK: Suffolk Co.: 
Robert Moses State Park, 28.x.84, B. M. Thiers & R. E. 
Halling no. 3883 (no. 47623, TENN). VIRGINIA: 

Northampton Co.: swamp near Norfolk, 20.viii.84, T. Flynn 
mown ls (no. 4/624, TENN). 


OBSERVATIONS. Marasmius paludigenus is characterized 
by: 1) yellowish brown or tan, sulcate pileus; 2) remote, 
broad, brownish orange or dark yellowish brown lamellae; 
3) glabrous, brown, non-insititious stipe; and 4) growth 
in swampy environments. In addition, long spores (mean 
length 19.4 um), absence of pleurocystidia and 
caulocystidia, and pileipellis morphology are distinctive. 
This species was collected by J. B. Ellis "among moss in 
swamps" in Newfield, New Jersey on several occasions 
between 1875 and 1882, although Ellis determined his 
specimens as Marasmius glabellus Peck. An Ellis 
collection was distributed by de Thumen as no. 505 in the 
Mycotheca Universalis Exsiccati (issued 1876). Several 
years later, Ellis distributed additional material 
(determined as M. glabellus) as no. 910 in his North 
American Fungi Exsiccati (issued 1883). All duplicates 
studied of the two exsiccata numbers mentioned above 
represent M. paludigenus. Marasmius glabellus is similar 
in stature, but differs in showing a darker pileus (i.e., 
usually retaining deep brownish orange or yellowish brown 
pigmentation in age and only rarely becoming pallid 
overall) and much paler lamellae which never develop dark 
yellowish brown tints as in M. paludigenus. Moreover, M. 
glabellus has numerous pleurocystidia, pileipelli composed 
entirely of Siccus-type diverticulate elements and smaller 
spores [x = 9 X.4.7 um,,s =0.94 X'0.36; E = 16-2727 
1.9, sQ = 0.13; n = 30. Holotype: Worcester & Croghan, 
Peck, July & August (NYS!)]. Another southeastern 
representative of sect. Sicci with sulcate pileus, 
distant broad lamellae and glabrous brown stipe is 
Marasmius similis Berk. & Curt. The latter differs from 
M. paludigenus in having a whitish pileus, pileipellis 
elements and cheilocystidia with densely nodulose 
diverticula, and in forming smaller spores [x = 12.3 X 3.9 
um, s = 0.91% 0,293.8 = 2.8-3.6;,Q = 3.2, sO’=) Of oe 
30. Isotype: South Carolina, Society Hill, Curtis 1319 
(FH! )]. 

An interesting feature of pileipellis morphology in 
M. paludigenus is the abundance of non-diverticulate 
elements in mature pilei and their relative absence in 
immature pilei. Indeed, in some mature basidiomata it is 
difficult to locate broom-cell-type elements in the pileal 
margin regions, whereas the entire pileipellis of immature 
basidiomata is formed almost exclusively of broom-cell- 
type elements. One might be easily misled as to the 
sectional disposition of this taxon if only a single 
tangential section of the pileus were examined. Were a 


85 


hymeniform pileipellis of smooth cells present, this in 
combination with other basidiomata characters of M. 
paludigenus, would suggest placement in sect. Globulares 
Kiihner. However, because of the abundant Siccus-type 
pileipellis elements present in young pilei and in the 
disc region of mature pilei, this species is accepted here 
Inesece si sicci’. 

Marasmius paludigenus has many characteristics in 
common with M. decipiens Halling, Desjardin & Tish. For 
example, both taxa are similar in basidiomata stature, 
pileus shape and coloration, lamellar width and spacing, 
stipe coloration, spore shape, hyphal tissue amyloidity 
and absence of pleurocystidia. And, although the 
cheilocystidia and pileipellis elements of these two 
species are similar in shape, diverticula are altogether 
lacking from these structures in M. decipiens, an 
important diagnostic criterion dictating placement of M. 
decipiens in sect. Globulares. Marasmius decipiens 
differs also in having paler lamellae, pubescent to 
strigose stipe base and longer spores [x = 22.9 X 4.6 um, 
Prmooe xe Onl 2c = 4-03) Ol=)5,2s0) =.0 1243) TLI0(90Z): = 
21.5-24.4 X 4.3-4.9 um, Q = 4.5-5.53; n = 25/6 specimens. 
Holotype: Tish 1602-F (NY!)]. See Halling et al. (1985) 
for a description of M. decipiens. 


MARASMIUS FALCATIPES Desjardin sp. nov. iS eh (ae oe 


Pileus 1-7(-11) mm latus, e convexo plano-convexus, 
ruguloso-striatus, subvelutinus; primo bubalinus, 
griseolo-brunneus, brunneo-aurantius vel brunneus, in 
aetate pallidior. Lamellae adnatae, confertae vel 
subdistantes, angustae, pallidae. Stipes 1-2.5 mm longus, 
<0.2 mm crassus, eccentricus vel centralis, teres, 
aequalis, falcatus, pruinosus, non insititius, bubalinus. 
Odor et sapor nullus. Basidiosporae (7.5-)8-10.5(-12) X 
3.5-5.2 um, ellipsoideae vel amygdaliformes, laeves, 
hyalinae, inamyloideae. Basidia tetraspora. 
Pleurocystidia rara vel abunda, refractiva, cylindrica, 
obtusata. Cheilocystidia elementis pileipellis similia. 
Pileipellis hymeniformis ex elementis M. sicco similibus; 
diverticuli hyalini et ferruginei. Trama ex hyphis 
dextrinoideis, fibulatis. Caulocystidia diverticulata. 
In foliis et ramis emortuis dicotyledonum vel 
monocotyledonum. Holotypus: North Carolina, Macon Co., 
Highlands, Horsecove, 10 Aug. 1987, D. E. Desjardin no. 
4415 (no. 47629, TENN). 


86 


Basidiomata (Fig. 17) marcescent, reviving. Pileus 
1-7(-11) mm broad, campanulate or convex, expanding to 
plano-convex or plane, rarely umbonate, often undulate in 
age, rugulose-striate; surface dull, dry, opaque, 
subvelutinous; context thin, whitish; color exceedingly 
variable, when young buff, greyish cream (4C2), grey 
(4E2), pale brownish grey (5E2-3), brownish orange (5G4), 
dark yellowish brown (5F4) or brown (6E4-6), the more 
deeply pigmented pilei fading in age to light yellowish 
brown (5D-E4-5), greyish yellow (5D3), pale brownish 
orange (5C3) or yellowish grey (4B3), but drying darker. 
Lamellae adnate, close or subdistant (7-14 reach the 
stipe), narrow or moderately broad (-1.5 mm), rarely 
forked or intervenose near the margin, white, buff or pale 
yellowish white (4A2) at first, remaining so in age or 
becoming pale yellowish grey (4B3) or orange white (5A2), 
not marginate; lamellulae in 1-2 series. Stipe 1-2.5 X 
<0.2 mm, typically eccentric, but often central or rarely 
nearly lateral, terete, equal, curved or geniculate, 
pruinose overall, arising from a small, white mycelial 
pad, white or buff overall when young, remaining so in age 
or the base darkening to cream (4A3), pale yellowish grey 
(4B3) or pale brownish orange (5C3). Odor and taste not 
distinctive. 

Basidiospores (Fig. 18) (7.5-)8-10.5(-12) X 3.5-5.2 
um [x = 9.4 X 3.9 um, s = 0.45 X 0.19; E = 1.9-2.9; Q = 
2.4, sQ = 0.12; TL90(90Z) = 8.5-10.3 X' 3.5-4.3 um, Q = 
2.2-2.73; n = 25/7 specimens], ellipsoid or amygdaliform, 
often with a slight suprahilar depression and abaxial 
bulge, with a prominent hilar appendix, hyaline, 
inamyloid. Basidia (Fig. 19) 16-22.5 X 5-6.5(-7.5) um, 
clavate, 4-spored. Basidioles (Fig. 19) broadly clavate 
or ventricose-fusoid. Pleurocystidia (Fig. 20) rare to 
numerous, 30-45 X 4.8-8 um, irregularly cylindric, obtuse, 
refractive, hyaline, inamyloid, arising from deep in the 
subhymenium and projecting up to 11 um beyond basidioles; 
base often curved. Cheilocystidia (Fig. 21) numerous, 11- 
17(-20) X 4.8-8(-9) um, diverticulate, similar to 
pileipellis elements but typically hyaline overall, only 
rarely with pale ferruginous diverticula; lamellar edge 
entirely sterile or with scattered basidia, basidioles and 
rare pleurocystidia-type elements. Pileipellis a 
hymeniform layer of Siccus-type elements (Fig. 22), 8.5-16 
X 4.5-8.5(-9.5) um, cylindric, clavate or irregular in 
outline, often lobed, diverticulate; basal portion of 
cells hyaline, thin-walled, inamyloid; diverticula 1-7(-9) 


Figs. 17-21. Features of Marasmius falcatipes [Desjardin 
WGl5, Holotype]. 1/7. “Basidiomata'i\(x3). 18. Basidio- 
spores. 19. Basidia and basidioles. 20. Pleurocystidia. 
21. Cheilocystidia. Scale bar = 10 um. 


Bigs) (22-2382 
4415, Holotype]. 22. Pileipellis elements. 

23. Elements of stipe vesture (thick-walled elements 
from stipe base). Scale’ bar’ = 107 tim. 


Features of Marasmius falcatipes [Desjardin 


89 


X 0.5-2 um, irregularly cylindric or contorted, rarely 
branched, obtuse, hyaline to pale yellowish and thin- 
walled or ochraceous to ferruginous and thick-walled, 
pigmented areas weakly dextrinoid. Pileal and lamellar 
tramal hyphae interwoven, 2.5-8(-10) um diam, inflated, 
non-gelatinous, hyaline, thin-walled, dextrinoid. Stipe 
cortical and medullary hyphae similar, parallel, 2.5-10 um 
diam, hyaline or pale yellowish, strongly dextrinoid, 
walls -2 um thick; hyphae from stipe base strongly 
sclerified. Stipe vesture (Fig. 23) of numerous 
diverticulate inflations projecting from the outermost 
layer of cortical hyphae, and with scattered elements 
similar to those of the pileipellis. Clamp connections 
common throughout basidiomata. 

HABIT, HABITAT AND DISTRIBUTION. Scattered to 
gregarious on senescent leaves or stems of various 
hardwoods, herbaceous vines or grasses in mixed deciduous 
woodlands. July - Sept. Uncommon. NC, SC, TN, VA. 

MATERIAL EXAMINED. UNITED STATES. NORTH CAROLINA: 
Buncombe Co.: Lake Powatah, near Asheville, 5.ix.87, DED 
no. 4472 (no. 47635, TENN). Haywood Co.: GSMNP, 
Cataloochee, 6.ix.87, DED no. 4485 (no. 47636, TENN); 
GSMNP, Cataloochee, 9.ix.87, DED no. 4490 (no. 47637, 
TENN). Macon Co.: Coweeta Hydrologic Lab, Ball Creek, 
4.ix.86, DED no. 4134 (no. 47631, TENN); Coweeta 
Hydrologic Lab, Ball Creek, 13.viii.87, DED no. 4456: (no. 
47634, TENN); Highlands, Horsecove, 30.vii.87, DED no. 
4413 (no. 47632, TENN); Highlands, Horsecove, 10.viii.87, 
DED no. 4415 (no. 47629, TENN - Holotype). SOUTH 
CAROLINA: Oconee Co.: Ellicott Wilderness Area, Chattooga 
Picnic area, 12.viii.87, DED no. 4448 (no. 47633, TENN). 
TENNESSEE: Blount Co.: GSMNP, Cades Cove, 17.ix.85, DED 
nos) 3485) (no. 47630, TENN). | VIRGINIA: no data, R. 
Vilgalys, US 900746 (BPI). Giles Co.: Jefferson Nat. 
Forest, 2.x.83, O. K. Miller 20804 (VPI). Montgomery Co.: 
Blacksburg, VPI, 14.ix.83, R. Vilgalys 83/195, US 900773 
(BPI). Washington Co.: Little Tumbling Creek, 17.ix.83, R. 
Vilgalys 83/202, US 900121 (BPI). 


OBSERVATIONS. Marasmius falcatipes exhibits 
substantial morphological and substrate variability. The 
stipe-pileus insertion ranges from central to nearly 
lateral, with the full range of variation exhibited within 
or between collections. Pileus coloration varies from 
buff, greyish yellow or brownish yellow to brownish 
orange, brown or grey. Basidiomata have been found 
growing on senescent leaves or stems of various deciduous 


90 


hardwoods or herbaceous vines (e.g., Rubus, Lonicera), as 
well as grass leaves. Moreover, there appears to be no 
correlation between degree of stipe eccentricity, pileus 
coloration and substrate preference. 

Marasmius falcatipes belongs in sect. Sicci, subsect. 
Siccini Singer (1965) because of Siccus-type pileipellis 
elements, non-collariate lamellae, non-insititious stipe 
and dextrinoid tramal hyphae. Singer (1976) established 
two series within subsect. Siccini based on the presence 
of well-differentiated pleurocystidia (ser. 
Haematocephali) or their absence (ser. Leonini). Although 
M. falcatipes possesses distinctive pleurocystidia, it 
appears to be most phenetically similar to several taxa 
belonging to ser. Leonini, viz., M. armeniacus Gilliam 
(1975a) and M. pusio Berk. & Curt. (1853). Marasmius 
armeniacus [Holotype: Gilliam 932a (MICH!)] and M. pusio 
var. pusio [Isotype: Ravenel 816 (FH!)] both differ from 
M. falcatipes in longer stipes (at least two times longer 
than the width of the pileus) and having more polymorphic 
caulocystidia. In comparison, the stipe of M. falcatipes 
is usually shorter than the width of the pileus. Marasmius 
pusio var. guatopoensis (Dennis) Sing. essentially differs 
only in lacking pleurocystidia (fide Dennis, 1961). 

If greater taxonomic emphasis is placed on the 
presence of a short, eccentric stipe, it could be argued 
that M. falcatipes belongs in sect. Neosessiles Singer 
(1958). This section was established to unify all taxa 
combining the following characters: 1) pleurotoid habit 
(i.e., stipe eccentric, lateral or absent); 2) non- 
collariate lamellae; 3) dextrinoid tramal tissue; and 4) 
hymeniform pileipellis of diverticulate elements. 
Subsequently, Singer (1965) emended the section to include 
both dextrinoid ("pseudoamyloid") tramal hyphae (subsect. 
Neosessilini) and inamyloid tramal hyphae (subsect. 
Spaniophyllini). If taxa currently placed in sect. 
Neosessiles are re-evaluated utilizing characters in 
common usage at sectional or subsectional levels, it 
appears that sect. Neosessiles is somewhat heterogeneous. 
It contains taxa with Siccus- or Rotalis-type pileipellis 
elements, insititious or non-insititious stipes, 
dextrinoid or inamyloid tramal tissue, and with or without 
pleurocystidia [see Singer (1976) for a thorough treatment 
of the neotropical species]. Examination of the holotype 
specimens of a number of species currently placed in sect. 
Neosessiles (Desjardin, unpublished data) indicates that 
this section includes discordant elements. For example, 
M. polycystis Singer (1976) exhibits all characters 


oO) 


diagnostic of sect. Sicci except for the presence of a 
short, eccentric stipe. Indeed, when compared with M. 
falcatipes, M. polycystis differs only in pleurocystidial 
morphology. In the latter, pleurocystidia are broadly 
ventricose and average 13.6 um in width [n = 45 cystidia. 
Known only from the holotype: Singer B1499 (F!)], while 
in M. falcatipes, pleurocystidia are cylindrical with a 
mean width of 5.8 wm [TL90(90Z) = 4.9-6.7 um, n = 20/10 
specimens). The extensive variability in stipe-pileus 
insertion exhibited by M. falcatipes indicates that stipe 
eccentricity may be of limited taxonomic value. These data 
suggest that M. polycystis is better placed in sect. 
Sicci, presumably closely allied with M. falcatipes. 
Since many species in sect. Neosessiles are known only 
from their type specimens, which in some cases consist of 
a single basidioma, it would be premature to dismantle or 
redefine the section until further material is available 
for comparative studies. 


ACKNOWLEDGEMENTS 


This contribution is dedicated to Dr. Harry D. 
Thiers, whose profound knowledge of the fungi, consummate 
teaching ability and infinite patience inspired the senior 
author toward the study of mycology. We thank the 
following for accompanying us during field studies and for 
contributing personal specimens to our research: Drs. 
Heidi and Timm Anke, Timothy Baroni, Andreas Bresinsky, 
Roy Halling, Egon Horak, Andrew Methven, Harry Thiers and 
Zang Mu. We thank Drs. David Frodin (PH), John Haines 
(NYS), Roy Halling (NY), James Kimbrough (FLAS), Gregory 
Mueller (F), Donald Pfister (FH), Amy Rossman (BPI) and 
Robert Shaffer (MICH) for facilitating loan of specimens. 
We are also most grateful to Mrs. Ellen Thiers for 
checking Latin diagnoses. This research was funded in 
part by grants-in-aid to the senior author from the 
Highlands Biological Station, Highlands, NC, and by 
National Science Foundation Doctoral Dissertation 
Improvement Grant BSR- 8612110. 


LITERATURE CITED 


Berkeley, M. J. & M. A. Curtis. 1853. Centuries of North 
American fungi. Ann. Mag. Nat. Hist. 12: 417-435. 

Dennis, R. W. G. 1961. Fungi Venezuelani: IV. 
Agaricales. Kew Bull. 15: 67-156. 

Desjardin, D. E. 1985. New marasmioid fungi from 


92 


California. Mycologia 77: 894-902. 

1987a. New and noteworthy marasmioid fungi 
from California. Mycologia 79: 123-134. 

1987b. Tricholomataceae I. Marasmioid fungi: 
the genera Baeospora, Crinipellis, Marasmiellus, 
Marasmius, Micromphale and Strobilurus. Pages 1-99 
in H. D. Thiers (ed.), The Agaricales (Gilled Fungi) 
of California. Mad River Press, Eureka, California. 

Elias, T. S. 1980. The complete trees of North America. 
Van Nostrand Reinhold Co., New York. 948 p. 

Gilliam, M. S. 1975a. New North American species of 
Marasmius. Mycologia 67: 817-844. 

1975b. Marasmius section Chordales in the 
northeastern United States and adjacent Canada. 
Contr. Univ. Michigan Herb. 11: 25-40. 

1976. The genus Marasmius in the 
northeastern United States and adjacent Canada. 
Mycotaxon 4: 1-144. 

Halling, R. E. 1983. A synopsis of Marasmius section 
Globulares (Tricholomataceae) in the United States. 
Bri trond anor oly. 3267 

Halling, RR. E.,) Desjardin,) D.-&°R.°V. Tish “196a8 
Additions to Marasmius section Globulares. Mycotaxon 
22469747 6ls 

Kornerup, A. & J. H. Wanscher. 1978. Methuen handbook of 
colour, ed. 3. Eyre Methuen, London. 252 p. 

Morgan, A. P. 1905. North American species of Marasmius. 
Hie Myco Lt nil a2 O hao Ae oye 

. 1906. North American species of Marasmius. 

SUNMV Cole h 22-9 hielo oA ORR 

Parmasto, E. & I. Parmasto. 1987. Variation of 
basidiospores in the Hymenomycetes and its 
significance to their taxonomy. J. Cramer, Berlin- 
Stutteartenl6gap. 

Pennington, L. H. 1915. New York species of Marasmius. 
New York State. Mus. Bull. 179: 52-79. 

Singer, R. 1953. Type studies on Basidiomycetes. VI. 
LilloarZori57a195. 

1958. New genera of fungi. VIII. Notes 
concerning the sections of the genus Marasmius Fr. 
Mycologia 50: 103-110. 

1965. Monographic studies on South American 
Basidiomycetes, especially those of the east slope of 
the Andes and Brazil. 2. The genus Marasmius in 
South America. Sydowia 18: 106-358. 

1976. Marasmieae (Basidiomycetes-Tricholoma- 
taceae). Flora Neotropica Monogr. 17: 1-347. 


MYCOTAXON 


(oath ne ean LAR Ac a a ud Sd cs a a 
Vol. XXXIV, No. 1, pp. 93-113 January 20, 1989 


A SYNOPSIS OF COLOMBIAN BOLETES 


ROY E. HALLING 


Cryptogamic Herbarium 
The New York Botanical Garden 
Bronx, NY 10458-5126 


RESUMEN 


En el presente quince especies de Boletaceae son cono- 
cidas en el norte de los Andes en Colombia; ellos per- 
tenecen a Austroboletus, Boletellus, Boletus, Gyrodon, Lec- 
cinum, Strobilomyces, y Tylopilus. Associaciones ecto- 
mycorrizicas con Quercus humboldtii y Alnus acuminata son 
asumidas en la mayoria de los casos. Por primera vez en la 
taxa se describen Boletus orquidianus, Leccinum andinum, 
y Tylopilus obscurus. 


INTRODUCTION 


Several weeks collecting during October-November 1986 and May 1987 in the 
Andean regions of Antioquia, Boyaca, Cauca, Cundinamarca, and Huila of Colom- 
bia resulted in the discovery of a number of interesting taxa (e.g., Halling & Ov- 
rebo, 1987a,b). In particular, repeated trips to forests of Quercus humboladtii 
Bonpl. showed that members of the Boletaceae were fairly well represented. Also, 
in one area near Popayan, Alnus acuminata H.B.K. was found to support Gyrodon 
monticola Sing., originally described from Argentina (Singer & Digilio, 1958). 

Recent reports on Boletaceae from South America include works by Horak 
(1977), Ovrebo (1983), Singer (1964, 1970, 1973), Singer & Digilio (1951, 1958, 
1960), Singer et al. (1983), and Wolfe et al. (1988). Generally, these papers discuss 
boletes found in temperate/subantarctic Nothofagus forests, Amazonia, or other 
lowland tropical/subtropical regions of South America. Except for the report of 
Boletellus ananas (Curt.) Murr. and Boletus fuligineotomentosus Sing. from oak 
forests near Cali, Colombia (Singer, 1970, 1973), other Boletaceae (excluding Phyl- 
loporus) from that country have not been treated. Even though some taxa men- | 
tioned below also occur in the north temperate zone (North America, Europe, or 
Japan), I suspect these boletes are also native in Colombia [unlike Suillus luteus 
(Fr.) S.F. Gray and Boletus piperatus Fr. which are found associated with exotic 
Pinus spp.]. As mentioned previously (Singer, 1963, 1964; Singer & Morello, 
1960), the oak forest mycota of Colombia remains essentially unsurveyed, thus this 
treatment of Colombian boletes must be considered a preliminary one. 

Descriptions of Colombian boletes are given for those taxa that are newly de- 
scribed or of questionable identity, poorly known or thought to be true South 
American endemics. For species of Gyrodon, extralimital South American 


94 


material has been compared. Color notations given as page, column, row (¢.g., 
6C6) are from Kornerup & Wanscher (1978). Other color names are general ap- 
proximations. 


1. AUSTROBOLETUS SUBVIRENS (Hongo) C. B. Wolfe, Bibl. Mycol. 69: 125. 
1979. FIG. 1 
= Porphyrellus subvirens Hongo, Acta Phytotax. Geobot. 18: 110. 1960. ns 


Pileus 1.5 cm broad, convex, dry, with dull-green, granulose to crustose scales, 
sometimes fasciculated and then forming minute suberect or repent areolae. Flesh 
white, without odor or taste. Tubes porphyry pinkish, not changing when bruised. 
Stipe 3 cm long, 3-4 mm thick, +equal, dry, white, green-subreticulate above, ir- 
regularly scaly below; scales sometimes coalescing and appearing alveolate-ridged 
(under a lens); interior white, unchanging. 

Basidiospores smooth at first, with wall becoming minutely pitted, eventually 
dissected by meandering subreticulate channels that occasionally isolate irregular 
truncate warts or ridges, especially around middle, with walls up to 1.4 um thick, 
14.7-18.2 x 6.3-8.8 um (mean Q=2.17), broadly subfusoid to elongate- 
amygdaliform, with a suprahilar depression, yellow brown in KOH, unreactive or 
dextrinoid in Melzer’s. Basidia broadly clavate to subsaccate, hyaline, (3)4- 
sterigmate, 24-35 x 12-15 um. Hymenial cystidia absent. Tube trama divergent 
from a central stratum, with gelatinized lateral stratum; hyphae hyaline, 3.5-6.3 4m 
broad. Pileus surface a trichodermium of long, cylindrical elements, sometimes fas- 
cicled, 5-10 um broad, smooth- and thin-walled, with dark green vacuolar pigment, 


Fics. | a,b. Scanning electron micrographs, basidiospores of Austroboletus sub- 
virens (Halling 5955). Standard line = 2 pm. 


w be 


dissolving in KOH, irregularly coagulated in Melzer’s and water mounts. Stipe sur- 
face lacking caulocystidia, with hyaline, narrow, subgelatinized hyphae, overlain 
with cylindrical elements forming reticulum, the latter morphologically similar to 
trichodermial elements of pileus. Clamp connections absent. 

Material examined: DEPT. HUILA: near Parque Nacional Puracé, Finca de Gana- 
deria Merenberg, under Quercus humboldtii, 21 May 1987, Halling 5955 (NY). 

The identity of this collection is tentative because only one, diminutive 
basidiocarp was found, and A. subvirens has been reported previously only from 
Japan and New Guinea. Despite the small size, other important characters such as 
the overall appearance of surface features, coloration, spore morphology and other 
microscopic features, correspond quite well to the protologue (Hongo, 1960), a 
later evaluation of the type specimen (Wolfe, 1979), and illustrations and com- 
ments on additional fresh material (Horak, 1980). However, more collections 
from Colombia are needed to assess the range of typical variation. 


2. BOLETELLUS ANANAS (Curt.) Murr., Mycologia 1: 10. 1909. 
= Boletus ananas Curt., Amer. J. Sci. II 6: 251. 1848. 


I have not seen specimens from Colombia, but information and a description 
of Colombian material can be found in Singer (1970) where one collection is cited: 
"VALLE. Rio Pance, 1800 m alt, 3 May 1968, Singer B6942 (F)." Also known to 
occur north to the southeastern United States and in southeast Asia. 


3. BOLETELLUS RUSSELLII (Frost) Gilbert, Bolets. 107. 1931. 
= Boletus russellii Frost, Bull. Buffalo. Soc. Nat. Sci. 2: 104. 1874. 


Only one immature basidiocarp was found in a 30 year-old, second-growth oak 
forest near Popayan. Despite the lack of spores, the long, coarsely lacerate- 
reticulate dull-reddish stipe with a viscid white base is an unmistakable field 
character. Detailed descriptions of material from the United States are available 
in Singer (1945) and Smith & Thiers (1970). 


Material examined: DEPT. CAUCA: Municipio de Tunia, km 93 of Cali-Popayan 
road, Reserva Forestal "El Guayabo," 1800 m, under Quercus humboldtii, 19 May 1987, 
Halling 5270 (leg. G. Mueller) (NY). 


4. BOLETUS ATKINSONIANUS (Murr.) Sacc. & Trott., Syll. Fung. 21: 236. 
1912. FIGS. 2-5 
= Ceriomyces atkinsonianus Murr., N. Amer. Flora 9: 144. 1910. TYPE: United 

States. NORTH CAROLINA: Pink Bed Valley, Jul 1908, Murrill & House 64 (Holo- 
type: NY!). 

Pileus 2.5-8.5 cm broad, convex to plano-convex, viscid when wet, matted 
tomentose or areolate when dry; surface brown (7E8) to reddish brown (9E8) on 
disc and brownish orange (6C5) at margin when young, brownish orange (7CS) to 
light brown (7DS) to dull reddish (8C4-8C5) with age, uneven to shallowly pitted- 
rugulose or wrinkled in some; margin sterile and projecting about 1-1.5 mm when 
young, not so much with age; flesh 4-11 mm thick, white, unchanging, slightly red- 
dish brown under pileus surface. Odor and taste mild. Tubes 5-17 mm long, un- 
changing when exposed, lemon yellow (2A8) to light yellow (3A5) when young, 


96 


Fic. 2. Habit of Boletus atkinsonianus. (Halling 5000) x1/2. 


more greenish yellow (3D7-3E7) with age; pores up to 1 per mm, light yellow (3A5) 
when young, mustard yellow to wax yellow (3B6-3B5) with age, unchanging when 
bruised. Stipe 7.5-12 cm long, 8-11 mm thick at apex, subclavate to broader below 
or sometimes +equal, strict to somewhat curved, solid; surface subviscid to tacky 
when wet, glabrous, lacking reticulations, but some broad and obscure rugulose 
zones present or coarse alveolate ridges at apex when very young, streaked with 
reddish brown (8D6), with some yellowish to grayish orange (6BS) ridges near 
apex to half the length on white ground color, reddish brown (8D6) more abundant 
at apex when young, becoming whitish there with age, reddish brown (8D6) more 
abundant below with age, but with a white base throughout development; interior 
white, solid, unchanging. | 

Basidiospores olivaceous in fresh deposit, yellowish brown in KOH, occasional- 
ly some dextrinoid in Melzer’s, smooth, thin-walled, subfusoid and inequilateral in 
profile to oblong ellipsoid, 11.9-16.8 x 4.9-6.3 4m (mean Q=2.7). Hymenophore 
lemon yellow in water mounts, this color disappearing in KOH mounts, often with 
dextrinoid encrusting pigment in Melzer’s when young. Basidia 24-32 x 8-11 ym, 
clavate, 4-sterigmate, hyaline. Pleurocystidia hyaline in KOH, hyaline or rarely 
pale orangish in Melzer’s, fusoid, thin-walled, 35-53 x 8-15 um. Cheilocystidia 
barely subfusoid to subclavate, hyaline in KOH, pale orangish in Melzer’s, not as 
abundant as pleurocystidia, 30-40 um long. Tube trama divergent from a central 
Stratum, with gelatinized lateral strata, hyaline, 3.5-7 um broad. Pileus trama 


ay, 


py) 


5 um 4 


Fics. 3-5. Cystidia of Boletus atkinsonianus (Halling 5026). 3. Caulocystidia. 4. 
Pleurocystidia. 5. Cheilocystidium. 


hyaline in KOH and Melzer’s, interwoven, with hyphae 5-9 4m broad. Pileus sur- 
face a tangled trichodermium in a gelatinous matrix, with elements subcylindrical, 
rarely inflated, often vermiform in general aspect, smooth, with vacuolar pigment, 
usually dissolving in KOH or yellow brown, coagulated and orangish in Melzer’s, 
(2.8)3.5-8.4(10.5) um broad. Stipe surface composed of vertically oriented, parallel 
hyphae, becoming gelatinized with age, hyaline in KOH, encrusted with dextrinoid 
pigment and penicillate crystals in Melzer’s, giving rise to caulocystidia, the latter 
often clustered, hyaline and smooth in KOH, sometimes with encrusting dextrinoid 
pigment, thin-walled, clavate to capitate clavate, generally gelatinizing with age, 
24-50 x 7-14(18) um. Clamp connections absent. 

Material examined: DEPT. ANTIOQUIA: Municipio Santa Rosa de Osos, near Llanos | 
de Cuiva, 6°45’N, 75°30’W, +2500 m, under Quercus humboldtii, 10 Nov 1986, Halling 
5000 (HUA, NY); 14 Nov 1986, Halling 5026 (HUA, NY); 25 Nov 1986, Halling 5057 
(HUA, NY); road between San José de la Montafia and Llanos de Cuiva, 6°50’N, 
75°3S°W, +2500 m, under Quercus humboldtii, 27 Nov 1986, Halling 5077 (HUA, NY). 

Boletus atkinsonianus was originally described from North Carolina, USA, ap- 
parently based on Atkinson’s (1901) concept of B. obsonium (Paul.) Fr., but seems 
not to have been described in detail since the description by Coker & Beers 
(1943). Diagnostic features include a viscid pileus and stipe with distinctive brown 
colors, projecting pileus margin, bright yellow hymenophore when young that, 
along with the flesh, is unchanging. Perhaps because of age and preservation, the 


98 


dextrinoid pigment in the stipe and young hymenophore of the type specimen is 
absent, but would seem to be a further distinction in fresher material. Most likely, 
B. atkinsonianus might be confused with B. viridiflavus Coker & Beers, but the lat- 
ter is a smaller fungus with the pileus mottled a greenish yellow (Halling 3788, 
NY). A color illustration of B. viridiflavus (Weber & Smith, 1985) from the south- 
eastern United States is reminiscent of the Colombian B. atkinsonianus. Guzman- 
Davalos et al. (1983) have reported the latter from Jalisco, México. 


5. BOLETUS FULIGINEOTOMENTOSUS Singer, Beih. Sydowia 7: 101. 1973. 


I have not seen specimens from Colombia, although I have collected near the 
paratype locality. Known only from Colombia, information and a description of B. 
fuligineotomentosus can be found in Singer (1973) where three collections are 
cited: "Typus a R. Singer (B 6958) in Columbia: Valle: Rio Pance, 1800 m alt., 3 V 
1968 lectus et in F conservatus.-Paratypus: Singer B 6466, Cauca: via a Cali 
Popayan versus, km 100, 29 IV 1968.-Etiam B 6790, El Guayabo 29 IV 1968, 
omnes in F.-Boleto subsolitario affinis." 


6. BOLETUS ORQUIDIANUS sp. nov. FIGS. 6-9 


Pileus viscidus, pileipelle hymeniformi; trama tuborum phylloporoideo; cystidia 
hymenii nulla; basidiosporae 10.5-13.3 x 4.2-5.6 um, laeves, interdum dextrinoideae. 

Pileus 2-2.5 cm broad, convex to plano-convex, reddish brown to brown (7F8- 
8E8), paler at the margin, moist to viscid, glabrous, not changing with NH4OH. 
Flesh white, not bluing, with miid odor and taste. Tubes depressed around the 


Fic. 6. Habit of Boletus orquidianus (Halling 4964, HOLOTYPE). x2. 


99 


stipe, olive yellow to grayish yellow (3C6-3CS) not changing to blue when exposed, 
5-7 mm long; pores up to 1 mm broad, concolorous, not changing to blue when 
bruised. Stipe 4-8 cm long, 4-5 mm broad at apex, + equal to slightly larger at base; 
surface white at apex and base, streaked brownish (7E7) in mid-region, lubricous 
but not viscid,glabrous. 

Basidiospores smooth, thin-walled, ellipsoid to subfusoid, golden yellow in 
KOH, a minority slightly dextrinoid in Melzer’s, 10.5-13.3 x 4.2-5.6 um. Basidia 
clavate, 4-sterigmate, hyaline, 28-35 x 12-15 um. Hymenial cystidia absent. Tube 
trama slightly bilateral (Phylloporus-type), with hyphae hyaline, 2.8-8.4(14) wm 
broad. Pileus surface a hymeniform epithelium embedded in an obvious gelatinous 
matrix, with elements hyaline or golden yellow to yellow brown in KOH, hyaline or 
sometimes with dull orangish contents in Melzer’s, broadly clavate to napiform or 
sphaeropedunculate, sometimes subisodiametric, appearing sphaerocyst-like in pa- 
radermal section, smooth and thin-walled, 17.5-31.5 x 14-24.5 um. Pileus trama in- 
terwoven, hyaline in KOH, pale orange in Melzer’s, with hyphae 4.2-9.8(14) um 
broad. Stipe surface composed of parallel and vertically oriented hyphae, 2.8-7 um 
broad, giving rise to scattered clumps of caulocystidia that are clavate to sub- 
cylindric, 21-35 x 10.5-14 um, rarely with an occasional dermatobasidium. Clamp 
connections absent. 

TYPE: COLOMBIA. DEPT. ANTIOQUIA: Municipio Urrao, Parque Nacional 
Natural "Las Orquideas," sector Calles, 1300-1400 m, 30 Oct 1986, Halling 4964 
(HOLOTYPE: HUA; ISOTYPE: NY). 


Boletus orquidianus is not associated with Fagaceae in Colombia, but rather 
with other hardwood trees in the submontane cloud forest. Boletus orquidianus 


O07 Qu 


Fics. 7-9. Microscopic features of Boletus orquidianus (Halling 4964, HOLOTYPE). 
7. Basidiospores. 8. Elements of pileus surface. 9. Caulocystidia. 


100 


keys to Xerocomus globuliger Sing. in Singer et al. (1983), which differs in a dry (not 
viscid) and granular to rivulose pileus surface, longer spores (13-17.5 um), and the 
presence of hymenial cystidia and pilocystidia. The viscid pileus and Phylloporus- 
type trama would also suggest Pulveroboletus (sensu Singer), but the hymeniform 
pileus surface of B. orquidianus is like that found in Marasmius sect. Globulares but 
with a gelatinous matrix. Furthermore, none of the pileus surface types (cutis, 
ixocutis, trichodermium, or palisade) described for Pulveroboletus by Singer (1986) 
are applicable to B. orquidianus. Additionally, the spores of B. orquidianus are 2-5 
um longer than the non-velate Pulveroboleti listed by Singer et al. (1983) and 
Pegler (1983). 


7. BOLETUS PULVERULENTUS Opat., Arch. Naturgesch. 2: 27. 1836. 


The dry, dark brown to nearly black pileus, yellow hymenophore and flesh that 
turn instantly to blue when bruised or exposed are key macroscopic features. Mi- 
croscopically, the abundant hymenial cystidia with dextrinoid granular content are 
likewise distinctive. Known previously from the north temperate zone, the single 
Colombian collection of several basidiocarps agrees in all respects with material 
from eastern North America. 


Material examined: DEPT. CUNDINAMARCA: near Pacho, Finca "La Ramada," under 
Quercus humboldtii, 11 May 1987, Halling 5251 (NY). 


8. BOLETUS PSEUDORUBINELLUS Smith & Thiers, Boletes of Michigan. 300. 
1970. FIG. 10 


Pileus 2-6.5 cm broad, dry, convex to plano-convex, violet brown (10ES) at 
first, then english red to reddish brown (8D8-8D6), matted subtomentose to 
matted subvelutinous, barely with finely appressed squamules near the margin with 
dull ochraceous color showing through. Flesh up to 1.5 cm thick, near pinkish 
white (7A2) but +paler with a faint tint of yellow, unchanging; odor and taste mild. 
Tubes near old rose (10DS) at first, near reddish brown (8D6) with age; pores con- 
colorous, unchanging, + 1 per mm, adnate or depressed around the stipe. Stipe 5-7 
cm long, 4-12 mm broad, equal or tapering downward, fibrous, curved or strict, 
fibrillose striate, near brick red (7D7), dry, with pinkish zone at apex, white or with 
some pale pinkish tones toward the base, with yellowish mycelium extending into 
substrate; interior yellowish, solid. 

Basidiospores smooth, thin-walled, pale olivaceous at first, but then bright 
greenish yellow in KOH, some becoming lightly dextrinoid in Melzer’s, 9.1-12.6 x 
4.2-4.9 um (mean Q=2.28), ellipsoid to subfusoid and inequilateral in profile. 
Basidia 4-sterigmate, clavate, hyaline, 19.6-31.5 x 8.4-10.5 um. Hymenial cystidia 
rare, narrowly fusoid ventricose, hyaline, 35-47 x 7-11 um, more abundant on the 
pores and then sometimes with amorphous pale brown content in KOH. Pileus 
surface a trichodermium of suberect, tangled, cylindrical cells, 5-9 4m broad, soon 
collapsing and becoming repent; hyphae hyaline to pale ochraceous yellow from 
pigment dissolving in KOH, but pigment remaining in Melzer’s and appearing 
granular-encrusting. Pileus tramal hyphae hyaline in KOH, somewhat collapsed 
and sometimes with a pale orangish tint in Melzer’s, otherwise inamyloid, some- 
times with scattered oleiferous elements. Caulocystidia often clustered, narrowly 


101 


fusoid ventricose, clavate, or sometimes spherical-saccate, thin-walled, 17.5-65 x 7- 
18 xm, without distinctive contents in KOH or Melzer’s. Clamp connections absent. 

Material examined: DEPT. CAUCA: Municipio de Tunia, km 93 of Cali-Popayan 
road, Reserva Forestal "El Guayabo," 1800 m, under Quercus humboldtii, 19 May 1987, 
Halling 5267 (NY). 

In the original description of Smith & Thiers (1970), B. pseudorubinellus was 
described as occurring near spruce in northern Michigan. Despite the difference 
in supposed mycorrhizal hosts, Halling 5267 agrees in overall colors, spore mor- 
phology, yellowish reaction in KOH, lack of an acrid taste, and general lack of 
hymenial cystidia. 


9. BOLETUS SUBTOMENTOSUS Fr., Syst. Mycol. 389. 1821. 
= Xerocomus subtomentosus (Fr.) Quél., Fl. Myc. 418. 1888. 


The olive to yellow olive (4D6), dry, subtomentose pileus, yellow tubes that 
change to blue when cut and exposed, compound pores, yellowish stipe with some 
occasional coarse ribbing at the apex, and yellowish basal mycelium are useful 
field characters. Also, the pileus surface does not react with NH4OH (blue to 
greenish). The descriptions of Smith & Thiers (1971) and Thiers (1975) fit the 
Colombian material well. 


Material examined: DEPT. ANTIOQUIA: Municipio Santa Rosa de Osos, road from 
San José de la Montana to Llanos de Cuiva, 6°50’N, 75°35’W, +2500 m, under Quercus 
humbolatii, 25 Nov 1986, Halling 5061 (HUA, NY). 


Fic. 10. Habit of Boletus pseudorubinellus (Halling 5267). x3/4. 


102 


10. BOLETUS TRUNCATUS (Sing., Snell, & Dick) Pouzar, Ceska Mycol. 20: 2. 


1966. FIG. 11 
= Xerocomus truncatus Sing., Snell, & Dick in Snell, Singer & Dick, Mycologia 51: 
573. 1959. 


Pileus 1.5-4 cm broad, convex to plano-convex, dry, velutinous to sub- 
tomentose, bronze brown (SES). Flesh pale yellow to white (3A3-2A3), not chang- 
ing to blue when exposed. Tubes pastel yellow to grayish yellow (2A4-2B5), com- 
pound, 5-9 mm long, with faint hint of changing to blue when exposed; pores con- 
colorous. Stipe 3-4 cm long, 4-6 mm thick, +equal, surface furfuraceous to 
pruinose-squamulose (red to pinkish) on yellow ground color above, pink at mid- 
dle, pale brown below, eventually with a yellowish white (4A2) ground color. 

Basidiospores truncated at the apex and then usually thicker-walled at the two 
apical angles and with markedly thinner wall between these angles, sometimes with 
faint but distinct longitudinal striae (with interference optics), yellowish brown in 
KOH, often fleeting amyloid and then some slowly and partially or completely dex- 
trinoid in Melzer’s, 11.2-15.4 x 4.9-5.6 um (mean Q=2.5). Basidia 4-sterigmate, 
hyaline to pale yellowish in KOH and Melzer’s, 24-42 x 9-12 um. Hymenial cystidia 
not abundant but conspicuously projecting, fusoid to subventricose, hyaline in 
KOH and Melzer’s, 35-68 x 9-12 um. Tube trama of the Phylloporus-type, obscure- 
ly bilateral, hyaline in KOH and Melzer’s, with hyphae 3.5-8.4 um broad. Pileus 
surface a palisade trichodermium, with elements ochraceous in KOH, brown in 
Melzer’s, generally encrusted, with individual hyphae barrel-shaped to isodiametric 
or nearly so, with apical cells subspherical to subcylindric, obtuse at the tips, (7)17- 
28 pm broad. Pileus trama hyaline, interwoven, with hyphae 7-17 um broad. Stipe 
surface composed of vertically oriented hyphae giving rise to isolated fascicles of 
caulocystidia, the latter subclavate to clavate or rarely subcapitate, with yellow 
brown contents or rarely hyaline, in KOH, 24-42 x 7-11 um, with occasional 2,4- 
sterigmate caulobasidia intermixed. Clamp connections absent. 

Material examined: DEPT. ANTIOQUIA: Municipio Santa Rosa de Osos, road from 
San José de la Montana to Llanos de Cuiva, 6° 50’N, 75°35’W, +2500 m, under Quercus 
humbolatii, 25 Nov 1986, Halling 5059 (HUA, NY). 

This is a tentative determination although the macroscopic features (very 
similar to B. chrysenteron Fr.) and truncate spores point toward B. truncatus. The 
presence of longitudinal striae are foreign to the protologue, current concepts and 
descriptions, but there appear to be other xerocomoid taxa in the United States 
which possess striate spores that may or may not be truncate. 


11. GYRODON EXIGUUS Singer & Digilio, Lilloa 30: 154. 1960. FIG. 12 


Pileus 1-3 cm, flabelliform to fan-shaped to infundibuliform, dry, matted fibril- 
lose scaly, especially at margin, with brown, appressed fibrillose scales on yellowish 
(4A5-4B5-5B5) ground color. Flesh yellow, up to 2 mm thick, slowly bluing, with 
mild odor and taste. Hymenophore decurrent, boletinoid, with compound pores, 
definitely radial in orientation, yellowish (4A6), bluing at first when bruised or cut, 
then changing to brown. Stipe lateral or sometimes eccentric, up to 1.5 cm long, 8 
mm thick at apex, yellow to pale orange (5A3) above, olive brown to watery 
brownish toward base, matted tomentose with a lens, arising from a dirty yellow- 


103 


Fic. 11. Scanning electron micrograph, basidiospores of Boletus truncatus (Halling 
5059). Standard line = 1.4 pm. 


brown superficial mycelium, the latter sometimes forming rhizomorphic strands 
over the substrate. 

Basidiospores olive brown in deposit, 7-9.8 x 4.9-6.3 um, (mean Q=1.55), 
hyaline to pale yellow brown in KOH, fleeting amyloid at first, but then some dex- 
trinoid or pale yellowish brown in Melzer’s, smooth, ovoid to broadly ellipsoid, 
rarely short-subcylindric. Basidia clavate, 4-sterigmate, hyaline, 22-30 x 6.5-9 um. 
Hymenial cystidia rare to scattered and inconspicuous, hyaline, subclavate with a 
broad mucro, or subcylindric, sometimes with short apical ramifications, 15-25 x 4- 
9 um. Tube trama hyaline, bilateral, with hyphae 2.1-8.4 wm broad. Pileus surface 
a tangled trichodermium with elements that form the fibrils possessing a red brown _ 
encrusting pigment in Melzer’s, the pigment soluble in KOH, sometimes with 
homogeneous ochraceous contents, thin-walled, +cylindric to filamentous, 3.5-8 
um broad. Pileus trama hyaline in KOH, hyaline to pale orange in Melzer’s, with 
hyphae interwoven, thin-walled, smooth, 3.5-10.5 um broad. Stipe surface a col- 
lapsed trichodermium with hyphae resembling those of the pileus surface, arising 
from a hyaline trama with elements 4-15 um broad. Clamp connections present. 

Material examined: COLOMBIA. DEPT. ANTIOQUIA: Municipio Urrao, Parque Na- 
cional Natural "Las Orquideas," sector Calles, 1300-1400 m, 28 Oct 1986, Halling 4942; 


30 Oct 1986, Halling 4969 (both HUA, NY). ECUADOR. PRov. PASTAZA: 2 km N of 
Rio Pastaza-Rio Topo confluence, 15 April 1987, Halling 5168 (QCA, NY). 


104 


The dingy yellowish pileus with brown appressed fibrils, yellow hymenophore 
and flesh that become blue with handling and exposure, eccentric to lateral stipe, 
and small size (for a bolete) are diagnostic macroscopic features. In my experi- 
ence, G. exiguus fruits on standing tree trunks or wet soil banks that are covered 
with rootlets, humus, etc. rather than on the forest floor. A mycorrhizal connec- 
tion has not been demonstrated for G. exiguus as it has for the following species. 


12. GYRODON MONTICOLA Sing. in Singer & Digilio, Lilloa 28: 256. 1957. 
FIG. 13 


Pileus 3-11 cm broad, viscid in wet weather, otherwise dry, dark cinnamon 
brown, matted fibrillose, convex or rarely a with subacute low umbo, with incurved 
margin at first. Flesh whitish, bluing when exposed, up to 2 cm thick. 
Hymenophore boletinoid, decurrent, with compound pores, shallow, up to 2 mm 
long, yellow, bluing when bruised. Stipe 2-6 cm long, 5-14 mm thick, + equal to 
subclavate or narrower downward, central to eccentric, curved or strict; surface 
dry, pinkish brown, more sordid toward the base, especially after handling, matted 
subtomentose to matted subfibrillose, arising from well-developed, dirty yellowish 
brown basal mycelium, developing scattered concolorous sclerotia, the latter hard 
but brittle, dull when fresh, wrinkled and shiny when dry, 1-2(3) mm broad, 
globose to subglobose. 

Basidiospores olive brown in deposit, 5.6-7 x 3.5-4.9 um (mean Q=1.46), 
smooth, hyaline to pale yellow brown in KOH, some lightly dextrinoid in Melzer’s, 
ovoid to short ellipsoid, sometimes inequilateral in profile. Basidia 31-42 x 7.5-9 
um, 4-sterigmate, clavate, hyaline or rarely with pale yellow brown pigment. Tube 
trama hyphae bilateral, hyaline, 3.5-9(13) um broad. Hymenial cystidia most 
abundant at and near pores, generally hyaline, sometimes with amorphous yellow 
brown content, thin-walled, narrowly fusoid-ventricose with ampullaceous apex, 
30-60 x 7.5-9 um. Pileus surface a trichodermium of loosely interwoven hyphae, 
with elements generally cylindrical, but sometimes irregularly inflated or strangu- 
lated, occasionally branched, thin-walled, smooth and hyaline to ochraceous in 
KOH, pale orange brown and with red brown encrusting pigment in Melzer’s, 
(2.1)3.5-7(9) 4m broad. Pileus trama hyaline, with pale rosy purplish pigment 
leaching out in KOH mounts, hyaline to pale yellowish orange in Melzer’s; hyphae 
4-8 um broad, smooth and thin-walled. Stipe surface barely differentiated from 
tramal hyphae, loosely interwoven, generally repent, hyaline or rarely with 
ochraceous content, smooth in KOH and Melzer’s, 3.5-7 um broad, subtended by 
tramal hyphae often inflated to 21 wm, leaching a pale rosy purplish pigment in 
KOH, with walls usually thickened to 1 um. Clamp connections present. 

Material examined: ARGENTINA. PRov. TUCUMAN: Dept. Tafi del Valle, +5 km 
N of Tafi del Valle, under Alnus acuminata, 18 Mar 1988, Halling 5849 (NY). 
COLOMBIA. DEPT. HUILA: E of Parque Nacional Puracé, Finca de Ganaderia Meren- 
berg, under Alnus acuminata, 21 May 1987, Halling 5287 (NY). 

Gyrodon monticola appears to be exclusively associated with Alnus acuminata 
in South America. In fact, Halling 5287 was found in nursery beds consisting solely 
of A. acuminata at Finca Merenberg. The Colombian material corresponds well 
with a collection (Halling 5849) from near the Argentine type locality. Gyrodon 


105 


monticola was also found by G. Mueller in Alnus forests of Ecuador (Napo Pro- 
vince, +8-9 km east of Papallacta, along road to Baeza), but voucher material was 
not retained because of its advanced age. The sclerotia of G. monticola are nearly 
the same morphologically as those reported for G. merulioides (Schw.) Sing. (as 
Boletinellus) by Cotter & Miller (1985) but seem to differ in color. 


Fics. 12-13. Habits of Gyrodon exiguus and G. monticola. 12. G. exiguus (Halling 
4969) x2. 13. G. monticola (Halling 5849) x}. 


106 


13. LECCINUM ANDINUM sp. nov. FIGS. 14-18 


A speciebus aliis sectionis Luteoscabrorum, nocta semper cyanescens, coloribus 
uniformiter succineis vel brunneo-flavidis, pileipelle trichodermium cystidioideum 
revocanti, dermatobasidiis bisterigmatis, et cum Querco humboldtii consortione sua dis- 
tinguenda. 


Pileus (4)9-11.5 cm broad, convex to plane, dry, minutely subtomentose when 
young, subviscid to tacky and glabrous with age, grayish yellow (4B4) at the 
margin, darker near brownish yellow (5C6) on the disc when young, brass yellow 
(4C7) at the margin with age and raw sienna (6D7) at the disc, honey yellow (5D6) 
to light orange (SAS) to cinnamon brown (6D6) to amber yellow (4B6) in 
splotches, or brownish yellow (SC6-5C7) to yellowish brown (5D8-6D7); margin 
often sterile and projecting. Flesh 7-20 mm thick, whitish to yellowish white to 
orange white (4A2-5A2), bluing slightly near the tubes when exposed, but not in- 
tensely, slightly reddish-brown under the pileus surface. Tubes 3-20 mm long, 
depressed around the stipe, light yellow (2A5-3A5-3A6) when young, olive yellow 
(3D6) with age, bluing when exposed, boletoid; pores up to 2-3 per mm, yellow to 
deep yellow (3A7-4A8) when young (slight brownish discoloration) apparently 
stuffed, bluing when bruised. Stipe 3-12 cm long, 1-2 cm thick at apex, + equal to 
subequal to subclavate to slightly enlarged below, strict or curved, dry, densely 
scabrous when young, less so with age; ground color pale yellow (4A3) when 
young, grayish yellow (4B4) with age, white at the base; scabers yellow (2A5) when 
young, yellow ochre to brownish yellow to light brown (SD8-5C8-5C7-5C6) to 
brownish orange to light brown (6C7-6D7,6,5) with age; interior solid, yellow 
(2A5) when young and bluing near the apex, marbled brown and white at the base. 

Basidiospores olive brown to umber brown in fresh deposit, 14.7-23.8 x 6.3-8 
um (mean Q=2.88), smooth, subfusoid to ellipsoid, usually inequilateral with a 
suprahilar depression in profile, ochraceous in KOH, with dextrinoid walls in Mel- 
zer’s. Basidia 31.5-40 x 12-15 um, 4-sterigmate, clavate, hyaline. Hymenial cystidia 
moderately abundant, more common near pores, thin-walled, hyaline or rarely 
with homogeneous ochraceous content in apex, ventricose rostrate to lageniform 
or barely subfusoid to narrowly ampullaceous, 50-73 x 5-11 wm. Tube trama 
bilateral from a central strand that is sometimes tinged yellowish in KOH, other- 
wise hyaline, the lateral strata becoming subgelatinous, with elements 2-7 um 
broad. Pileus surface a loosely interwoven trichodermium, with hyphae seemingly 
embedded in a hyaline matrix, yellow ochraceous in KOH, sometimes with 
ochraceous homogeneous contents, hyaline to pale ochraceous yellow in Melzer’s, 
consisting of +filamentous, branched, occasionally cylindric, suberect smooth- 
walled elements, with endcells 2.8-5.6 4m broad, sometimes cystidioid and then 
subcapitate to inflated or obtuse to slightly tapered, intercalary cells sometimes 
subisodiametric, arising from a layer of broader sphaerocyst-like hyaline hyphae, 
12-30 x 10-21 um broad. Pileus trama interwoven with hyaline hyphae, 3.5-10.5 #m 
broad. Stipe surface covered with isolated scabrosities, these sometimes fused- 
subreticulate, composed of versiform caulocystidia, narrowly lageniform to 
ventricose rostrate, subcylindric or sometimes filamentous with short-branched 
apices, hyaline and thin-walled, often arising from subisodiametric cells, 30-78 x 3- 
12 wm; caulobasidia 2-sterigmate, broadly clavate, generally hyaline, intermixed 
with caulocystidia. Clamp connections absent. 


107 


Figs. 14-15. Habits of Leccinum andinum. 14. Halling 5001 xli/2. 15. Halling 
5052 (HOLOTYPE) x1/2. 


5 um 


Fics. 16-18. Microscopic features of Leccinum andinum (Halling 5052, HOLOTYPE). 
16. Basidiospores. 17. Elements of stipe surface. 18. Hymenial cystidia. 


TYPE: COLOMBIA. DEPT. ANTIOQUIA: Municipio Santa Rosa de Osos, 
near Llanos de Cuiva, 6°45’N, 75°30’W, +2500 m elev, under Quercus humboldtii, 
25 Nov 1986, Halling 5052 (HOLOTYPE: HUA; ISOTYPE: NY). 

Additional material examined: DEPT. ANTIOQUIA: Municipio Santa Rosa de Osos, 
near Llanos de Cuiva, 6°45’N, 75°30’W, +2500 m elev, under Quercus humboldtii, 10 
Nov 1986, Halling 5001 (HUA, NY); 14 Nov 1986, Halling 5027 (HUA, NY); 
Municipio San José de la Montana, +13 km S of San José de la Montafia along road to 
Labores, +2500 m, under Quercus humboldtii, 26 Nov 1986, Halling 5064 (F, HUA, 
NY). 

Macroscopically this Leccinum approaches Boletus morrisii Pk. with regard to 
overall color schemes, but differs in bluing reactions (reddening in the latter), 
white basal mycelium (yellow in B. morrisii), and the pores are yellowish or rarely 
brownish in L. andinum (reddish in B. morrisii). Furthermore, L. andinum has 
larger spores (12-15.4 x 3.4-4.3 um in B. morrisii [TENN 26054, 42113]), and the 
erect trichodermium composed of encrusted, broad, compact elements of B. mor- 
risti contrasts markedly with the narrow diameter hyphae of the loosely interwoven 
trichodermium in L. andinum. The association with Quercus, the yellowish con- 
text, and yellow hymenophore clearly indicate section Luteoscabra, but consistent 
bluing reactions have not been reported in that section before. Other taxa of 
Luteoscabra that sometimes are cyanescent include L. subglabripes (Pk.) Sing. and 


109 


its variants (Singer, pers. com.), but that species is differently colored on the pileus 
and stipe, has smaller spores, and the hyphae of the pileus surface form a 
hymeniform epithelium. 


14. STROBILOMYCES CONFUSUS Singer, Farlowia 2: 108. 1945. FIG. 19 


A few immature basidiocarps with an intact veil were found along with one 
basidiocarp that was over-mature. The spores from the latter are unmistakable 
and possess the "sparassoid" type of ornamentation characteristic for S. confusus. 
The dry, suberect, black squamules on the pileus, the gray and soon blackening 
hymenophore, reddening flesh on exposure, and woolly stipe surface further dis- 
tinguish the species. Excellent descriptions are available in Singer (1945) and 
Smith & Thiers (1970). 


Material examined: DEPT. HUILA: E of Parque Nacional Puracé, Finca de Gana- 
deria Merenberg, under Quercus humboldtii, 21 May 1987, Halling 5280 (NY). 


Fic. 19. Scanning electron micrograph, basidiospores of Strobilomyces confusus 
(Halling 5280). Standard line = 2 pm. 


15. TYLOPILUS OBSCURUS sp. nov. FIGS. 20-23 


Pileus usque 13 cm latus, brunneus vel griseobrunneus vel atrogriseus, tomentosus vel 
subvelutinus, atrogriseus quum dessicatus. Contextus pallidus, immutabilis. Tubuli pal- 
lidi tum subroseocinnamomei, immutabiles; pori brunneo-umbrini vel ochraceobubalini 
tactu cinnamomei. Stipes ttotus conspicue alveolatoreticulatus, pagina sub- 
roseocinnamomei; reticulum pileo concolor ad apicem pallidum aetate subcanescens com- 
positum. Sporae 10.5-15.4(20) x 4.2-5.6 wm. Cystidia pigmentifera parietibus tenuibus. 


110 


Pileus 4-13 cm broad, convex then plane, dry, tomentose to subvelutinous to 
matted tomentose to matted velutinous, brown to dark brown to grayish brown to 
dark gray (7E-F5,4,3); margin slightly incurved to decurved, fertile, uplifted and ir- 
regular with age. Flesh near grayish orange (SB3) or whitish, unchanging, 1.5-2 cm 
thick, odor and taste mild. Tubes up to 12 mm long, subdecurrent or depressed, 
orange white (SA2) when young, near light pinkish cinnamon (6CS) at maturity, 
not changing when exposed or bruised; pores 1-2 per mm, cocoa brown to burnt 
umber to teak brown (6E-F6,5) when young, ochraceous buff (5B4) with maturity, 
near cinnamon (6D7) when bruised. Stipe 8.5-15 cm long, 1.3-4 cm thick, sub- 
clavate to +equal, solid within and marbled white plus brown or black; surface 
near light pinkish cinnamon (6CS), alveolate-reticulate nearly to the base; 
reticulum concolorous with cap surface, concolorous with hymenophore above, 
grayish and compound with age. 

Basidiospores tinted flesh-pinkish in a light deposit, subfusoid and inequilateral 
to ellipsoid or subcylindric, sometimes constricted or bent near the middle, smooth 
and thin-walled, ochraceous in KOH, usually pale ochraceous or a few dextrinoid 
in Melzer’s, 10.5-15.4(20) x 4.2-5.6 um (mean Q=2.71). Basidia clavate, 4- 
sterigmate, hyaline, 24-35 x 9-12 4m. Hymenial cystidia conspicuous, thin-walled, 
with granular, homogeneous, or coagulated and refractive orange brown content in 
Melzer’s and uniform yellow brown or coagulated content in KOH, or more rarely 
hyaline and these more often near the pores, lageniform to fusoid ventricose to 
ventricose rostrate to subclavate rostrate, 42-63 x 8.5-12 um. Tube trama bilateral; 
hyphae diverging from a pale ochraceous central stratum, 3.5-10.5 um broad, with 


Fic. 20. Habit of Tylopilus obscurus (HOLOTYPE) x3/4. 


pia 


22 


10 zm . - } fs: ; 4 
aay si @ 
aN 
21 
5 um 
= oc theres 
aie 


Fics. 21-23. Microscopic features of Tylopilus obscurus (HOLOTYPE). 21. 
Basidiospores. 22. Elements of pileus surface. 23. Hymenial cystidia. 


the lateral stratum becoming gelatinized. Pileus surface a palisade trichodermium, 
becoming more tangled and less erect with age, not encrusted but hyaline or with 
brown to olive brown vacuolar pigment, with elements cylindric to broadly clavate, 
subfusoid to cystidiiform or rarely subisodiametric, 11-45 x 5-15 ym, with obtuse 
apices. Pileus trama hyaline, interwoven, 3.5-10.5 um broad. Stipe surface formed 
of long-cylindric, thin-walled, usually hyaline hyphae, 3.5-10.5 um broad, that occur 
between the ridges of the reticulum, the latter composed of clavate to subfusoid, 
lageniform caulocystidia with contents similar to those of the hymenium. Clamp 
connections absent. 


TYPE: COLOMBIA. DEPT. ANTIOQUIA: Municipio Guarne, Centro Experi- 
mental Piedras Blancas, +14 km E of Medellin, 2350 m, Cupressus, Eucalyptus, 
Pinus, and Quercus nearby, 11 Nov 1986, Halling 5008 (HOLOTYPE: HUA, 
ISOTYPE: NY). 


Tylopilus obscurus is not close to any of the neotropical Tylopili discussed by 
Singer et al. (1983). However, there are a few somber-colored Tylopili with reticu- 
lated stipes known from the Old World that are similar (viz. T. niger (Heinemann 
& Goossens) C. B. Wolfe, T. nigerrimus (Heim) Hongo & Endo and T. nigropur- 
pureus (Corner) Hongo). They differ from T. obscurus in important features: all 
have flesh that changes color when exposed, and some have shorter spores (T. 
nigropurpureus, T. niger), thick-walled cheilocystidia (7. niger), or olivaceous pig- 


112 


mentation (7. nigerrimus). An undescribed species from the eastern United States 
has black to gray tubes and pores, and tubes and flesh that change colors with ex- 
posure (Wolfe & Halling, in ed.). 

Exotic plantings of Pinus patula, Eucalyptus, and Cupressus, along with stands 
of native Q. humboldtii, were located within 50 m of the collecting site. Although 
not unquestionably associated with Quercus, T. obscurus was growing in an area 
once recently forested by Q. humboldtii and may be associated with the oak roots. 
Clearly this is circumstantial evidence for an association with oak, and deserves 
further investigation. 


ACKNOWLEDGEMENTS 


I am grateful to Gregory Mueller, Betty Strack, Clark Ovrebo, and Barbara 
Thiers for providing bolete collections while in Colombia. Also, the support from 
National Science Foundation grant #BSR-860024 made this study possible. The 
generosity, patience, and help of the staff and students, particularly Lucia 
Atehortua, Linda Albert de Escobar, Ricardo Callejas, Beatriz Echeverry and 
Patricia Velasquez in Medellin, Margarita Pulido in Popayan, and Gunther Buch 
at Finca Merenberg are likewise very much appreciated. The assistance of the 
curators, staff, and students at QCA and LIL enabled access to collecting sites. 
Esperanza Franco kindly assisted with the Spanish summary, and Donald and 
Susan Black provided graphic and technical expertise. Ron Petersen arranged for 
a loan of specimens from TENN. In addition, I thank Tim Baroni, Ernst Both, 
Egon Horak, and Rolf Singer for many helpful discussions on bolete identities. 
Naturally, I shall always be indebted to Harry Thiers for sharing his excitement 
about boletes and giving me the chance to wonder about them too. 


LITERATURE CITED 


Atkinson, G. F. 1901. Studies of American Fungi, Mushrooms Edible, Poisonous, Etc. 
Ed. 2. Andrus & Church, Ithaca. 

Coker, W. C. & A. H. Beers. 1943. The Boletaceae of North Carolina. Univ. North 
Carolina Press, Chapel Hill. 

Cotter, H. V. T. & O. K. Miller, Jr. 1985 (1986). Sclerotia of Boletinellus merulioides 
in nature. Mycologia 77: 927-931. 

Guzman-Davalos, L., G. Nieves, & G. Guzman. 1983. Hongos del estado de Jalisco, 
II. Especimenes depositados en el herbario ENCB, la. parte. Bol. Soc. Mex. Mic. 
18: 165-181. 

Halling, R. E. & C. L. Ovrebo. 1987a. A new species of Rozites from oak forests of 
Colombia, with notes on biogeography. Mycologia 79: 674-678. 

. 1987b. Some agarics and boletes of Colombian oak forests. [abstr.] MSA 

Newsletter 38(1): 27. 

Hongo, T. 1960. The Agaricales of Japan 1-2. Acta Phytotax. Geobot. 18: 97-112. 

Horak, E. 1977. New and rare boletes from Chile. Bol. Soc. Argent. Bot. 18: 97-109. 

. 1980. Supplementary remarks to Austroboletus (Corner) Wolfe (Boletaceae). 
Sydowia 33: 71-87. 

Kornerup, A., & J. H. Wanscher. 1978. Methuen Handbook of Colour. 3rd ed., Methuen 
Ltd., London. 

Ovrebo, C. L. 1983. New records of fleshy fungi from Venezuela. Mycotaxon 18: 355- 
356. 

Pegler, D. N. 1983. Agaric flora of the Lesser Antilles. Kew Bull. addit. ser. 9: 1-668. 

Singer, R. 1945. The Boletineae of Florida with notes on extralimital species I. The 
Strobilomycetaceae. Farlowia 2: 97-141. 


113 


. 1963. Oak mycorrhiza fungi in Colombia. Mycopath. et Mycol. Appl. 20: 239- 
2 


nN 


. 1964. Boletes and related groups in South America. Nova Hedwigia 7: 93-132. 
. 1970. Strobilomycetaceae (Basidiomycetes). F/. Neotropica Monogr. 5: 1-34. 
. 1973. Diagnoses fungorum novorum agaricalium III. Beith. Sydowia 7: 1-106. 
. 1986. The Agaricales in Modern Taxonomy. Ed. 4. Koeltz Sci. Books, Koenig- 
stein. 
, I. Araujo, & M. H. Ivory. 1983. The ectotrophically mycorrhizal fungi of the 
neotropical lowlands, especially central Amazonia. Beih. Nova Hedwigia 77: 1-352. 
, & A. P. L. Digilio. 1951 (1952). Prédromo de la flora agaricina Argentina. 
Lilloa 25: 5-461. 
. 1958. Las boletaceas Austrosudamericanas. Lilloa 28: 247-268. 
. 1960. Las boletaceas de Sudamerica tropical. Lilloa 30: 141-164. 
, & J. H. Morello. 1960. Ectotrophic forest tree mycorrhizae and forest com- 
munities. Ecology 41: 549-551. 
Smith, A. H. & H. D. Thiers. 1970. The Boletes of Michigan. Univ. of Michigan Press, 
Ann Arbor. 
Thiers, H. D. 1975. California Mushrooms: A field guide to the boletes. Hafner Press, 
New York. 
Weber, N. S. & A. H. Smith. 1985. A field guide to Southern Mushrooms. Univ. 
Michigan Press, Ann Arbor. 
Wolfe, C. B., Jr. 1979. Austroboletus and Tylopilus subg. Porphyrellus. Biblioth. Mycol. 
69: 1-148. 
, R. Singer, & R. Walsh. 1988. Notes on neotropical Austroboletus species. Myco- 
logia 80: 46-53. 


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MYCOTAXON 


Vol. XXXIV, No. 1, pp. 115-118 January 20, 1989 


CLADONIA THIERSIT: A NEW LICHEN FROM CALIFORNIA 


SAMUEL HAMMER! 


Department of Biology 
San Francisco State University 
San Francisco, CA 94132 


SUMMARY 


Cladonia thiersii is described from northern California. It 
is compared to C. subsubulata Nyl. and C. squamosa (Scop.) 
Hoffm. var. subsquamosa (Nyl. ex Leight.) Vain. The 
identity of a long-disputed specimen from California 
labeled C. santensis in the Tuckerman collections in FH is 
clarified. 


Cladonia sect. Perviae (Fr.) Matt. 
Cladonia thiersii sp. nov. Fig. 1 


Type: USA. California: Marin Co., Point Reyes National Seashore, 
Kehoe Beach, on humus, E facing, 50 m, 1988, Hammer 2286 
(Holotype: SFSU; Isotype FH); contains thamnolic acid. 


Thallus primarius squamulosus, persistens, crassus, sub- 
coralloides, partim infossus, ramosus infra substratum, ad 
12 mm longus et 5 mm latus. Podetia fragilia, pulvinos 
densos formans, cinereo-virescentia, ad 30 mm alta et 4.5 
mm lata, subcylindrica, parce dichotome ramosa, axillis 
perforatis vel clausis et sufflatis, cortex crassus vel 
rugulosus, verruculosus, squamulosus. Hymenia fusca; con- 
idiomata gelatinum hyalinam continentia. Acidum tham- 
nolicum continens. 


Primary thallus squamulose; primary squamules persistent, thick, 
forming extensive mats over surface of substratum, barely distinguish- 
able from one another, deeply established in substratum, 4-12 mm long, 
1-5 mm wide, irregularly dichotomously branched below surface of sub- 
stratum, deeply laciniate, subcoralloid to irregularly lobate above; edges 


1 Present address: Farlow Herbarium, Harvard University, Cambridge, MA 02138. 


116 


sinuate, to 0.3 mm thick, accessory lobules occasionally present; upper 
surface glaucescent green, sometimes with bluish tinge, subpruinose; 
ventral surface white, smooth to subfibrillose, upturned, esorediate or 
with granular soredia beneath upturned lobes. Podetia from upper sur- 
face of primary squamules, 4-30 mm tall, to 4.5 mm wide, subcylindri- 
cal, sometimes appearing inflated, forming scyphus-like openings at 
apices, sparingly to moderately branched; branches arising laterally or 
from margins of apical formations; cortex continuous to subcontinuous, 
sometimes disappearing, usually becoming quite thick above basal por- 
tions and appearing chinky-areolate to rugose to verruculose above; 
areoles or verruculae becoming elevated and appearing as closely ap- 
pressed squamules, or enlarging and imbricating, or becoming upturned, 
in some podetia becoming abundant, breaking up and appearing as 
corticated granules; apices enlarging, often abundantly covered with 
peltate squamules, gradually opening, exposing smooth, whitish to chest- 
nut brown interiors of podetia; closed apices enlarging and appearing 
inflated. Apothecia brown, borne in clusters around margins of apical 
openings, to 0.9 mm in diam; ascospores 8/ascus, hyaline, non-septate, 
oblong-ellipsoid, 6-16 x 3-6 wm; pycnidia brown, borne on margins; 
conidia hyaline, non-septate, straight to arcuate, 0.5-2 x 4-8.8 ym. 

Anatomy: Primary squamules 200-300 um thick; cortex 35-60 um 
thick, of tightly interwoven to adglutinated hyphal cells 7-10 wm in 
diam; algal layer 40-90 wm thick; medulla of densely packed, irregularly 
arranged, subcylindrical, sparingly branched hyphal cells 3-4.5 wm thick. 
Podetial wall 340-450 um thick; outer layer of closely agglutinated cells, 
40-60 wm thick; exterior medullary layer 250-300 um thick; internal 
cartilaginous tissue 50-90 wm thick. Hymenium 20-44 pm thick; sub- 
hymenium 28-48 wm thick; asci 25-28 wm thick, clavate; paraphyses 
subcylindrical unbranched, 2-3 septate, 28-39 x 2-3 um. 

Spot test reactions: K* bright yellow, persistent; KC* yellow; P* 
deep yellow to orange. 

Chemical constituents: Thamnolic acid with accessory substances F 
and G. 

Specimens examined: CALIFORNIA. Marin Co.: Hammer 2286 
(TYPE), Hammer 2302, 2395, 2407, 2411; Mendocino Co.: Hammer 1522 
(all SFSU); San Francisco Co.: Bolander 30 (FH). 

Cladonia thiersii is an intermediate species between C. subsubulata 
(=C. carassensis Vain.) and C. squamosa var. subsquamosa. TLC 
analyses demonstrate identical chemical constituents for this lichen and 
for the above named species. Cladonia thiersii is distinct on the basis of 
its morphology. Its podetia lack the wide lateral openings that are com- 
mon in C. subsubulata. The axils of C. thiersii may be open or closed, 
whereas they are always open in C. subsubulata. The primary thallus of 
C. thiersii is persistent, the squamules thick and deeply established in 
the substratum, unlike C. subsubulata or C. squamosa var. subsquamosa, 
whose squamules lack these characters. The podetia of C. thiersii be- 
come inflated when there is no apical opening, a characteristic I have 


117 


Scale bar =5mm 


Cladonia thiersii Hammer sp. nov. 


Fig. 1 


118 


not observed in other members of sect. Perviae. Many podetia are 
densely squamulose, approaching C. sguamosa var. subsquamosa. In C. 
thiersii, this state is often accompanied by a thick cortex, whereas in C. 
squamosa var. subsquamosa, densely squamulose podetia are usually 
decorticated. 

This lichen has a long and confused history in California. A 
specimen collected by Bolander in 1863 from Mission Dolores (Bolander 
30, FH) matches the type specimen of C. thiersii. Bolander sent his 
specimen to Tuckerman who called it C. santensis. The determination 
of this and several other specimens as C. santensis was considered by 
Nylander (Tuckerman, 1872) and later by Vainio (1887). Robbins 
(1927) summarized the controversy over Tuckerman’s treatment of C. 
santensis and stated, "Tuckerman conceded to C. santensis a wide lati- 
tude." Robbins cited seven distinct species included in Tuckerman’s 
concept of C. santensis, and determined Bolander 30 to be "a young 
state of C. crispata." This latter specimen, however, contains thamnolic 
rather than squamatic acid, which is found in C. crispata. Cladonia 
thiersii bears a superficial resemblance to C. crispata but differs in its 
chemistry and morphology. 


ACKNOWLEDGEMENTS 


I thank Dr. Teuvo Ahti for examining the type specimen of C. 
thiersii and for his helpful comments. I thank Drs. John W. Thomson 
and Richard C. Harris for their correspondence regarding C. santensis. 
I am grateful to Dr. Donald H. Pfister (FH) for the loan of specimens. 
For information on the correspondence between Edward Tuckerman and 
Henry Bolander, I thank Anna M. M. Reid. I am indebted to Ellen 
Thiers for her assistance with the Latin diagnosis. 


LITERATURE CITED 


Robbins, C. A. 1927. The identity of Cladonia beaumontii. Rhodora 29: 
133-138. 

Tuckerman, E. 1872. Genera lichenum: an arrangement of the North 
American lichens. E. Nelson, Amherst. 

Vainio, E. 1887. Monographia Cladoniarum universalis (1). Acta Soc. 
Fauna Flor. Fenn. 4: 1-510. 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 119-128 January 20, 1989 


STUDIES IN AGARICUS IV: 
NEW SPECIES FROM COLORADO 


RICHARD W. KERRIGAN 


Department of Biological Sciences 
University of California 
Santa Barbara, CA 93106 


Abstract 


The history of Agaricus study in Colorado 
is reviewed. Two new species of Agaricus from 
the Rocky Mountains of Colorado are described. 
They are: Agaricus amicosus, section Spissicaul- 
es, and Agaricus cuniculicola, section Arvenses. 


The recorded study of Agaricus in Colorado began 
shortly before the close of the nineteenth century, when E. 
Bethel sent some dried mushrooms which had been collected 
near the town of Craig to the New York State Botanist, C. 
H. Peck. This material formed the basis of A. tabularis 
Pk. (Peck, 1898). Peck (1905) subsequently renamed the 
Colorado fungus A. praerimosus Pk., addressing the conflict 
of priority posed by the earlier A. tabularis Pers. within 
the framework of the just-codified "rules of botanical 
nomenclature." 


E. B. Sterling, a New Jersey resident (and namesake of 
the 1902 species A. sterlingii Pk.) visited Denver in 1903, 
collected mushrooms in and around the city, and sent his 
dried specimens to Peck. Based on this material, Peck 
(1904) published four new species of Agaricus, which are 
given here with their later synonyms (Peck, 1905), if any: 
A. solidipes Pk., A. rutilescens Pk., A. sphaerosporus Pk. 
(= A. pilosporus Pk.), and A. cothurnatus Pk. (= A. chlamy- 


dopus Pk.). 


All five of these species were based on material col- 
lected in the plains to the east and west of the Rocky 
Mountains. With the exception of A. praerimosus, the 
"fairy rings" of which were studied extensively by Shantz 
and Piemeisel (1917), their natural history and their phe- 
notypic variation in the field are undocumented. Today 
they are but very poorly known. 


120 


Kauffman (1923) discussed two species of Agaricus (as 
Psalliota) found in the high Rockies. One was determined 
as A. semotus Fr., whereas the other formed the basis of a 
revision of A. rutilescens Pk. Smith (1940) reassessed 
Kauffman's treatment of A. rutilescens. Both of these 
reports are pertinent to A. amicosus (described below), and 
will be discussed in that context. 


No published reports on Agaricus of Colorado (other 
than type studies) have appeared since Kauffman's paper. 
There has been, however, a substantial amount of field 
research on the agarics of the Rockies, conducted by pro- 
fessional and amateur mycologists over the last several 
decades. Both A. H. Smith and H. D. Thiers, in particular, 
have spent portions of numerous "seasons" in this region, 
and both have related their impressions of the striking 
elements of the Rocky Mountain Agaricus mycota to me. My 
own observations from portions of three seasons spent in 
these mountains confirm and extend those of my recent 
predecessors. 


Materials, methods, and conventions: Formulae for 
most of the reagents mentioned below are given in Kerrigan 
(1985). Some new reagent formulations are: 


Syringaldazine: 10mg in 10 ml of 95% ethanol (a sat- 
urated solution) 
Ethanol: 95%, aqueous 


Color terms in single quotes are from Kornerup and 
Wanscher (1978). Other details of microscopy and terminol- 
ogy are given in Kerrigan (1985). Vouchers are deposited 
at. SFSU .or UCSB. 


Agaricus amicosus Kerrigan, sp. nov. Figs .dj920 


Pileus 7-18 cm latus, maturitate late convexus, 
fibrilloso-squamosus, albidus, fibrillis appressis, initio 
pallide fulvus, aetate rufobrunneis praeditus; contextus ex 
albo rufescens; odor mitis, dulcis. Stipes 6-12 cm longus, 
apice 1.1-2.5 (-3.2) cm et basi 3.0-4.5 cm latus, clavatus 
vel modice bulbosus, e farcto cavus; contextus albus modice 
rubescens; superficies alba, tactu rufula vel flavida; 
velum pendens, album, margine bruneolum; sporae (5.0-5.4-) 
6.0-6.7 ~(-7.2-8.0) x ((4.4—-) 4.8-5.3 ,(-6.2)  [tipeeee eee 
26.5-30.5 x 9-10.5 um, tetraspora; cheilocystidia 13.5-20 x 
6 Se21 2S um. Holotypus: R. W. Kerrigan 1258, Robber's 
Roost C.G., Grand Co., Colorado. 21 August 1983. SFSU; 
Isotypus NY. 


PILEUS 7-18 cm broad, at first hemispherical or semi- 
columnar to semi-cuboidal, becoming broadly to very broadly 
convex, the disc sometimes truncate or slightly depressed; 


121 


Figs. 1-2. Agaricus amicosus. Basidiocarps, x 
aes RWK 2258" (Holotype)':/" "2. “RWK' 1259. 


surface dry, variably fibrillose-squamose, the squamae ca. 
5 mm long by 2-20 mm broad, usually appressed, somewhat 
concentrically arranged, or the squamae obscure and the 
surface essentially fibrillose, colored pale buff ('6B3') 
when young and protected to medium deep brown ('6C5'-'6D5'- 
'6D6'-'6E7'-'7E7') in age, background white, becoming pink 
to reddish brown when incised or bruised; context white, 
becoming light reddish to slightly vinaceous ('9B5'-'10C6') 
when exposed, especially above stipe and below disc, moder- 
ately firm, 7-22 mm thick, odor fruity/spicy. 


iby 


LAMELLAE free, close (ca. '13 per cm at 1) /cm {rom 
stipe), to 9 (-13) mm broad, when young pallid pinkish, 
becoming deep dingy pinkish with a slightly paler margin, 
then bruising rose color, later becoming drab brown, ulti- 
mately dark blackish brown. 


STIPE 6-12 cm long x 1.1-2.5 (-3.2) cm above, 3.0-4.5 
cm at base, clavate or essentially equal above a moderate 
to indistinct bulb; interior white, lustrous, becoming 
rather uniformly medium red ('9B5'-'9B6'-'10C6'-'10C7') 
when sectioned longitudinally, stuffed-hollow, the cavity 
to 6 mm broad; surface glabrous, with fine striations, 
white, becoming bright reddish orange ('9D7'-'8C7') or 
sometimes bright orangish yellow ('5A7') below, when bruis- 
ed; basal 1-2.5 cm rooted in litter layer. 


VEILS forming a moderately thin, pendant, subapical 
(to supramedian) white annulus with a thick (to ca. 3 mm), 
grooved, often brownish (like pileus surface) UV margin, 
also often forming a white, appressed volval boot with an 
appressed or slightly flaring brownish margin, or rarely a 
series of brown-tipped rings or only an obscure velar rem- 
nant above the base. 


SPORES (5.0-5.4-) 6.0-6.7 (-7.2-8.0) x (4.4-) 4.8-5.3 
(-6.2) wm, mean size = 6.29 x 5.02 um (N=340, C=12), dark 
brown, broadly ellipsoid to ellipsoid, hilar appendix 
Slightly prominent, germ pore not evident. BASIDIA 26.5- 
30.5 x 9-10.5 um, clavate, predominately tetrasporic, ab- 
sent from lamellar margin; sterigmata 2.5-3 um long. CHEI- 
LOCYSTIDIA 13.5-30 x 6.5-11.5 wm, clavate to cylindro-clav- 
ate, or stocking-shaped, becoming brownish in age, then 
rehydrating poorly. PILEIPELLIS of straight, subparallel 
cylindrical elements of dimensions 33-43 (-78) x (4-) 8-9 
pm. 


CHEMICAL REACTIONS: KOH yellow or not on pileus sur- 
face, faintly to moderately strongly yellow on basal stipe 
context, slightly yellowish to negative elsewhere; Aniline 
x HNO, ("Schaffer reaction") red to orange (1/1 tests); 
ETOH orange on pileus surface (hence o-tolidine, syringald- 
azine, 1-napthol and 2,4-D orange in the absence of other 
color changes); o-tolidine blue (or orange; see ETOH) on 
pileus surface, blue on stipe base surface, violet else- 
where; l1-napthol bright pink to pinkish-orange everywhere 
(or with some purplish streaks in lower stipe context), to 
red on pileus surface (except disc sometimes purple); syr- 
ingaldazine orange (see ETOH). [Note: the extracellular 
laccase isozyme phenotype of isolate RWK 1438 is given in 
Kerrigan and Ross (in press).] 


Gregarious, often in rings or arcs, in deep litter 
under Picea and Abies, at elevations of 8500 to 10,000 
feet, in the Rocky Mountains of Colorado. August. Rou- 
tinely encountered. 


125 


Material studied: COLORADO. Boulder Co.: RewiW: 
Kerrigan 1438. Grand = Cons: RWKieL 25/5 onc OO pet 25956 (263° 
wa eee 23 1D, 81277. 2302;,° 1430 741433:,014385591436, «1446. 
San Miguel Co.: RWK 1448, 1452. Misc.: RWK 1447. 


Observations: In my experience, A. amicosus is the 
most frequently observed Agaricus in the high-elevation 
conifer forests of Colorado. Not only is it fairly common, 
but also the robust habit and red-staining context and 
surfaces of this brown-capped mushroom are distinctive. 
The impressions related by A. H. Smith and H. D. Thiers 
confirm that this striking species is quite familiar to 
those who study this habitat during the August rainy sea- 


son. Judging from conversations with several collectors, 
it appears that A. amicosus has often been taken for A. 
haemorrhoidarius Schulz. in Kalchbr. One difference be- 


tween these two species is the much narrower spore of A. 
haemorrhoidarius: 3-3.5 wm according to Cappelli (1984). 
Another way to distinguish A. amicosus from A. haemorrhoid- 
arius is by chemical spot-tests on the pileus surface. The 
KOH (yellow), ETOH (orange), and positive Schaffer's reac- 
tion of A. amicosus are not shared by A. haemorrhoidarius, 
and suggest affinities with section Spissicaules (Heinen. ) 
Kerrigan. The occasional tendency of the stipe cuticle to 
turn bright yellow-orange (e.g. RWK 1448) rather than 
bright red also appears to support this relationship. 
While perplexing to the uninitiated, this rather singular 
trait may ultimately shed light on the chemistry of these 
color reactions in the genus as a whole. 


Agaricus amicosus is quite similar to the European A. 
lanipes (Moll.) Moll., another member of section Spissi- 
caules. Agaricus amicosus differs from A. lanipes in habi- 
tat (high-elevation spruce-fir forests vs. lower-elevation 
oak forests), in chemistry (Schaffer's reaction positive 
vs. negative), and in spore size. The spores of A. lanipes 
are given by Moller (1950) as 5.5-6.5 x 3.75-4 um, and by 
Cappelli (1984) as 5.5-6.5 x 3.5-4.5 wm. The spores of A. 
amicosus are slightly longer and substantially broader; the 
lowest mean spore width value recorded for A. amicosus 
(collection RWK 1302), with a 95% confidence interval of 
+0.10 wm, is 0.37 wm larger than the most extreme spore 
width recorded by Cappelli, and is 0.87 wm broader than the 
broadest spores noted by Moller. Taking the mean spore 
size of A. lanipes, from Cappelli's data, as 6 x 4 um, that 
species has a mean spore volume of 50.3 um? (calculating 
the volume of the spore as (4/3)m(L/2) (W/2)“). Agaricus 
amicosus, with a mean observed spore size of 6.29 x 5.02 
um, haS a mean spore volume of 83.0 um, 65% greater than 
that of A. lanipes. I suggest that the difference in spore 
size between the two species is most indicative of a long 
evolutionary divergence that warrants recognition of A. 
amicosus at the species level. 


Kauffman's (1923) emended description of A. rutiles- 
cens is also evocative of A. amicosus. Kauffman described 
a frequently-encountered, large, brownish-capped, rufescent 


124 


mushroom under spruce and fir at both sites he studied 
(Leal, in Grand: Co., and Tolland, in Gilpin Co.) The 
points of discrepancy in his description, relative to A. 
amicosus as I have observed it, are a solid stipe [Kauff- 
man's emphasis], a thin, narrow annulus, and spores which 
were reported as 5-6 x 4-4.5 wpm. Smith later corrected 
Kauffman's spore measurements on the material to 5-6.5 x 4- 
5.5 um, in good agreement with my own observations (ironi- 
cally, Kauffman regarded Peck's spore measurements as being 
in error). The features of the annulus undoubtedly can be 
influenced by weather and environment, and in any event 
were described subjectively. Smith (1940) determined that 
Kauffman's material probably belonged to more than one 
species, since one collection had spores measuring 7-9 x 5- 
6 um (I have also found such a large-spored collection, RWK 
1449, among material of my own which was initially deter- 
mined as A. amicosus). Smith recommended adherence to the 
type concept of A. rutilescens. I have examined neither 
the type of A. rutilescens nor Kauffman's material, and 
prefer to follow Smith's advice. It seems likely that 
Kauffman may have had A. amicosus in hand, as least in 
part, when he redescribed A. rutilescens. 


The epithet amicosus, while suggestive of the gregari- 
ous habit of this mushroom, was actually chosen to cele- 
brate the many friendships which are a hallmark of Harry 
Thiers' career. 


Agaricus cuniculicola Kerrigan, sp. nov. Figs. 3% 


Pileus 5-11 cm latus, ex acute convexo planus, 
appresso-fibrillosus vel fibrilloso-squamulosus, pallide 
coloratus, fibrillis initio brunneis demum  fuscis 
praeditus; contextus albus, interdum aurantio-flavescens; 
odor amygdalinus; stipes 6-14 cm longus, apice 0.8-1.5 cm 
et basi 2-3 cm latus, bulbosus vel abrupte bulbosus, e 
farcto cavus; vela pendentia, interdum recurvo-pendentia, 
margine et pagina inferiore aurantia vel bubalina; sporae 
(5.0-5.4-) 6.2-6.9 (-7.6-8.0) x (4.2-) 4.6-5.6 (-6.2) pm; 
basidia (17.5-) 22.5-27.5 x 8-11 um, tetraspora; cheilocys- 
tidia~7-30.5 x 6.5-18.5 “um. Holotypus: R. W. Kerrigan 
1271, Willow Creek Pass summit, Grand Co., Colorado. 22 
August 1983. SFSU; Isotypus NY. 


PILEUS 5-11 cm broad at maturity, less than half the 
diameter of the stipe in primordia, acutely convex when 
young, ultimately becoming semi-plane with a slightly de- 
pressed or slightly umbonate disc; surface dry, innately 
appressed-fibrillose, becoming appressed-fibrillose or 
minutely fibrillose-squamulose (except the disc remaining 
entire), the squamules ca. 1-3 mm long x 1-2 mm broad, 
these hyphae initially medium brown (about '6C4'-'6C6!- 
'705'-'7D6') or paler, becoming darker in age (to about 
'7E8' or darker) on a whitish (to yellowish in age) back- 


ZS 


Fig. 3. Agaricus cuniculicola RWK 1271 (Holo- 
type). Basidiocarps, x 1/3. Note three basidi- 
ocarps that formed in small tunnels, with pilei 
just protruding above substrate surface. 


ground; context white, unchanging when young, becoming 
orangish in older material when exposed, 6-10 mm thick, 
odor of almonds. 


LAMELLAE free, close (ca. 16 per cm at 1 cm from 
stipe), 6-12 mm broad, initially pallid, then greyish-drab 
and faintly marginate or not, color unchanging when bruis- 
ed, finally dark blackish brown. 


STIPE 6-14 cm long x 0.8-1.5 cm broad above, 2-3 cm 
broad at base, bulbous to abruptly bulbous; interior white, 
unchanging in young material, or becoming orangish in older 
material, when exposed, lustrous, fibrous, stuffed-hollow 
to hollow, the cavity 3-4 mm broad; exterior white, lus- 
trous, becoming yellowish when bruised, finely striate 
above, covered below with numerous scattered, small, white, 
floccose velar squamules, these deciduous in age, lower 
surface lustrous, finely fibrous-striate; basal bulb cover- 
ed with coniferous litter. 


126 


VEILS forming a thin, pendant to pendant-recurved, 
sub-apical, white (or later orangish-yellow) annulus, with 
a finely striate upper surface and a scurfy, sometimes buff 
colored lower (UV) surface, the margin slightly thicker, 
somewhat eroded, orangish-buff; the UV also leaving scat- 
tered scurfy squamules over most of the lower stipe. 


SPORES (5.0-5.4-) 6.2-6.9 (-7.6-8.0) x (4.2-) 4.6-5.6 
(-6.2) um, mean size 6.58 x 5.05 wm (N=150, C=5), dark 
brown, broadly ellipsoid to ellipsoid (with many hyaline 
subglobose immature spores often also present), hylar ap- 
pendix not prominent, germ pore not evident. BASIDIA 
(17.5-) 22.5-27.5 x 8-11 um, clavate to cylindro-clavate, 
predominately tetrasporic, extending to edge of lamellar 
margin; sterigmata 3.5-4 um long. CHEILOCYSTIDIA 7-30.5 x 
6.5-18.5 um, globose, ovoid, ellipsoid, or various, catenu- 
late, abundant but restricted to the center of the lamellar 
margin. PILEIPELLIS of parallel to subparallel hyphae 
composed of straight, cylindrical elements of dimensions 
(15-) 33-47 (-56) x 6.5-9.5 pm. 


CHEMICAL REACTIONS: KOH yellow; o-tolidine blue ev- 
erywhere; 1-napthol purple at base of stipe and on cuticles 
(slowly so on context near base of stipe), pink elsewhere. 


Gregarious to scattered in litter of Picea and/or 
Abies, particularly in deep piles of decaying cone scales 
that result from the feeding activities of squirrels; the 
primordia sometimes form several cm beneath the surface, on 
the walls of short tunnels apparently excavated by small 
mammals; from ca. 8500-10,000 feet in the Rocky Mountains 
of Colorado. August. Fairly frequently encountered. 


Material studied: COLORADO. Boulder Co.: R. W. 
Kerrigan 1439. Grand Co.: RWK 1271, ©1272, 53276 pee eee 
307 W143 lee lLAS 24455 San Miguel Com: RWK 1453. 


Observations: Agaricus cuniculicola, when slender, 
most closely resembles A. smithii Kerrigan (= A. perrarus 
sensu A. H. Smith, 1940) of the.coastal Pacific Northwest, 
differing macroscopically only in having a brownish cuticu- 
lar pigment when immature, rather than the tawny orangish 
pigment of A. smithii. The overall aspect of these two 
species, including a narrowly convex pileus when young, a 
bulbous stipe base, and frequently a pendant-recurved an- 
nulus, suggest a close relationship between them. Other, 
more robust collections which resemble pale specimens of A. 
augustus Fr., or brownish forms of A. perobscurus Kerrigan, 
may also belong to A. cuniculicola. Such material that I 
have obtained has unfortunately been too immature for mi- 
croscopic analysis. If these variants are all conspecific, 
then A. cuniculicola has a fairly wide range of aspects. 
This makes field identification problematic, particularly 
since members of Agaricus section Arvenses Konrad & Mau- 
blanc are well represented in the high Rockies. 


n27 


Microscopically, the spores of A. cuniculicola are 
shorter than those of A. smithii. Mean values for spore 
length from eleven collections of A. smithii ranged from 
8.3 wm down to 7.5 wm, whereas mean spore lengths for col- 
lections of A. cuniculicola ranged from 6.3 to 6.8 um. The 
mean spore width of A. cuniculicola varied more between 
collections than did mean spore length, ranging from 4.6 t9 
eee PTE The difference in mean spore volumes (116.3 pum 
for A. smithii vs. 89.2 wm” for A. cuniculicola) is about 
30% of the mean spore volume of A. cuniculicola. 


The pink color formed by 1-napthol when placed on some 
of those areas of the context which turn blue with o-tolid- 
ine has not been recorded previously in other species; a 
blue tolidine reaction usually correlates with a purple 
napthol reaction. This unusual combination of reactions 
may have to do with the biochemistry of the orangish hue 
that develops in older context when exposed. 


Although not restricted to tunneled piles of cone- 
scales, in my experience A. cuniculicola fruits most abun- 
dantly there. Its association with this "naturally-manu- 
factured" feature of the high coniferous forests of the 
Rockies is striking enough to have been noted by other 
collectors, among them H. D. Thiers (pers. comm.). This 
association is reflected in the species epithet. 


ACKNOWLEDGMENTS 


I would like to express my gratitude to those who 
interested me with their tales of the Colorado mountain 
species of Agaricus (including my mentor and his mentor), 
and to those who guided my way in the field, offered shel- 
ter and company, and made it financially feasible to work 


in their state. Marilyn Shaw, Linnea Gillman, Duane H. 
Mitchel, and many other members of the Colorado Mycological 
Society, Emmanuel Salzman and Fungophile, Inc., deserve 


special mention. I thank David Farr for checking the pri- 
ority of the proposed names, Robert Patterson for the co- 
vert loan of material housed at SFSU, and Roy Halling and 
Wayne Ferren for their comments on the manuscript. I am 
particularly grateful to Ellen Thiers for rendering the 
taxonomic diagnoses into Latin. 


LITERATURE CITED 


Cappelli, A. 1984. Agaricus. Biella Giovanna, Saronno. 
560 pp. 

Kauffman, C. H. 1923. The mycological flora of the higher 
Rockies of Colorado. Pap. Michigan Acad. Sci. 1:101- 
150 + 5 pl. 

Kerrigan, R. W. 1985. Studies in Agaricus III: New spe- 
cies from California. Mycotaxon 22:419-434. 


128 


, and Ross, I. K. In press (1988). Extracellular 


laccases: Biochemical markers for Agaricus systemat- 
ics. Mycologia 80(5). 


Kornerup, A., and J. H. Wanscher. 1978. 


of Color. 3rd Ed. Eyre Methuen, 
Moller, F. H. 1950. 


Friesia 4:1-60. 


Peck, C. H. 1898. New species of fungi. Bull. Torrey. 
BoG: IClubn2oe32 i328. ‘ 


. 1904. New species of fungi. Bull. Torrey Bot. 
Club ese DTA ais 


. 1905. Report of the state botanist 1904. New 
York State Mus. Bull. 94:1-58. 
Shantz, H. L., and R. L. Piemeisel. 1917. Fungus fairy 
rings in eastern Colorado and their effect on vegeta- 
tion. «id .sAgric.URes . (11: 191-245) (+3050 Fay 


Smith, A. H. 1940. Studies in the genus Agaricus. Pap. 
Michigan Acad .\Scilsi25: 107-138 Onl 


Methuen Handbook 
London. 
Danish Psalliota species. [Part 1] 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 129-131 January 20, 1989 


A NEW, LIGNICOLOUS SPECIES OF ENTOLOMA 
(Entolomataceae, Agaricales) FROM CALIFORNIA. 


David L. Largent 
Department of Biological Sciences 
Humboldt State University, Arcata, California, 95521 


SUMMARY 


A new species of Entoloma (sensu stricto), £. lignicola, that grows on 
conifer wood is described from northern California. 


INTRODUCTION 


In November of 1986, at least 200 basidiomes of a species of Hntoloma (sensu 
stricto) were found growing on a conifer log in northern California. A collection of 
nearly 50 basidiomes in all stages of development was made. Because of the following 
set of features, this species was considered distinct and unusual: woody substrate, 
consistently convex, tan pileus with a margin extending beyond the lamellae, close, 
narrow lamellae, curved, slender stipe, obscurely angled, isodiametric, small basid- 
iospores, and a narrow cutis as the pileipellis. A careful review of the pertinent 
literature resulted in the conclusion that this species had not been previously 
described. 

Therefore, it is with pleasure that I dedicate this species, E’ntoloma lignicola, to 
Harry D. Thiers. 


Entoloma lignicola Largent, sp. nov. 


Pileus 20-45 (-55) mm latus, convexus vel lato-convexus, glaber, tannus, haud 
hygrophanous, margine decurvato, integro, non-striato. Lamellae adnatae, albae, 
angustatae, confertae. Stipes 3-5 (-6) mm crassus, 25-60 mm longus, glaber, sub- 
albus. Odor et sapor farinceus. Sporae 5.1-6.1 x 4.1-5.1 um, isodiametricae, obscuro- 
angulatae. Hymenial cystidia desunt. Pileipellis cutiformis, cellulis terminalibus 
cylindro-clavaticis. Stipitipellis cutiformis; caulocystidia desunt. Pigmentum in 
vacuolis hypharum. Fibulae in hyphis pileipellis, pileal trama, lamellar trama, 
stipitipellis, stipe trama, et basi basidiorum adsunt. Basidiocarpi dispersi vel gre- 
garii ad lignum in sylvis sub Lithocarpum, Arbutum, et Pseudotsuga . Holotypus 
(DLL 8747) in HSU conservatus. 

Pileus: 20-45 (-55) mm in diameter, 10-20 mm in height, consistently broadly 
convex and without an umbo, glabrous, dull, dry, with an incurved to decurved, 
non-translucent-striate margin that consistently extends beyond the lamellae by 
1-3.5 mm, not hygrophanous, colored at first an off-white to a pale buff (6,A-B,3 to 
6,B,3), soon becoming and remaining a uniform tan (6,C-D,5); pileal flesh concolor- 
ous with the surface, 1.5-2.5 mm thick. Lamellae: adnexed to adnate at first becom- 
ing typically adnate, narrow to somewhat moderately broad (8-7 mm high, 10-830 mm 
long), typically close to rarely subdistant, colored white at first, with a smooth 
margin concolorous with the surface. Stipe: 3-5 mm thick at the apex, 3-6 mm thick 


130 


at the base, 25-60 mm long, most often equal but at times tapered towards the apex, 
usually entirely glabrous but at times with a slight pruinose apex, decidedly curved 
towards the light and away from the ground, thus negatively geotrophic and 
positively phototrophic, colored an off-white, obscurely longitudinally striate, witha 
sparse to moderate, whitish basal tomentum. Odor and taste: farinaceous, especially 
when the flesh of the pileus is crushed. 

Spores 5.1-6.1 x 4.1-5.1 um, average length 5.44 wm, average width 4.85 um, Q 
(average length/width) 1.12, length/width 1.0-1.38, length minus width 0-1.5 um, 
isodiametric, multiangular with the angles typically obscure, even in outline and 
with no protrusions or outgrowths. Basidia clavate, 25.4-33.5 x 6.1-8.1 um, length/ 
width 3.3-5.0, 4-spored. Hymenial cystidia absent. Subhymenium in cross section a 
tightly interwoven layer, up to 24.6 wm wide, composed of small, slender hyphae, 
3.7-4.9 um wide. Lamellar trama in cross section composed of subparallel hyphae, 
71.4-172.4 x 12.3-17.3 um. Pileal trama in radial section composed of interwoven 
hyphae above the stipe and hyphae parallel to the pileal surface above the lameliae; 
hyphae 36.9 -135.5 x 7.4-29.6 um, with the broader hyphae more abundant beneath 
the pileipellis. Pileipellis in radial section a cutis, 2-6 hyphae thick, with the hyphae 
more slender than, and thus differentiated from, the underlying hyphae of the pileal 
trama, composed of slightly entangled hyphae over the stipe and parallel hyphae 
elsewhere; terminal cells cylindro-clavate to somewhat broadly clavate, 24.6-69.0 x 
6.2-9.9 um, length/width 3.3-11.2. Stipitipellis composed of a very narrow layer of 
loosely interwoven hyphae overlying a layer of parallel hyphae; caulocystidia absent. 
Pigmentation apparently vacuolar, distinctly not incrusting or membranal. Lactif- 
ers quite rare, more common in the trama of the stipe and the lamellae than in the 
pileus trama. Clamp connections extremely rare on the hyphae of the pileipellis, 
moderately abundant on the tramal hyphae of the pileus and stipe, abundant on the 
tramal hyphae of the lamellae and at the base of the basidium, rare on the hyphae of 
the stipitipellis. 

Distribution and Habitat: Rare, collected only once; scattered to gregarious on a 
slightly decomposed conifer log in a Tanbark Oak (Lithocarpus densiflorus (Hook. & 
Arn.)Rehd.), Douglas fir (Pseudotsuga menziesii (Mirbel)Franco), Madrone (Arbutus 
menziesii Pursh.) forest; Humboldt County, California; mid-November. 

Material Studied: CALIFORNIA: Humboldt Co.: DLL 8747 (HOLOTYPE: HSU; 
16 November 1986, 5.0 miles east from the junction of Big Hill Road and State 
Highway 96) 

An unsuccessful attempt at identifying this species was made utilizing the follow- 
ing pertinent literature: Horak (1978, 1978, 1980), Kiihner & Romagnesi (1953), 
Noordeloos (1981, 1987), and Romagnesi & Gilles (1979). 

On the basis of its small basidiomes, convex, tan pileus, narrow, adnate lamellae, 
rather slender stipe, farinaceous odor, small, obscurely angular, isodiametric basid- 
iospores, cutis as the pileipellis, and lignicolous habit, Hntoloma lignicola appears to 
be a unique species. 

Because of its non-hygrophanous, non striate, glabrous pileus, it appears related to 
Entoloma niphoides Romagn. ex. Noordel., HE. eulividum Noordel., FE. sinuatum 
(Bull.:Fr)Kumm., and FE. prunulordes (Fr.:Fr.)Quél. However, all of these species 
have much larger spores, a humicolous habit, a pileus that is up to 100-125mm broad, 
and a T’richoloma-like stature. 

Because of its basidiome colors, convex pileus with thin flesh, and farinaceous 
odor, Entoloma lignicola appears also to be related to FE. rhodopolium and 
E. speculum (Fr.:Fr.)Kummer. However these species are distinct because of their 
hygrophanous pileus with a striate margin, humicolous habit, and larger basidio- 
spores that are distinctly angular. 

On the basis of its small, obscurely angular basidiospores, Entoloma lignicola 


131 


could be placed in Entoloma section Turfosa (Romagn.)Noordel. However, all species 
in this section possess a hygrophanous pileus with a striate margin, grow in the 
humus, and possess basidiomes that are much darker in color. 

Colors from Kornerup and Wanscher (1978) are quoted by page (first number), 
horizontal row (letter), and vertical column (last number). 


LITERATURE CITED 


Horak, E. 1973. Fungi Agaricini Novazelandiae Entoloma (Fr.) and related genera. 
In Beih. Nova Hedwigia 43, J. Cramer, Vaduz. 

———— . 1978. Entoloma in South America. Sydowia 28: 171-236. “1974/1975” 

———— . 1980. Entoloma (Agaricales) in Indomalaya and Australasia. In Beih. 
Nova Hedwidgia 65, J. Cramer, Vaduz. 

Kornerup A, & J. H. Wanscher. 1978. Methuen Handbook of Colour. Methuen, 
London. 

Kuhner, R. & H. Romagnesi. 1953. Flore analytique des Champignons superieurs- 
Paris. 

Noordeloos, M. E. 1981. Entoloma subgenera Entoloma and Allocybe in the Nether- 
lands and adjacent regions with a reconnaissance of their remaining taxa in 
Europe. Persoonia 11: 1538-256. 

———— . 1987, Entoloma (Agaricales) in Europe. In Beih. Nova Hedwigia, 91, J. 
Cramer, Berlin-Stuttgart. 

Romagnesi, H. & G. Gilles. 1979. Les Rhodophylles des foréts c6tiéres du Gabon et de 
lat Cote d’Ivoire avec une introduction générale sur la taxinomie due genre. In 
Beih. Nova Hedwigia 59, J. Cramer, Vaduz. 


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Vol. XXXIV, No. 1, pp. 133-151 January 20, 1989 


NOTES ON THE GENUS PROTUBERA (PHALLALES) 


David Malloch 
Department of Botany 
University of Toronto 

Toronto, Ontario, Canada, M5S l1Al 


ABSTRACT 


The genus Protubera includes species of Phallales that 
are essentially gasteromycetous. Basidiomata are composed 
of a thin peridium enclosing a gelatinous matrix. Fertile 
glebal plates or chambers extend inward from the peridial 
eveeeeaua weventually §fITT all’ ior part of: the: interior. 


There are seven described species of Protubera: jag 
Rerroavagumer., borealis, PP. \brunnea, | PP. clathroidea,.’ P. 
fametcensis,  P. maracuja’ (the type species), anaes uP 


nipponica. An eighth species, P. sabulonensis is described 
as new to accommodate collections from primary sand dunes 
on Sable Island, Nova Scotia,. Canada. All of the species 
Peeve eliy characterized, including» notes on, type. and 
other collections, illustrated in part and differentiated 
by a dichotomous key. 


INTRODUCTION 


TamerosO  Altred Méller, while’ collecting fungi near 
Blumenau, Santa Catarina, BY az i, encountered a 
gasteromycetous form that closely resembled an unexpanded 
"egg: OL one of the Phallales. This fungus was common in 
the region and fruited abundantly throughout the year. He 
was able to study it closely throughout its development and 
determined that it remained in the closed gasteromycetous 
Permeetiont cChrough “to: its ‘final: dissolution. After his 
return to Germany he determined this fungus to represent a 
Rewemeenus, Of | Phallales' and described’ it as 'Protubera 
maracuja Moller (Moller, LS 9 Sic: In that paper 
developmental stages and mature basidiomata were well 
illustrated, and these were later reproduced by Fischer 
¢190.0))-. Since then this fungus has been collected a number 
Ditames and .reported from ‘various parts: of, the) world; 
Furtado & Dring (1967) accumulated and presented most of 
these records, although they accepted as correctly 
identified only material from southern Brazil. 


As described by Médller, basidiomata of P. maracuja 
first appear as brown, spherical to elongated structures. 
They are’ attached to the soil by stout rhizomorphs and 
consist of a compact’ peridial layer surrounding a 
gelatinous center. During maturation, hyphal plates or 
lobes grow into the central gelatinized tissue from the 


134 


peridium, and give rise to internal labyrinthine cavities 
lined with basidia. Cross sections of mature basidiomata 
reveal a number of radially arranged fertile lobes 
connected by narrow strands to the inner surface of the 
peridium and extending halfway or more into the center. 


Lloyd ° (1920) ~*‘received ‘additional  materiaieowers 
mMaracuja from Rev. Rick in Brazil,-who stated @thaGii gs 
not rare, but added nothing to Médller's description. In 
the same paper Lloyd reported on a species collected in 
Stellenbosch, ' South Africa by Miss’ A. V. 3 Duehieeeand 
described it as a new species, Pp. africana slovar This 
species was said to differ from P. maracuja in having a 
single glebal mass filling the center of the basidiomata 
mather Tthan Pradiactingws plates: The glebal mass _ was 
described as "compact and minutely porous to the eye" and 
similar to that of Rhizopogon species. 


A third species, P. borealis Imai was proposed by Imai 
(1936) to accommodate a collection from soil in Cryptomeria 
faponica '(L:). DD. Don. forests (near, Sapporo none eee 
Japan. P. borealis was reported’ to be” similarieco 
maracuja and to differ from it in having slightly @iarce, 
spores and a nearly white peridiun. 


Kobayasi (1938) described the new species P. nipponica 
Y. Kob. for a collection from (soil in the @vaegeuoee. 
Kunitati, Honshu, Japan, said to be similar to P. maracuja 
and P. borealis but differing in having a holloweeenee. 
caused by the disintegration of the gelatinous matrix. The 
glebal plates were described as greatly elongated and 
extending nearly to the center of the basidiomata. 


Imai & Kawamura (1958) obtained material of P. 
maracuja from Dr. Rick in Brazil and compared i cCeagi ec 


with the two Japanese species. Based on the nature of the 
peridial and glebal tissues these authors proposed two new 
genera for the Japanese species; Protuberella for P, 


borealis and Kobayasia for P. nipponica. Interpretation of 
this paper is rather difficult, but the following@heavaga: 
reflect Imai & Kawamura's views: 


Ai 2, Peridium composed of three distinct 
Lay:GT Sitaie abs tte ny pee er ie We Protubera maracuja 
be Peridium composed of two layers .... 2 
an Basidiomata hollow at the 
COT RCTs sts inathriats cr its alain Kobayasia nipponica 
4.3 Basidiomata gelatinous at the 
CENCE wy at an un tis omar as ths one ns Protuberella borealis 


A fifth species of Protubera, P. clathroidegmpeenae 
was proposed by Dring (1964) for a form collected in sandy 
places in ;Togo’ and alsio ‘one reported as P. maraciw aa 
Pakistan, by Ahmad X1952)°° In PP. clathroidea themiguenan 
plates’ ~appear “to be “united into (4a; single “masa This 
species was reported to be similar to P. africana Ulovomuee 


135 


to have similarly broad basidiospores. Unlike P. africana, 
P. clathroidea has basidiomata with a sterile gelatinous 
center. 


Two further species, P. brunnea (Zeller) Zeller and P. 
jJamaicensis (Murrill) Zeller must be included in this 
discussion. Protubera jamaicensis was originally described 
Bemechem scype species “of sProtophallus Murrill (Murrill, 
£9:1.0).. Protophallus was apparently described without 
knowledge of Protubera and was maintained as distinct from 
it by Zeller (1939) and Imai & Kawamura (1958) apparently 
on the basis of a simple rather than branched columella 
(central gelatinous area). Zeller (1948) later rejected 
the distinction and transferred _ the two species of 
Protophallus to Protubera, a decision accepted by Dring 
(1964). Protubera jamaicensis and P. brunnea were not 
originally compared with species of Protubera and are thus 
not easily distinguished from them on the basis of their 


diagnoses. Protubera jamaicensis was reporteds) <to Yebe 
Piaeweteri zed. by a viscid “peridium and) P. ,brunnea® by 


broadly ellipsoidal basidiospores (5-6 X 2.5-3 pm). 


During October, 1982, I spent three weeks studying 
fungi on Sable Island, Nova Scotia. ThiUsvaistande: Ustea 
narrow bar of sand about 200 km east of the Nova Scotia 
mainland that supports a surprisingly varied vegetation. 
theweroredunes on «the north ‘side of the island iare 
relatively unstable and support a vegetation dominated by 


Ammophila breviligulata Fern. (Graminae) and Lathyrus 


maritimus (L.) Bigel. (Leguminosae). LALs *thapitatrewas 
generally rather poor for fungi, but I nevertheless found 
abundant basidiomata of a species of Protubera. Although 


descriptions were available for all of the described 
species of Protubera it quickly became apparent’ that 
species concepts were confused or not well defined and that 
identification of my unknown specimen was nearly 
impossible. Because of this, it became necessary to 
examine representative material of some of the _ species 
similar to the Nova Scotia collections and to arrive at a 
more satisfactory concept of existing species. The 
following notes are based on these examinations. Colors 
designated in upper case are according to the Methuen 
Handbook of Colour (Kornerup & Wanscher, 1978). 


THE SPECIES OF PROTUBERA 


PROTUBERA AFRICANA Lloyd, Mycological Notes 64: 987. 1920. 


Figs, 412 3% 
BASIDIOMATA (Fig. 1) more or less spherical, smooth, 
pale brown; in cross’ section revealing four separate 
tissues, 1) a thin brown peridium, 2) a soft gelatinous 
Davyexrmeabouc. 3: mm thick; ) 3)'' a ‘thin *internal membrane 
surrounding 4) a central glebal mass. RHIZOMORPHS stout, 


extending into the soil. ODOR not specified. 


136 


OUTER PERIDIAL LAYER compact, 52-96 pm thick, 
brownish, composed of tightly interwoven non-gelatinized 
hyphae, never enclosing druse-like crystalline masses. 


GELATINOUS AREAS 600-700 um thick, composed of interwoven, 
septate, gelatinous hyphae, containing numerous oleiferous 
hyphae, sometimes containing a tissue similar to the 
peridium that is oriented parallel to the peridium. GLEBAL 
MASS filling the center of the basidiomata, reticulate- 
poroid as in species of Rhizopogon, bearing basidial 
hymenia within the chambers, connected to the peridium by 
narrow bands of tissue. BASIDIA (Fig. 2) 8-spored, short- 
steriigemate Mw2h33 7 Oy An2e7e2arpm. BASIDIOSPORES (Fig. 3) 
ellipsoidal, smooth, with a conspicuous basal apiculus, 
greenish olive in mass, 4.6-6.2 (mean = 5.1 + 0.44) X 2.0- 
2.8 (ep = 294 b7001L7) cpm, with imeanyD/dt="2..09) taGy nae 


MATERIAL EXAMINED: SOUTH AFRICA: Stellenbosch, 
Papagaaisberg, in damp clayey soil, Duthie 233 (Lloyd 
22143)),°'20.V1I.1919, HOLOTYPE (BPI); possibly ‘samemlocatice 
as holotype, Duthie 243 (Lloyd 22144) (BPI). 


The macroscopic part of the description is based on 
that of Lloyd (1920) who presumably had a description from 


Miss Duthie, the collector. The holotype packet also 
contains a colored drawing of fresh material, redrawn in 
Figs Soule This drawing indicates that there are strands 


connecting the peridium to the gleba as in other species, 
although .<“Murrill): states) (‘that’) no.4 such: \strtuctume sms uc 
present. I was unable to observe these in the herbarium 
specimens, but the parallel layer that resembled to me a 
second embedded peridium may in fact have been one of these 


connections. This is the only species of Protubera where 
the glebal masses appear to be combined into a single 
ecentralivunicy As Lloyd pointed out, the resemblance of the 


gleba to that of Rhizopogon species is striking. 


PROTUBERA BOREALIS (Imai) Imai & Kawamura, Si. Rep. 
Yokohama Nat JeuUniwv Sesecta wie 7 340 LO5.& 
= Protuberella borealis Imai, Bot. Mag., Tokyo 50:223. 
1936. 


Figs. 4-6. 


BASIDIOMATA (Fig. 4) arising from a rhizomorph, uneven 
to wrinkled. or clearly  sulcate, (grayish white §to spate 
sordid, 20-50 mm in diameter; in cross section there appear 
to be four tissues, 1) a thin pale compact peridium, 2) a 
rigid, gelatinous peridium-like layer 1-3 mm thick, 3) a 
thin gelatinous "filling", and 4) centripetally arranged 
glebal plates extending inward from the peridium. 
RHIZOMORPHS white, gelatinous within, branched. ODOR not 
recorded. 


PERIDIUM compact, 230-360 um thick, pale, composed of 
interwoven, occasionally clamped hyphae with globose to 
oblong or cylindrical cells 8-20 um in diameter, without 
crystalline masses. GELATINOUS AREAS composed of 


137 


interwoven gelatinous hyphae up to 9 um in diameter, rather 
firm near the peridium and glebal masses and highly 


gelatinous and watery elsewhere, sometimes containing 
capitate hyphae 12-18 um in diameter. GLEBAL MASSES 
Pectactune “inward from’) the’ peridium’, shorcnmand often 
cerebriform. BAS LOLA PACE Tp )wec Lavaten Con its bevulariy 


Cylindrical, 8-spored, clamped at base, 14-20 X 2.8-4.9 ‘um. 
Rao vesr ORES) (Fig: (6) "ellipsoidal, “smooth, (with'a) broad 
beanaieapiculus, ‘Grayish ‘yellow’ to” Olive ‘brown’ (4CD3) "in 
mass in rehydrated material, 3.8-4.7 (mean = 4.2 + .020) X 
Pees (mean = 91.8 °° 0.13) ‘pm, with mean 'D/d '=' 2.36 + 
Oe205"3 


MATERIAL EXAMINED: JAPAN: Tottori Pref., Tottori-City, 


Kokoge, gregarious in stand of Quercus myrsinaefolia Blume 
in woods dominated by Pinus densiflora Siebold & Zaccarini, 
Nagasawa, 16.VII.1980 (TMI-6948) (TRTC). 


The material examined here seems close enough to the 
Gtaeenan cescription of P. borealis to) cause’ Vittle” doubt 


Seancovuts identity. The holotype was collected in forests 
of Cryptomeria japonica (L.) D. Don. near Sapporo, Ishikari 
Prov., Hokkaido, Japan. The one point of disagreement is 


ine spore, size: that given by Imai’ (1936) was 3.75-4.5 XK 1»2- 
oem which is broader than’ the’ spores of ithe’ Tottori 


material. The species is distinguished from other 
Procupera, species by its’ combination of short’ glebal 
masses, soft or almost watery gelatinous material and 


relatively narrow basidiospores. 


PROTUBERA BRUNNEA (Zeller) Zeller, Mycologia 40: 644. 
1948. 
# Protophallus brunneus Zeller, Mycologia 31: 28. 
1939. 


BASIDIOMATA arising from a rhizomorph, = spherical, 
smooth but somewhat felty, white, becoming sordid, 
reticulate with a few lines, 15-35 mm in diameter; in cross 
section there appear to be four separate tissues, 1) a thin 
tough peridium, 2) a rigid gelatinous peridium-like layer 
feomemme chick’, 93) ta ‘thin (gelatinous "filling, and | 4) 
centripetally arranged glebal plates extending inward from 


the peridium. RHIZOMORPHS white, apparently unbranched, 
with a peridium-like cartilaginous sheath and a gelatinous 
center, extending nearly into the center of the 


basidiomata. ODOR not specified. 


PEREDIUM* compact’, about 175-250 ym) thick,|) composed of 


interwoven and anastomosing non-gelatinous hyphae, with 
crystalline masses not reported. GELATINOUS AREAS composed 
of interwoven anastomosing, highly gelatinous’ hyphae. 


GLEBAL MASSES arising as ingrowths of the peridium, less 
gelatinous than the surrounding tissues, extending nearly 
to the center of the basidiomata, giving rise to a regular 
hymenium of basidia internally. BASIDIA elongate-clavate, 
4-6-spored. BASIDIOSPORES broadly ellipsoidal, smooth, 
Drown 1n mass’; 9-6 Xo2.5=3. pm. 


138 


MATERIAL EXAMINED: none. The above description has 
been taken from that of Zeller (1939) and transformed into 
the standard format of this paper. 


Protubera brunnea appears to be distinctive in having 
the rhizomorph extending well into the center of the 


basidioma and in having large broad basidiospores. The 
type (now at FH) was collected by Roland Thaxter in a park, 
Buenos Aires, Argentina in February and March 1906. The 
type locality is not soi distant \from that of |) Byemagecuae 
that, it) does not! storce: comparison \withiie. However, the 
two diagnostic features mentioned above, as well as the 
white, filamentous peridium OLGaars brunnea, seem to 


distinguish them. 
PROTUBERA CLAITHROIDEA Dring,.Mycol..(Pap. CMIC Ee 9siae 1964. 
Figs. 7-8. 


BASIDIOMATA (Fig. 7) arising from. a rhizoid, = ovoi1gere 
subglobose, smooth, covered with an incomplete network of 
shallow, grooves,, dirty white, 15 X 15-30 X 60 \mmjMinwervogs 
section there appear to be five separate tissues, 1) a thin 
white membranous peridium, 2) a rigid gelatinous peridiunm- 
like layer , 3) a thin white membranous layer enclosing the 
gleba, 4) fused glebal masses attached to the peridium by 
centripetally extended "sutures" and 5) a central 
gelatinous, 'iitling RHIZOMORPHS white, occasionally 
branched. ODOR when ripe of ethyl acetate (apples). 


BASIDIOSPORES (Fig. 8) ellipsoidal to ovoid, smooth, 
hyaline, with a-broad' sterigmatalk scar, (3.5) ()4=6506@- 730 
C3 = Roe = G15 50 Sie my, 


MATERIAL EXAMINED: none. Description adapted from 
thatyofeDrine ss G1L964)¢ 


It is: difficult ‘to compare’ P. ‘clathroidea with ae. 
described species of Protubera because of the lack of 


microscopic detail reported in its original diagnosis yeine 
holotype was collected in sandy places in Togo. As noted 
above, the species also seems to occur in Pakistan. Aside 


from. the, habitat, it. is. distinguished /by its Wiconwiwea. 
peridial masses surrounding a central sterile gelatinous 
zone (called a columella by Dring) and by = tte ogoed 
basidiospores. 


PROTUBERA JAMAICENSIS (Murrill) Zeller, Mycologia 40:644, 


P9485 

= Protophallus jamaicensis Murrill,  Mycologiagggesn 
1910 
| a BY ag 8) ee Bee En 


BASIDIOMATA (Fig. 9) arising from a Trhigesorens 
spherical, smooth, viscid, avellaneous, becoming white at 


139 


maturity, 20-40 mm in diameter; in cross section there 
appear?’ to be three separate tissues, 1) a thin’ white 
compact peridium, 2) a gelatinous inner layer that forms a 
peridium-like tissue 1-1.5 mm thick and extends inward to 
the center, and 3) centripetally arranged glebal plates 
extending inward from the peridium almost to the center. 
RHIZOMORPHS arising from the substrate, white, unbranched, 
extending to the center of the basidiomata. ODOR lacking. 


PERIDIUM compact, 200-280 wm thick, composed of rather 
densely interwoven hyaline hyphae in a gelatinous matrix. 
GELATINOUS AREAS composed of interwoven gelatinous hyphae. 
GLEBAL MASSES arising as ingrowths of the _ peridiun, 
filamentous, extending nearly to the center of the 
basidioma, olivaceous to deep brown, giving rise to a 
tortuous hymenium of basidia internally. BASIDIA (Fig. 10) 
8-spored, clavate to subcylindrical. BASIDIOSPORES (Fig. 
11) ellipsoidal, smooth, with a broad basal apiculus, brown 
iemageews:, 1-4:,4° (méan = °3 16 240025) °Xi1.3-1.9 Cmean = 1.6 
Pome opm, with meantD/d =—\2.26 £°0.223. 


MATERIAL EXAMINED: JAMAICA: near Cinchona, in shaded 
soil rich in humus on the bank of the Clyde River, Murrill, 
Jveieugo9. HOLOTYPE (NY); “same general locality" as the 
holotype (Coker & Couch, 1928), on very rotten wood, Coker, 
1900 (NY). 


The above description was based upon the reports of 
Maen to 10:), CGoker: & Couch °(1928) and’ Zeller (1939) and 
augmented by my own examination of the holotype and Coker's 


material from the same locality. A colored illustration 
made by a Miss Taylor, who was studying at Cinchona at the 
time, is contained in the holotype packet and was also 
used, A second illustration, now torn in two, illustrates 
a cross sectional view of a basidioma (redrawn in Fig. 9) 
and of one glebal mass. It is not clear whether this was 
done by Miss Taylor or Dr. Murrill. The herbarium material 


at the New York Botanical Garden is not in good condition: 
the holotype seems to be nothing more than shell-like 
peridial fragments devoid of gleba or spores and the Coker 
material is half of a basidioma dried down from liquid 
preservative. There is a permanent microscope slide mount 
of P. jamaicensis at NY that is not clearly from either of 
the two localities. Like the Coker collection, however, it 
came originally from the herbarium of S. M. Zeller and may 
be part of the material that Coker had collected and later 
Sséne. ctorhim. On the other hand, a note in the holotype 
packet states that part of the collection had been 
preserved in liquid and it may be that this is what Zeller 
had used for his slide. This slide is the only material on 
which 8-spored basidia can be seen clearly. 


The elongated glebal plates and basidiospore 
gamensitonss suggest that. it* ‘is’ close {to /P:. nipponica: in 
penoweelrte is ‘difficult torditscover ‘characters’ useful’ 'in 
separating the two species. The only feature that may 
serve this purpose is the peridium, which is described as 


140 


viscid in P. jamaicensis. Coker & Couch (1928), however, 
make no mention of this character at all. Another feature 
that may prove, diagnostic)’ is the extenslom® Gfimwene 
rhizomorph of P. jamaicensis well into the center of the 
basidioma in the manner of a columella: this does not seem 


COMOccUr Nim fo nipponicar 


PROTUBERA MARACUJA Méller in Schimper, Bot. Mittheil. aus 
den’ Tropen 7ivb0 mo. Ts957 


Figs.) L225) 


BASIDIOMATA (Fig. (12) arising ‘from \ ayy rhtzomoroun: 
smooth, pale leather brown when young, becoming wrinkled 
and darker at maturity, up to 50 mm in diameter; in cross 
section there appear to be four separate tissues, 1) a thin 
brown compact peridium, 2) a rigid gelatinous peridium-like 
layer 2-3 mm thick, 3) a thin gelatinous "filling" and 4) 
centripetally arranged glebal plates growing inward from 
the peridium. RHIZOMORPHS extending into the humus, nearly 
white to pale reddish below, up to 3 mm in diameter, 
becoming branched and extending out into a network up to l 
meter long. ODOR of ripe passion fruit when mature. 


OUTER RERIDIAL LAYER» (Fig. ').13)) compact, * 17Se22500um 
thick, appearing to be pseudoparenchymatous in cross 
section, orange-brown, with cells ca. 10-60 pm in diameter, 
with inner cells apparently enclosing large druse-like 
masses of irregular colorless crystals (Fig. 14). 
GELATINOUS AREAS composed of septate, interwoven, rather 
contorted and apparently clampless hyphae 1.8-3.7 pm in 
diameter, with hyphae less gelatinized in the area adjacent 
to the peridium and the glebal masses. GLEBAL MASSES 
arising as ingrowths from the peridial layer, filamentous, 
less gelatinized than the surrounding tissues, extending 
one-third to two-thirds of the way to the center, giving 


rise to a regular hymenium of basidia internally. BASIDIA 
8-spored, short-sterigmate. BASIDIOSPORES GEL gs doe} 
ellipsoidal, smooth, with a broad basal apiculus, Brown 
(7E6-8))\\in \mass); '}2.9-3.8° (mean) =, 343° 250020) ee 


(mean )=— 1,70 +) 0513): pm, with mean D/d'’= 1.9532 00 Pia 


MATERIAL EXAMINED: BRAZIL, Santa Catarina, Blumenau, 


BU Sis y ila 2 Protubera africana. Et gicgek's Basidioma, 
redrawn from colored illustration in holotype packet. Fig. 
Pay Basidia. bd EY AN Go Basidiospores. Figs. 4-6. 
Protubera borealis. Bl pig!) ney) Basidioma, redrawn from 
Kobayasi (1938) : Fig’. ou Basidia. Fig. 6. 
Basidiospores. Figs, /-8.. Protubera clathroidea vm higaeas 
Basidioma, redrawn from Dring (1964). Fig. 8. 
Basidiospores, redrawn from Dring (1964). Figseeaocseue 
Protubera jamaicensis. Bigpied 9: Basidioma, redrawn from 
illustration in holotype packet. Bier gu Basidiun, 
probably collapsed. Fig. Hy Basidiospores. The 


basidiomata are X 1; the remainder X 1500. 


141 


es 
os 
Ces 
Ca 


142 


in soil in woods, Méller, 1890. HOLOTYPE,:(B) 4. SADDETIOCNAL 
MATERIAL EXAMINED BUT NOT INCLUDED IN THE ABOVE 
DESCRIPTION: BRAZIL: Sao Paulo: Sao Paulo, Reserva 
Biolégica, Parque Estadual das Fontes do Ipiranga, Sales & 
Vital, §8.V.1961. (SP:.62010); Milanez & Altimay Lez yee 
(SP | 62033); Zago. & Furtado, 23-XI1.1960 C3 eee 
Guidiccl, OA Bs De (Cio ys, CS Pen eS. cools Serra da \Gantarert ra, 
Picada’ das Jaboticabeiras, Zampieri, 13.1.1966 (Sree tGg7 


Serra da Cantareira, Servico Florestal do Estado, 
Hernandes, T3966 CS Pees LOS Sr Serra da Cantareira, 
Johnston, Ta ie 66 (SP 91084); Ubatuba, Estacgao 
Experimental do Instituto agronomico de Campinas, 
Hernandes, Lilo 66% Paranapiacaba, Esta¢gao Bioldégica do 
alto. da, Serra, Hernandes; 11.1966% Sao Paulo veepepenve 
Bioldégica, Parque Estadual das Fontes do Ipiranga, 
SKVOLrtzov., Wee EN Byes bee Ode All, of (the above’ aren ea teetie 


Instituto de Botanica, Sao.Paulox(SP). 


The above description may be somewhat narrow as it 
includes only Méller's and my observations on) themtype, 
Furtado and Dring (1967), however, presented a description 
of this species based on a number of additional collections 
ErQm Sp razgiin In that paper Furtado and Dring stated that 
their material confirmed "in every respect" the description 
of Moéller, However, Méller described his material as 
having brown basidiomata while those of Furtado and Dring 
were white or grayish. Since both authors seem to have 
observed numerous specimens it is unlikely that differences 
in gage» account ifors thick evanvarion: Méller's original 
material came from temperate forests in Santa Catarina 
while that of Furtado and Dring was from tropical ?rain 
forests further north; perhaps humidity or habitat has some 
effect on color. I have examined 10 of the specimens cited 
by Furtado and Dring (1967) and find that there is also-—a 
difference in spore size from the holotype. In Fo) maracuya 
the spores never exceeded 3.8 pm long while those of the 
Furtado and Dring material averaged 4.1 wm and could be as 
Lonigu.as) <bean. Spore diameters of the two groups were 
approximately the same and thus the D/d ratios were 1.7-2.4 
in ,the holotype and 1.9-2.8 “in (the Furtado Mandir. 
collections... The report of .4-spored basidia) in’ their 
material by Furtado and Dring seems to be in error; I found 
only basidia with 8 spores. 


The most striking feature of P. maraculya ieee 
presence of numerous masses of crystals in )theseinge: 
peridial, celiseyC hie wi AD.. These are very large and easily 
seen with lower power microscope objectives. I have not 
observed these in any other species and believe that they 
may be diagnostic. In this respect it is interesting that 
all of the Furtado and Dring specimens examined also 
contained these crystals. Méller (1895) supposed these to 
be calcium oxalate but did not offer evidence other than 
to) mention tnatwcheyordissol ved. (in eC: The supposedly 
pseudoparenchymatous peridium in P. maracuja may also be 
diagnostic. All of the material of this speciésiithauea 
examined, including the "controversial" collections, had 


143 


this feature. LEpiSevervyadaglirtoult Glo previverthiisy ti seuwe 
and it is quite possible that it is not truly 
pseudoparenchymatous. However, that is the way it appears 


in herbarium specimens and it is quite unlike the clearly 
hyphal peridia of other species. 


PROTUBERA NIPPONICA Y. Kobayasi in Nakai & Honda, Novae Fl. 
Menatee 2 S25), LIS Se 


= Kobayasia nipponica (Kobayasi) Imai & Kawamura, Sci. 
Repe, LOkKoOnama Nat. .Univ /> Sect... dl o/ s5-. 1958. 


Figs. 16-18. 
BASILDIOMATA \(Fig, 16). .arising from a -rhizomorph, 


spherical to somewhat irregular in shape, somewhat wrinkled 
or cracked above, sulcate toward the base, white, 25-50 mm 


in diameter; in cross section there appear to be three 
femaratcewtissues,, 1),,ay-thin compact peridium,« 52) ai irigid 
gelatinous peridium-like layer 1eO-ipsoe Lamm Path ick. 3) 
centripetally arranged glebal plates extending inward from 
thes peridium; the center is hollow. RHIZOMORPHS white, 
simple or branched. ODOR not reported. 


PERIDIUM compact, 199.0239 Anpm Jethicks composed of 
interwoven and occasionally clamped hyphae. GELATINOUS 
AREAS composed of loosely interwoven gelatinous hyphae 2-5 
pam in diameter, very firm and rubbery near the peridium and 
the glebal plates and softer to almost watery elsewhere. 
GLEBAL MASSES arising as ingrowths of the peridiun, 
filamentous, densely packed, extending nearly to the center 
of the basidiomata, giving rise to a regular hymenium of 


Desidisamunternally. BAS IDUAM  CHRigin ols) S8-sporeds) short: 
sterigmate, irregularly clavate to cylindrical or 
ventricose, clamped at base, 22-41 xX HL FET ter 5 -4)'O) pm. 


Basidiospores (Fig. 18) narrowly ellipsoidal, smooth, with 
a broad basal apiculus, Olive to Olive brown (3-4EF4) in 
mace nao. Oo. (mean, = 4.0) 5.0.24) X).1.5-253 (mean = 1:8 + 
Orpto) gums with mean’ D/d. = 2.23 + 0.211; 


MATERIAL EXAMINED: JAPAN: Tottori Pref., Tottori-City, 
Kokoge, In woods of Pinus densiflora, Nagasawa, 21.1X.1976 
Gieiew2o0); Shimane Pref., Ochi-gun, Mizuho-cho, in forest 
of Pinus densiflora, Maeda, 9.X.1980 (TMI-6925). 


The holotype was collected on the ground in 
unspecified woods in Kunitati (a western suburb of Tokyo), 
Honshu, Japan (Kobayasi, 1938). Additional collections 
were described by Imai & Kawamura (1958) from Kubokawa- 
Caewewenochi., Pref...) Shikoku ‘and, Shingi.,, Omi» Prov., (in Shiga 
Pret.)% Honshu, Japan. It has been very well illustrated 
by Kobayasi (1938), Imai & Kawamura (1958) and Imazeki & 
Hongo (1965). 


As discussed above, the most difficult species to 
temineuish from oP. nipponica is 2. jamaiecensis,..\) The only 
other species of the genus reported from Japan is P. 
borealis, which has shorter glebal plates. Kobayasi also 


144 


distinguished the two species on the basis of the hollow 
center of P. nipponica basidiomata and the fact that the 
glebal chambers (|of P: nipponica are green) an eiiguce 
preservative while those of P. borealis are leather brown. 


PROTUBERA SABULONENSIS Malloch, sp. nov. 


Rigs toto 


Basidiomata ad obovoidea irregulariter 
tuberoidea, ad levis araeolata, ad basem frequenter 
SUlcatia Alba, 2704 OAD eT LO a) erate Peridium 16-65 pm 
crassum, luteobrunneum, a hyphae intertextis compositum. 
Trama gelatinosa, firma, a hyphae gelatinosae intertextae 
composita. Gleba ad peridium in laminae irregularae 
centripetaliter radiata, 3-15 X L=5 mm. Basidia 
octospora, ad subclavata cylindracea, 16-25 X 3.9-5.0 pm. 
Basidiosporae leniter obovoideae, laevae, ad basim 


truncatae’, flavobrunneae, 3,8-5.0'X 1.0-276 im, 


HOLOTYPUS: CANADA: Nova Scotia, Sable Island, 600 m NW 
of West Light, in loci aranosi a Ammophila breviligulata 
associata, Malloch .30.9'°82/4, 30 September sy igo. In 
herbarium cryptogamicum University of Toronto conservata. 


BASIDIOMATA (Fi gis: D9), 23-24) arising from a 
rhizomorph, obovoid to somewhat irregular in shape, smooth 
to finely cracked above, often coarsely sulcate below or 
grooved when in contact with grass culms, dry, rubbery in 
texture,’ initially white, with surface easily “diseororing 
when touched to reveal colors near Grayish yellow to Olive 
brown (4CF4) to Yellow brown (5E5), with some) tincs eo. 
Brownish orange (7C4) in age, 21-64 mm high, 10-55 mm 
broad; in cross section there appear to be three separate 
tissues, 1) a compact white peridium, 2) a firm gelatinous 


tissue filling the’ bulk) /of") the, *basid loner arndir ay 
centripetally arranged glebal plates extending in from the 
peridiunm. RHIZOMORPHS white, usually branched, with 


peridium and gelatinous matrices continuous with those of 
the basidiomata, not extending. into the interior of the 


basidiomata, extending into the substrate. ODOR of 
Bidig cue 2a ioue Protubera maracuja. Fi girl ian Basidioma, 
redrawn from Méller (1895). op aN od eS Cross section of 
peridium showing outer hyphal layer, "pseudoparenchymatous" 
layer and inner gelatinous’ matrix; note crystalline 
inclusions in the inner cells of the "pseudoparenchyma". 
Ea gee Lan Crystalline inclusion from peridial Celisteeae. 
oe Basidiospores, with upper group of 8 as attached to 
basidiun. Figs). (16- U8". Protubera nipponica. Figirr.on 
Basidioma, redrawn from Kobayasi (1938). Fig. Lr, 
Basidia. ‘Fig. 18. Basidiospores. Figs, 19-227 (Peegupere 
sabulonensis. Pagel Oh Basidioma. Ri gree o Cystidium- 
like element from gelatinous matrix. Fil gat rein Basidia. 


Fig. 22. Basidiospores.: The basidiomata are ‘X 17" Figeeia 
is X 75 and the remainder X 1500. 


145 


146 


overripe bananas when old. 


PERIDIUM 16-65 pm thick, yellowish brown, composed of 
parallel to somewhat interwoven, apparently clampless 
hyphae 1.7-6.0 um in diameter, never enclosing crystalline 
masses. GELATINOUS AREAS hard and rather agar-like, nearly 
colorless to Yellowish grey (4B2) or Olive brown (4DE4) 
depending upon the stage of glebal maturity, composed of 
hyphae that are interwoven, very loosely arranged, 
clampless to occasionally clamped, hyaline, and 1.3-9.0 pm 
in diameter, often containing oleiferous hyphae that are 
sometimes swollen at the ends to form cystidium-like 
elements (Fig. 20). GLEBAL MASSES arising from "plates" of 
nearly parallel and hardly gelatinous hyphae similar to 
those of the peridium that extend inwardly at right angles 
to. >the. surtace, cerebriforn, composed of labyrinthine 
cavities lined with basidia, rather irregular in size and 
shape but approximately 3-15 X 1-5 mn. BASIDIA (Fig. 21) 


8-spored, borne in a parallel hymenium, ventricose to 
subclavate, clamped at base, with short sterigmata, 16-25 X 
3 9 Oe OL ems BASIDIOSPORES (Fig. 22) narrowly obovoid, 


smooth, truncate at the broad sterigmatal scar, inamyloid, 
Grayish yellow (4BC3) to Yellowish brown (5EF6) in mass 
when fresh, 3.8-5.0 (mean = 4.4 + 0.22). X 1.9 - 2963 Cneane= 
2.3 £.0.14) pm, with mean) D/d = 1.940250 [face 


MATERIAL EXAMINED: CANADA: Nova Scotia, Sable Island, 
600 /m. NW) of West , Light (UIM, Grid@ene, 21TTU56/686), 
scattered to gregarious in sand in dense marram community 
on side of sand dune in association with Ammophila 
breviligulata Fern., Lathyrus maritimus (L.) Bigel. , and 
Solidago semprevirens ‘L., Malloch 30.9.82/4, 300i xai9675 
HOLOTYPE (TIRTC);° 2.5: km: E ‘of West Light (UI Mga 
21TTU599687), in sand in small dune depression in dense 
Marram community, Malloch 1.10.82/4, 1.X.1982 (TRTC). 


P. sabulonensis is unique among species of Protubera 
in having a peridium less than 70 wpm thick, a very dense 
and agar-like gelatinous matrix, quite broad basidiospores 
(mean D/d < 2.0) and habitat on oceanic sand dunes. eS a 
should not be confused easily with any other species. 


The habitat of P. sabulonensis is unlike that of other 
species, with the possible exception of P. clathroidea. 
Both localities were behind the crests of large barrier 
sand dunes densely covered with Ammophila breviligulata 
("Dense Marram Community" of Freedman, Catling & Lucas, 
1982). Although well covered with grasses, these sites are 
all very dry and have no soil other than sand. All of the 
basidiomata were tightly adpressed to the base of the 
Ammophila culms and were deeply grooved along the area of 


Figs. LO) a. Protubera sabulonensis. Fig. re 
Basidiomata in sand beside culms of Ammophila 
brevilicguldta, |) Fie. 26. Basidiomata., Both Xela 


147 


148 


contact. The rhizomorphs extended into the sand and 
appeared to be attached to dead and rotting culms and 
rhizomes. Of all species of Protubera, P. sabulonensis 


appears to be the most densely and uniformly gelatinous. 

It is likely that this extremely gelatinous nature of tne 
basidiomata protects them against drying in the very xeric 
sites they occupy. 


Most other species of Protubera occur in humus in 


forests; however, there is no indication that they are 
ectomycorrhizal. I. know .of no confirmed “records. 
ectomycorrhiza formation by members of the Phallales. In 
addition, the occurrence of species of Protubera is 
frequently in association with trees known to form only 
vesicular-arbuscular mycorrhizae, such as Cryptomeria 
japonica. The roots of Ammophila breviligulata in the 


communities where P. sabulonensis occurred were only weakly 
mycorrhizal, and this appeared to be brought about by the 
usual aseptate VAM fungi. 


The relationship of species of Protubera to the 
Phallales has not been disputed since the first species was 
described. In spite, of this little has been) said wabout 
odors. Most species of Protubera have been described as 
odorless. It» is significant, \though’, that th Memes 
entirely so: old basidiomata of P sabulonensis smell of 
overripe bananas, those of P. clathroidea smell of apples 
(Dring, 1964), while’ P. maracuja smells of |cipejipase van 
Peuicy The fruity odor of P. maracuja was striking enough 
to Méller’ that ‘he, used it as the basis of hiemispecgc 
epithet; ie. maracuja, the local Brazilian name for passion 
Prue, It seems possible that these species attract and 
are dispersed by frugivorous - insects. Dring (1964) 
reported that insects attacked older basidiomata of P. 
clathroidea and it is likely that this happens in other 
species but has passed unnoticed. The fact that some 
species have been reported as odorless may only be because 
the collectors smelled young basidiomata; if the fungus is 
attracting dispersers with these odors it will =noteiieewy 
do so before the spores are fully mature. It is tempting 
to consider Protubera species as dispersed by frugivorous 
insects while species of other Phallales attract feeders on 


less attractive substances. It is interesting nase. 
regard, however, thats PilAe We) PUsakrmit1o5s5e in ‘their 
discussion, of Phallus impudicus L.. ex Pers), stave ewe 
Mutinus' caninus’: (Huds.) Fr, has >the’ same odovmegeueee 
impudicus and that while it is strong and offensive in the 
latter, M. caninus has_ been said by some mushroom 
collectors to have the odor of fruit. This suggests, at 


least, that the same substances may play a part in the 
odors of all Phallales. 


KEY TO THE SPECIES OF PROTUBERA 
There is much yet to be learned about the taxonomy of 


Protubera. Most collections have not been adequately 
studied in the fresh state and we thus lack information 


149 


about elongation of glebal plates and changes in the 


gelatinous consistency of tramal tissues. Some of the 
species outlined above may be based on isolated stages in 
development and may be synonymous with one another. In 


Bpecew or this “uncertainty  1t is’ possible to ~make some 
decisions about taxonomic relationships among the species 
and to identify, cautiously, an unknown specimen. The 
following key is a step in this direction. 


4 LF Peridium appearing to be pseudoparenchymatous 
in cross section, often enclosing druse-like 
crystalline masses in the innermost parts 
TI ae acc tars adi slo. c.tupes Cite hele ts P. maracuja 

1% Peridium usually clearly composed of inter- 
woven hyphal elements, not enclosing large 
SUNT TLG WM SS OOS 0 alt el tie tation ar els tate ince Fo hedee coe Lod ty 2 


25. Glebal masses apparently confluent, not 
appearing as centripetally arranged 
MeL OW COU SITOL Che AD Gti LUM irre Co uetel., 3 

hs Glebal masses distinct and extending 
inward from the peridium as individual 
Dera eSMOL MASS OS isi c es phe spies! it sini vies fe, Sih 6 4 


ze Glebal mass hollow; basidiospores broader than 
RMT ete esairs sea vest eid + oe a )8) lie) P. clathroidea 

oN Glebal mass solid, reticulate or poroid, 
resembling that of Rhizopogon species; basid- 
iospores narrower than 3pm .. P. africana 


4. Pas vGLospoOres! Ow KYi2) 2-3 OPE Wl ce lens 
ee Ot Tee vane he Pee Gs Sadia opin G0 P. brunnea 
4. Ree VOS DOL CSM SMALL Ge Meal csiers cision eeu cytes aks 5 


he Glebal masses elongated and extending nearly 
to the center of the basidiomata; gelatinous 
tissues soft to watery, sometimes absent at 
RUMI TLCGLA Pe aro %s ew aia a tet escheat sis dnl Sah Uae catia” big el Sits Unl  ads s 6 

ot Glebal masses rarely extending more than 
halfway toward the center of the basidiomata; 
gelatinous tissues firm and present at the 
Perera mate sDOre Ma Cur Lily ou tee jais gr éceduen ekele okie. « 7 


be Surface slightly viscid; rhizomorph 
extending well into the basidioma ..... 


PES eT aah aie, Sheds fo utes: Citi P. jamaicensis 


6. Surface dry; rhizomorph not extending 
into the basidioma .~.... Po ynipponica 
Ti Peridium > 200 pm thick; gelatinous tissues 


firm but not hard and agar-like; mean 
basidiospore length/width ratio > 2.0 ...... 
MT eRe Sel te cae tas ca lid te Rens P. borealis 

Tis Peridium < 100 pm thick; gelatinous tissues 
hard and agar-like; mean basidiospore 
Paneer /Widtiuractor<).27,077.',)...00P. sapulonensis 


150 


ACKNOWLEDGEMENTS 


I thank Dr. E. Nagasawa for sending me representative 
material of the two Japanese specimens and accompanying me 
to the collection locality of one of them and Dr gee 
for translating some Japanese texts and clarifying for me 
the citations of Japanese localities. I am grateful to Dr. 
A. R. Lock of the Canadian Wildlife Service for arranging 
my visit to Sable IsYand and, to Mr. R. B. Taylovsst ore 
generous help before, during and after that visit. The 
help of the curators of the New York Botanical Garden, the 
National Mycological Herbarium at Beltsville, Maryland, the 
Botanischer Garten und Botanisches Museum, Berlin-Dahlem 
and the ,Instituto de Botanica at Sao Paulo) is@igrare tn 
acknowledged. This work was supported by an Operating 
Grant from the Natural Sciences and Engineering Research 
Council of Canada. 


LITERATURE CITED 
Ahmad, S. 1952. Gasteromycetes of West Pakistan. Punjab 


Univ. Press, Lahore. 
Coker aWi A Gwen ae Nae GOMehe LOZ. The gasteromycetes of 


the eastern United States and Canada. Univ-3En 
Carolina Press, Chapel Hill. 

Drineie Dee 1964. Gasteromycetes of west tropical 

ALY icar, Mycol) Papers CMI 98. 

Piscney yas 1900. Hymenogastrineae in Engler, A. & K. 
Prantl. (eds.)Die nattirlichen Pflanzenfamilien I(1**): 
9 OF 

Freedman, -Bii, Poy MM. Catlingar dé) 2 alucass 9.68 The 


vegetation -of Sable Island ,in Taylor, Ri Beegcedua 
Terrain management and biological studies on Sable 
Vs Vandi @ bo.oiLe. Sable Island Environmental Advisory 
Committee, Halifax. 

Furtadow a) 2 es4 CDi ee Mone cel tT) ae LOG The rediscovery of 
Protubera maracuja, with additional descriptive notes. 
TY ansee). Bra CoM Yy COLCRSS O Coain oO aU Uie) Uae 


IMiivets hays SNA 193.65: Symbolae ad floram mycologicam Asiae 
onientalis (Bot. Mag TokyOn 25 Onur) Oac 2 cee 
Imai, S. & A. Kawamura. 1958. ‘On the Japanese species of 


Protubera. Sci. Rep. Yokohama Univ., Ser? 2 eee 
imazeki, Ro &) sl Hongo. 1965" Coloured illustrations of 
fungi of Japan II. Hoikusha Publ. Co., Osaka. 
Kobayasie.) 5y% 1:93 8. Hymenogastrineae et Phallineae in 
Nakai, T. & M. Honda (eds.). Novae Flora Japan, 2. 
KOrnerup uA ea ne ewans Chex. . 1978. Methuen Handbook of 
colour. Eyre Methuen, London. 
LDL6OyVd) auCn aac T9207 The genus Protubera. Mycological 
Writings 6(64): 986-987. 
M6 Llern ola. 1895. Brasilische Pilzblumen in Schimper, A. 


F. W. (ed.). Botanische Mittheilungen aus den Tropen, 
VOU c. Gustav Fischer Verl., Jena. 

Murrill, W. As’ 1910. . A new phalloid genus. Mycolomgamae 
Zioini2 OF, 

Pilate aA poet Orme ea ke L585 Mushrooms. Spring Books, 


London. 


151 


Bevery *o).. M. LOS ON New and noteworthy gasteromycetes. 
my co Loria %3 12332". 
Pemeet so. M. 1948. Notes on certain gasteromycetes, 


including two new orders. Mycologia 40: 639-668. 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 153-179 January 20, 1989 


NOTES ON CLAVARIADELPHUS. III. 
NEW AND NOTEWORTHY SPECIES FROM NORTH AMERICA ! 


Andrew S. Methven 
Botany Department 
Eastern Illinois University 
Charleston, IL 61920 


SUMMARY 


Clavariadelphus americanus is proposed as an 
autonomous taxon based on Clavariadelphus pistillaris var. 
americanus. It is redescribed and a neotype specimen 
designated. Four new species from North America, Cc. 


caespitosus, C. flavidus, C. occidentalis, and Cc. 
pallido-incarnatus are described and illustrated. 


Although Clavariadelphus Donk (1933) has been treated 
in several publications (Corner, 1950, 1970; Wells & 
Kempton, 1968; Smith, 1971; Petersen, 1972; Petersen et 
al., 1974), delimitation of infrageneric taxa in North 
America has proven difficult in many cases. A 
considerable part of the problem is the result of poor 
documentation of subtle variations in basidiocarp color, 
size, shape, growth habit, and micromorphological 
structures. Additionally, there has been a tendency for 
systematists to categorize collections of Clavariadelphus 
into three widely, often misapplied, epithets, C. ligula 
Csoueerft.s) Fr.) Donk, Cy pistillaris)(L.: Fr.) Donkj\tand 
C. truncatus (Quel.) Donk. Lack of type specimens for 
certain taxa, including the three listed above, has 
further compounded the confusion surrounding the taxonomic 
and nomenclatural limits of infrageneric taxa in North 
America. 


lohis work represents a portion of a dissertation 
submitted to the Graduate School of The University of 
Tennessee, Knoxville, in partial fulfillment of the 
requirements for the degree of Doctor of Philosophy. 


154 


During an examination of material collected by the 
author, and dried specimens from cooperating herbaria, 
some new taxa were recognized and are described herein. 
Additionally, C. americanus, recognized as an autonomous 
taxon based on C. pistillaris var. americanus Corner, is 
redescribed and a neotype specimen designated. 

Microscopic features are described from sections 
revived in 95% ethanol and water, then mounted in 3% KOH, 
Melzer's reagent or cotton blue. Measurements were made 
in 3% KOH. Colors terms followed by alphanumeric 
designations are from Kornerup and Wanscher (1978), those 
from Ridgway (1912) are bracketed by quotation marks. 
Recipes for macrochemicals may be found in Marr and Stuntz 
(1973), Marr (1979), and Singer (1986), and abbreviations 
are as follows: KOH = 3% potassium hydroxide; FeC1, = 102 
ferric chloride; EtOH = 95% ethanol; Melzer's = Melzer's 
reagent; PCR = p-cresol; PYR = pyrogallol; SYR = 
syringaldazine; TYR = l-tyrosine; GUA = guaiac tincture; 
PHN = phenol; NH40H = 10% ammonium hydroxides; NAP = 
1-naphthol; ANO = aniline oil. 

Abbreviations used in citation of basidiospore size 
are as follows: L_= mean length; W_ = mean width; 

E = length/width ratio; E. = mean of E. Herbarium 
acronyms are from Holmgren et al. (1981) except for the 
following individuals who lent specimens from their 
private herbaria: Wm. Bridge Cooke -- Herbarium of Wn. 
Bridge Cooke (= WBC); P. E. Kempton -- Wells and Kempton 
Herbarium (= WK); Clark L. Ovrebo -- Herbarium of Clark L. 
Ovrebo (= CLO). 


Clavariadelphus americanus (Corner) Methven, stat. nov. 
Figs. 1,3-5 


Clavariadelphus pistillaris var. americanus Corner. 
1950 a ¢Anns | Bot. wMemt ili 692" 


Clavaria pistillaris var. americana (Corner) 
Leathers. 1955. The genus Clavaria Fries in Michigan. 
Ph.D. dissertation (ined.), University of Michigan, Ann 
Arbor. 344 pp. 


NEOTYPE (des. mihi.): TENN: United States, 
Massachusetts, Berkshire Co., White Oaks Road, vicinity of 
Williamstown, 16 Aug 1986, legit A. S. Methven No. 4724 
(TENN 47403). 


153 


Figs. 1-2. 1. Basidiocarps of Clavariadelphus 
americanus (Methven 4715). x1/2. 2. Basidiocarps of 
Clavariadelphus caespitosus (TENN 34744). x 3/4. 


156 


BASIDIOCARP 3-15 cm high, inserted 1-4 cm in the 
substrate, 3-12 mm diam basally, up to 30 mm diam 
apically, simple, initially subcylindric to narrowly 
clavate, enlarged upward in age, then clavate to broadly 
clavate; mycelial hyphae interwoven or aggregated into 
rhizomorphic strands up to 1 mm diam, off-white to pallid; 
base terete, smooth, pruinose, white or pallid where 
covered, otherwise yellowish white (4A2) to orange white 
(5A2), “light ochraceous-buff," "pale ochraceous-buff," 
"pale pinkish buff"; hymenium initially smooth, 
longitudinally rugose to rugulose in age, initially pale 
orange (5A3-2) to greyish orange (5B4-3), "light 
ochraceous-buff," "pale ochraceous-buff," "cartridge 
buff," "light buff," "pinkish buff," then brownish orange 
(6C6-4), "cinnamon-rufous," "cinnamon buff," "cinnamon," 
"orange-cinnamon," finally light brown (7D6-4) to brown 
(7E6-4), “mikado brown," "verona brown," "fawn color,” 
"army brown's; apex obtuse to broadly rounded, smooth, more 
or less concolorous with the hymenium; surface staining 
slowly, irregularly brown (6E6-4, 7E6-4) to dark brown 
(7F6-4), "verona brown," "mikado brown," "mar's brown," 
"russet'" where cut or bruised, staining more conspicuously 
downward; flesh initially solid, becoming soft and spongy 
upward as the apex enlarges, white to pallid, on exposure 
staining slowly, irregularly brown (6E6-4, 7E6-4) to dark 
brown (7F6-4), "verona brown," "russet," "mar's brown," 
"mikado brown". ODOR AND TASTE not distinctive. 


MACROCHEMICAL REACTIONS: FeCl., FeCl. + EtOH, ANO, 
PYR, PHN, GUA, PCR, SYR = positives; KOH, NH, OH, TYR = 
negative. 


MYCELIAL HYPHAE 2.5-5 yim diam, branched, clamped, 
uninflated; walls thin or irregularly thickened to 1 pn, 
hyaline in KOH, refractive under phase contrast, smooth, 
echinate or echinulate; projections up to 1 ym high, 
hyaline, insoluble in KOH, refractive under phase 
contrast, acyanophilous; clamps uninflated or inflated 
(-8 wm), sometimes medallion or ampulliform; hyphal 
contents amorphous, hyaline in KOH. TRAMAL HYPHAE 4-12 yim 
diam, more or less parallel to longitudinally interwoven 
basally, more loosely interwoven upward, radially 
interwoven beneath the subhymenium, uninflated, inflated 
(-16 pm) or broadly undulate, branched, clamped; walls 
thin or irregularly thickened to 1 ym diam, hyaline in 
KOH, refractive under phase contrast, smooth; clamps 
uninflated or inflated (-16 ym), sometimes medallion or 


157 


aie mee e®ee ae 


ee 
° 
@ 0%. %e 
eevee 


wien te pe e 


e 

e 

weteree & 
o® 


ee e 
. PaCS o%e % ote e e 


© 
o oo of & Ch herd 
e8 ee '@ "be 
e ey 
e*%e* .% 
@e 


Representative hymenium for taxa of 


basidia an 
Representative mycelial hyphae for taxa of subg 


d leptocystidia. 


Clavariadelphus 


4. 


10 pm. 


Clavariadelphus. Scale bars 


Clavariadelphus sect. 


158 


ampulliform; hyphal contents amorphous, hyaline in KOH. 
GLOEOPLEROUS HYPHAE 2.5-5 ym diam, arising from generative 
hyphae at clamp connections, scattered throughout the 
trama, more abundant downward, uninflated, inflated 

(-8 ym) or strangulated, branched, clamped; walls thin, 
hyaline in KOH, smooth; clamps uninflated or inflated 

(-8 pm), sometimes medallion or ampulliform; hyphal 
contents amorphous, subopalescent, yellow in KOH, 
refractive under phase contrast, cyanophilous. 

SUBHYMENIUM rudimentary; hyphae 2.5-5 ym diam, interwoven, 
branched, clamped, uninflated; walls thin, hyaline in KOH, 
smooth; clamps uninflateds; hyphal contents amorphous, 
hyaline in KOH. HYMENIUM thickening, extending from near 
the base of the basidiocarp over the apex, composed of 
basidia and leptocystidia: LEPTOCYSTIDIA 55-80 x 2.5-5.5 
jum, scattered among and scarcely projecting beyond the 
basidia, cylindric, subcylindric or strangulated, inflated 
apically at maturity, then narrowly clavate, at times 
apically or subapically branched, clamped; walls thin, 
hyaline in KOH, smooth; clamps uninflateds; hyphal contents 
amorphous, pale yellow in KOH, acyanophilous: BASIDIA 
60-110 x 8-12.5 pm, narrowly clavate to clavate, inflated 
apically at maturity, then broadly clavate, clamped; walls 
thin or irregularly thickened, hyaline in KOH, smooth; 
clamps uninflated; basidial contents multiguttulate and 
refringent to aguttulate and amorphous, pale yellow in 
KOH; sterigmata (2) -4, 7-9.5 yim long, broadest basally, 
narrowed to an obtuse apex, incurved. BASIDIOSPORES white 
in deposit, 8-12 x 4.5-6.5 pm (L_ = 9.6 pm; W_ = 5.2 pm; 
E=. 1652.13) EY = 128)s broadty. ovate to amyegdaliform; 
walls thin, hyaline in KOH, smooth; contents 
multiguttulate and refringent to aguttulate and amorphous, 
pale yellow in KOH; acyanophilous, inamyloids; hilar 
appendage oblique with an obtuse ‘apex. 


HABIT, HABITAT, AND DISTRIBUTION: Scattered to 
gregarious, less commonly in caespitose clusters; 
terrestrial; duff; mixed coniferous-deciduous forests in 
association with Quercus and Pinus, eastern North America: 
Illinois, Maine, Massachusetts, Michigan, Minnesota, New 
York, North Carolina, Nova Scotia, Ohio, Ontario, 
Pennsylvania, Tennessee, Vermont, and Wisconsin. 


SPECIMENS EXAMINED: CANADA. NOVA SCOTIA. Kings 
Co.: Kentville Research Station Ravine, 4 Aug 1973, 
Harrison No. 12137 (ut Clavariadelphus pistillaris) 
(MICH). Colchester Co.: Upper Brookside, 27 Aug 1931, 


159 


Smith No. 1417 (ut Clavaria pistillaris); Folleigh Lake, 
29 Aug 1931, Wehmeyer No. 1417a, (ut Clavaria 
pistillaris). ONTARIO. Haliburton Co.: Wren Lake, 

20 Sep 1984, Methven No. 3248 (TENN). Nipissing Co.: 
Algonquin Provincial Park, 21 Sept 1984, Methven Nos. 
Bio0.eo2>) (TENN). “UNLTED STATES. ILLINOIS.’ Jackson 
Co.: Lake Murpheysboro State Park, 7 Nov 1984, Methven No. 
3587 (TENN). MAINE. Aroostook Co.: Near Madawaska Lake, 
18 Aug 1956, Bigelow No. 4095 (ut Clavariadelphus 
pistillaris) (MICH). Hancock Co.: Acadia National Park, 
Mt. Desert Island, 1935, Morse s. n. (ut Clavariadelphus 
pistillaris) (TENN 31518). MASSACHUSETTS. Berkshire Co.: 
Oblong Road, vicinity North Adams, 15 Aug 1986, Methven 
No. 4715 (TENN); White Oaks Road, vicinity Williamstown, 
16 Aug 1986, Methven No. 4724 (TENN 47403) [NEOTYPE]. 
MICHIGAN. Emmet Co.: Pellston Hills, 25 Aug 1956, Thiers 
No. 4394 (ut Clavariadelphus pistillaris) (MICH). Gratiot 
Cossmectnaca, 15 Sep 1961, Potter! No. 13146 (MICH). 
Jackson Co.: Clear Lake, 13 Aug 1960, Shaffer No. 2604 
(ut Clavariadelphus pistillaris) (MICH). Marquette Co.: 
Canyon Lake Bay, 4 Sep 1970, Harrison No. 9614 (ut 
Clavariadelphus pistillaris) (MICH). Oakland Co.: Ormand 
Road, 12 Aug 1937, Smith No. 7012 (ut 'Clavaria 
pistillaris) (MICH; NCU); Haven Hill, Highland Recreation 
Area, 20 Sep 1972, Weber No. 3723 (ut Clavariadelphus 
pistillaris) (MICH); New Hudson, 26 Aug 1937, Smith No. 
7302 (ut Clavaria pistillaris) (MICH; NCU); Haven Hill, 

16 Sep 1961, Smith No. 64300 (ut Clavariadelphus 
pistillaris) (MICH); Horner Hill, 25 Sep 1975, Smith No. 
86661 (ut Clavariadelphus pistillaris var. americanus) 
(MICH). Ostego Co.: Vanderbilt, Pigeon River, Red 
Bridge, 10 Aug 1969, Smith No. 77794 (ut Clavariadelphus 
pistillaris) (MICH). Presque Isle Co.: Little Presque 
Isle Point, 28 Aug 1978, Ovrebo No. 634 (CLO). Washtenaw 
Co.: Ann Arbor, Horner Woods, 1 Aug 1973, Smith No. 84324 
(ut Clavariadelphus pistillaris) (MICH); South of Highland 
Lake, 2 Oct 1970, Weber No. 3768 (ut Clavariadelphus 
pistillaris) (MICH); Waterloo, Bush Road Trail, 2 Aug 
1973, Nimke No. 401 (ut Clavariadelphus pistillaris) 
(MICH). MINNESOTA, Rice Co.: Wheeling Township, 
Nerstrand Woods State Park, 24 Jul 1962, Weaver No. 
7-24-62-N-1 (ut Clavariadelphus pistillaris f. americanus) 
(MICH; TENN 33669). NEW YORK. Washington Co.: Vaughns, 
11 Aug 1915, Burnham No. 68 (ut Clavaria pistillaris) 
(NCU); Vaughns, 2 Sep 1917, Burnham No. 139 (ut Clavaria 
pistillaris) (NCU). NORTH CAROLINA. Macon Co.: 
Highlands, 10 Sep 1971, Harrison No. 11093 (ut 


160 


Clavariadelphus pistillaris) (MICH); Horse Cove, Jul 1964, 
No. 31411 (ut Clavariadelphus pistillaris) (TENN); 
Cliffside Lake, 2 Sep 1986, Desjardin No. 4101 (TENN); 
Highlands, 30 Aug 1932, No. 9599 (ut Clavaria pistillaris) 
(NCU). Orange Co.: Vicinity of Ginghaul Castle, 22 Oct 
1947, No. 14321 (ut Clavaria pistillaris) (NCU); Chapel 
Hill, Battle's Branch, near Indian Springs, 21 Oct 1915, 
No. 1913 (ut Clavaria unicolor) (NCU); Chapel Hill, Battle 
Park, 20-23 Sep 1945, No. 13933 (ut Clavaria unicolor) 
(NCU); Chapel Hill, Battle Park, 24 Sep 1945, No. 13920 
(ut Clavaria pistillaris var. unicolor) (NCU); Chapel 
Hill, Battle Park, 26 Sep 1945, No. 13950 (ut Clavaria 
pistillaris var. unicolor) (NCU); Chapel Hill, Meeting of 
Waters, 26 Sep 1945, No. 13951 (ut Clavaria pistillaris 
var. unicolor) (NCU); Chapel Hill, Meeting of Waters, 

28 Sep 1945, No. 13960, (ut Clavaria pistillaris var. 
unicolor) (NCU); Chapel Hill, Meeting of Waters, 3 Oct 
1945, No. 13995 (ut Clavaria pistillaris var. unicolor) 
(NCU); Chapel Hill, Mason Farm, 25 Oct 1946, No. 14068 (ut 
Clavaria pistillaris) (NCU); Chapel Hill, New Hope Creek 
Farm, 24 Oct 1947, No. 14324 (ut Clavaria pistillaris) 
(NCU). Swain Co.: Almond, 21 Sep 1971, Harrison Nos. 
11233, 11234, 11235 (ut Clavariadelphus pistillaris) 
(MICH). Transylvania Co.: Pisgah National Forest, White 
Pines Campground, 13 Sep 1966, No. 32337 (ut 
Clavariadelphus pistillaris) (TENN). OHIO. Hamilton Co.: 
Miami-Whitewater Forest Park, Timberlakes Area, 2 Oct 
1965, Cooke No. 36192 (ut Clavariadelphus pistillaris) 
(WBC:. TENN. 32445). © Pike ‘Coz: Pike Lakes State Parkes 
Sep 1973, Cooke No. 48742 (WBC; TENN 47398). 

PENNSYLVANIA. Chester Co.: West Chester, Sep-Oct, 1887, 
North American Flora, Second Series No. 1922b (ut 
Craterellus pistillaris) (PAD). TENNESSEE. Blount Co.: 
Nales Creek, 26 Sep 1959, No. 22939 (ut Clavaria 
pistillaris) (TENN). Knox Co.: Ball Camp Pike, 6 Oct 
1934, No. 6542 (ut Clavaria pistillaris) (TENN); Ball Camp 
Pike, 9 Oct 1934, No. 6543 (ut Clavaria pistillaris) 
(TENN). VERMONT. Windham Co.: Grafton State Forest, 

30 Aug 1961, Shaffer No. 3510 (ut Clavariadelphus 
pistillaris) (MICH). WISCONSIN. Dane Co.: Blue Mounds, 
Aug 1903, No. 162 (ut Clavaria pistillaris) (NCU). 


Corner (1950) described C. pistillaris var. 
americanus from frondose wood in eastern North America. 
Although Corner (1950) published a Latin diagnosis for 
var. americanus he did not designate a type specimen 
having examined no North American specimens at the time. 


161 


Instead, Corner based his variety on previous literature 
reports of C. pistillaris in North America, noting that it 
differed "in colour, form, and small spores from the 
European species." 

In an attempt to bring order to the taxa similar to 
C. pistillaris in North America, Wells and Kempton (1968) 
proposed C. cokeri based on "Coker's excellent notes, the 
good spore deposit, Miss Eaton's fine illustration (Coker, 
1932 [sic], Pl. 23)" and Coker's specimen (No. R 27, NCU). 
Wells and Kempton (1968) did not include Corner's var. 
americanus in their preliminary study of Clavariadelphus 
in North America, concluding that var. americanus was 
"based on literature" and because "we have seen no 
specimens which could be referred to it." Corner (1970), 
in a critical rebuttal, noted that Wells and Kempton 
dismissed "Coker's contribution and discarded the idea of 
var. americanus because it was based on literature and 
because they had seen no specimens which could be referred 
to it." Corner (1970) went on to say that var’. 
americanus was based "not on literature (? belles lettres) 
but on the scientific researches of Harper, Coker, and 
Wehmeyer by direct citation and of Burt and Doty by 
reference in my Table XXI (Monogr. p. 282)." In Corner's 
own words then, his protologue (Corner, 1950) for var. 
americanus was taken from literature and in no part from 
specimens he had examined. After examining the 
collections for which Corner (1950) cited spore 
measurements in Table XXI, I found them to be 
representative of several discrete taxa. Coker's specimen 
from Connecticut (No. R 27, NCU) is the holotype of C. 
cokeri Wells and Kempton, Wehmeyer's collections (Smith 
1417, Wehmeyer 1417a, MICH) are C. americanus, Doty's 
specimens are C. occidentalis, two of Coker's collections 
cited from New York (Burnham 68, 71, NCU) are C. 
americanus and C. truncatus, respectively, and Coker's 
collections from North Carolina are C. americanus (Nos. 
1913, 3793, NCU) and C. unicolor (No. 4770, NCU). It is 
little wonder, then, that Wells and Kempton were confused 
by Corner's concept of var. americanus. 

Following an examination of specimens labeled as C. 
pistillaris from eastern North America I believe one 
element of Corner's conglomerate taxon is more accurately 
recognized as autonomous. In order to provide maximum 
taxonomic data, one of my collections has been selected as 
a neotype for this taxon. Although the basidiocarps of C. 
americanus are similar in coloration to those of C. 
pistillaris, they are generally not as robust, the flesh 


162 


has a sweet or nondistinctive taste, the basidiospores are 
smaller (10.5-14 x 6-7.5 pm for C. pistillaris), and it 
grows in mixed deciduous-coniferous forests in association 
with Pinus and Quercus. 

In eastern North America, C. americanus is most 
likely to be confused with C. cokeri and C. flavidus. 
Clavariadelphus cokeri is distinct in its caespitose to 
fasciculate growth habit in hemlock forests, pinkish buff 
to rose pink basidiocarps, which are at times branched 
apically, and longer, narrower basidiospores (L_ = 10.3 
pm; We = 4.4 pm). Clavariadelphus flavidus occurs in 
mixed coniferous-deciduous forests, produces bright yellow 
to lemon yellow basidiocarps, flesh without a distinctive 
taste, and the smallest basidiospores in subg. 
Clavariadelphus (7-10.5 x 4.5-5.5 ym). 

In cases where the apex of the basidiocarp has been 
damaged during development, and becomes more or less 
truncate or turbinate as a result, C. americanus might be 
confused with C. unicolor (Rav. apud Berk.) Corner, which 
is found in similar habitats. Clavariadelphus unicolor is 
distinct, however, by its violet-brown to lavender-brown 
basidiocarps which become truncate early in development, 
often appearing cantharelloid or craterelloid at maturity, 
and longer, narrower basidiospores (L_ = 10.3 pm; W_ = 5.0 
ym). As long as several basidiocarps at different stages 
of development are available, there is little problem in 
segregating these two taxa. 


Clavariadelphus caespitosus Methven, sp. nov. Figs. 2, 6 


Receptaculum ad 15 cm altum, apice 5-15 mm latum, 
simplex, fusiforme vel clavatum; apex subacutus, laevis 
vel rugosus; hymenium laeve vel rugosum, juventute 
cinnamomeum, maturitate ferrugineum vel castaneum; caro 
spongiosa, albida, brunnescens; odor et sapor nullus3; 
sporae 8-11.5 x 4.5-6.5 pm (L_ = 9.8 pms W_ = 5.6 pm; 

E = 1.7-1.93; E = 1.8)3 late Ovatae vel amygdaliformes, 
laeves, albae; "hyphae monomiticae, intertextae, fibulatae. 
Gregaria vel cespitosa ad terram in sylvis coniferis 
Americae borealis occidentalis. 


HOLOTYPUS: TENN: United States, Idaho, Bonner Co., 


Upper Priest River, 21 Sep 1968, legit A. H. Smith s. n. 
(TENN 33983). 


163 


BASIDIOCARP 3-15 cm high, 3-15 (20) mm diam apically, 
simple, intially subcylindric, enlarged upward in age, 
then fusiform to clavate; base terete, pruinose, orange 
white (5A2, 6A2), "pale ochraceous salmon," "buff-pink," 
"seashell pink"; hymenium initially smooth, longitudinally 
rugose to rugulose in age, initially greyish red (7B3-2) 


to dull red (8C3-2), "cinnamon buff," "cinnamon," "japan 


rose," "“vinaceous-buff," "light vinaceous—fawn," finally 
reddish brown (8D5-4), "mikado brown," "cacao brown," 
"fawn-color," "vinaceous-russet"; apex subacute to 
narrowly obtuse, remaining more or less pointed throughout 
development, smooth, concolorous with the hymeniun; 
surface staining slowly, irregularly brown (6E6-4, 7E6-4) 
to dark brown (7F6-4), "verona brown," "mikado brown," 
"russet," "mar's brown" where cut or bruised, staining 
more conspicuously downward; flesh initially solid, 
becoming soft and spongy upward as the apex enlarges, 
white to pallid, on exposure staining slowly, irregularly 
brown (6E6-4, 7E6-4) to dark brown (7F6-4), "verona 
brown,'’ "mikado brown," “russet," "mar's brown". ODOR not 
distinctive. TASTE not distinctive or slightly bitter. 


MACROCHEMICAL REACTIONS: FeC1l.,, FeCl, + EtOH, NAP, 
ANO, PYR, PHN, GUA = positive; KOH = yellow to golden 
yellow; NH4OH = negative. 


MYCELIAL HYPHAE 2.5-4 yim diam, branched, clamped, 
uninflated; walls thin or irregularly thickened to Il pm, 
hyaline in KOH, refractive under phase contrast, smooth, 
echinate or echinulate; projections up to | pm high, 
hyaline, insoluble in KOH, refractive under phase 
contrast, acyanophilouss; clamps uninflated or inflated 
(-12 wm), sometimes medallion or ampulliform; hyphal 
contents amorphous, hyaline in KOH. TRAMAL HYPHAE 3-12 pm 
diam, more or less parallel to longitudinally interwoven 
basally, more loosely interwoven upward, radially 
interwoven beneath the subhymenium, uninflated, inflated 
(-16 ym) or broadly undulate, branched, clamped; walls 
thin or irregularly thickened to 1 ym, hyaline in KOH, 
refractive under phase contrast, smooth; clamps uninflated 
or inflated (-16 jm), sometimes medallion or ampulliforn; 
hyphal contents amorphous, hyaline in KOH. GLOEOPLEROUS 
HYPHAE 2.5-6.5 pm diam, arising from generative hyphae at 
clamp connections, scattered throughout the trama, more 
abundant downward, uninflated, inflated (-9.5 jm) or 
strangulated, branched, clamped; walls thin, hyaline in 
KOH, smooth; clamps uninflated or inflated (-12 sm), 


164 


sometimes medallion or ampulliform; hyphal contents 
subopalescent, yellow in KOH, refractive under phase 
contrast, cyanophilous. SUBHYMENIUM rudimentary; hyphae 
2.5-5 pm diam, interwoven, branched, clamped, uninflated; 
walls thin, hyaline in KOH, smooth; clamps uninflated; 
hyphal contents amorphous, hyaline in KOH. HYMENIUM 
thickening, extending from near the base of the 
basidiocarp over the apex, composed of basidia and 
leptocystidia: LEPTOCYSTIDIA 65-125 x 2.5-5 ym, scattered 
among and scarcely projecting beyond the basidia, 
cylindric, subcylindric or strangulated, inflated apically 
at maturity, then narrowly clavate, at times apically or 
subapically branched, clamped; walls thin, hyaline in KOH, 
smooths; clamps uninflated; contents amorphous, pale yellow 
in KOH, acyanophilous: BASIDIA 70-125 x 8-13.5 pn, 
narrowly clavate to clavate, inflated apically at 
maturity, then broadly clavate, clamped; walls thin or 
irregularly thickened, hyaline in KOH, smooth; clamps 
uninflated; basidial contents multiguttulate and 
refringent to aguttulate and amorphous, pale yellow in 
KOH; sterigmata (2) -4, 7-11 ym long, broadest basally, 
narrowed to an obtuse apex, incurved. BASIDIOSPORES white 
in deposit, 8-11.5 x 4.5-6.5 pm (L = 9.8 pm; W_ = 5.6 pm; 
E = 1.7-1.9; E_ = 1.8)3 broadly ovate to amygdaliform; 
walls thin, hyaline in KOH, smooths; contents 
multiguttulate and refringent to aguttulate and amorphous, 
pale yellow in KOH; acyanophilous, inamyloid; hilar 
appendage oblique with an obtuse apex. 


HABIT, HABITAT, AND DISTRIBUTION: Gregarious, 
densely gregarious or caespitosus clusters; terrestrial; 
duff; coniferous or mixed coniferous-deciduous forests; 
California, Idaho, Oregon, and Washington. 


SPECIMENS EXAMINED: UNITED STATES. CALIFORNIA. Del 
Norte Co.: Jedediah Smith Redwoods National Park, 11 Sep 
1967, No. 33193 (ut Clavariadelphus pistillaris) (TENN). 
Humboldt Co.: Fickel Hill, 14 Sep 1984, Saylor No. 2184 
(SFSU). Mendocino Co.: Jackson State Forest, 23 Nov 
1985, Methven No. 4396 (TENN). Yuba Co.: Bullard's Bar 
Recreation Area, Schoolhouse Campground, 15 Dec 1983, 
Methven No. 2681 (TENN). IDAHO. Bonner Co.: Upper 
Priest River, 4 Oct 1970, No. 34685 (TENN); Priest Lake, 

2 Oct 1966, No. 31954 (TENN); Priest Lake, 28 Sep 1968, 
Smith No. 76695 (MICH); Priest River, 19 Sep 1968, No. 
33937 (TENN); Upper Priest River, 21 Sep 1968, Smith s. n. 
(TENN 33983) [HOLOTYPE]; Priest Lake, 29 Sep 1968, Smith 


165 


No. 76727 (MICH); Upper Priest River, 19 Sep 1968, Smith 
No. 76548 (MICH); Upper Priest River, 25 Sep 1968, Smith 
No. 76667 (MICH; TENN 47399). Kootenai Co.: Couer 
d'Alene, 7 Oct 1970, Nos. 34724, 34644 (TENN); Couer 
d'Alene National Forest, Deception Creek Experimental 
Forest, 22 Sep 1969, Smith no. 70662 (ut Clavariadelphus 
ligulus) (MICH). OREGON. Clackamas Co.: Mt Hood 
National Forest, 20 Oct 1984, Methven Nos. 3519, 3520 
(TENN). WASHINGTON. Pierce Co.: Mt Rainier National 
Park, 15 Oct 1984, Methven No. 3540 (TENN). 


Figs. 5-8. 5. Basidiospores of Clavariadelphus 
americanus (TENN 47403, Neotype). 6. Basidiospores of 
Clavariadelphus caespitosus (TENN 33983, Holotype). 

7. Basidiospores of Clavariadelphus flavidus (TENN 47401, 
Holotype). 8. Basidiospores of Clavariadelphus 
occidentalis (TENN 47402, Holotype). Scale bars = 10 yn. 


166 


Clavariadelphus caespitosus is one of several taxa 
within sect. Clavariadelphus passing under the name C. 
pistillaris sensu lato. Clavariadelphus caespitosus is, 
however, quite distinct from that taxon by its slender, 
narrowly clavate, greyish red to dull red basidiocarps 
with subacute apices, caespitose growth habit in 
coniferous or mixed coniferous-deciduous forests, and 
smaller basidiospores (10.5-14 x 6-7.5 gm for C. 
pistillaris). In western North America C. caespitosus 
might be confused with C. subfastigiatus or C. 
fasciculatus Methven and Guzm4n. Clavariadelphus 
subfastigiatus differs in its pallid flesh-color to light 
cinnamon basidiocarps with obtuse apices, forest green 
staining reaction with KOH, scattered to gregarious growth 
habit in coniferous forests, and smaller basidiospores 
(8-10.5 x 5-6 jim). Clavariadelphus fasciculatus, known at 
this writing only from central Mexico, is distinct by the 
strongly fasciculate growth habit of its basidiocarps, 
larger basidiospores (11-14 x 5-7 jim), and acerose- to 
acicular-shaped crystals encrusting the walls of the 
mycelial hyphae. 


Clavariadelphus flavidus Methven, sp. nov. Fig lar 


Receptaculum 10-15 cm altum, apice 10-12 mm latum, 
simplex, cylindricum vel anguste clavatum; apex subacutus 
vel obtusus, laevis; hymenium laeve vel rugosum, laete 
aureum vel flavidum; caro spongiosa, albida, brunnescens; 
odor nullus, sapor amarus; sporae 7-10.5 x 4.5-5.5 jim 
(L. = 8.4 yum; Wo = 5.0 pms E = 1.4-2.03 Evi = 17) ee ace 
ovatae vel amygdaliformes, laeves, albae; hyphae 
monomiticae, intertextae, fibulatae. Sparsa ad terram in 
sylvis frondosis-coniferis Americae borealis orientalis. 


HOLOTYPUS: TENN: United States, Massachusetts, 
Berkshire Co., Williamstown Reservoir, 14 Aug 1986, legit 
A. S. Methven No. 4714 (TENN 47401). 


BASIDIOCARP 10-15 cm high, 10-12 mm diam apically, 
simple, cylindric to narrowly clavate; mycelial hyphae 
interwoven or aggregated into rhizomorphic strands up to 
1 mm diam, white to pallid; base terete, smooth, pruinose, 
off-white to pale cream; hymenium intially smooth, 
longitudinally rugulose in age, yellow (3A7-6) to 
yellowish orange (4A7-5), "apricot yellow," "light orange 
yellow," "light cadmium," "primuline yellow"; apex 


167 


subacute to obtuse, smooth, concolorous with the hymenium; 
surface staining slowly, irregularly brown (6E5-4, 7E5-4) 
to dark brown (7F5-4), "rood's brown," "vandyke brown," 
"army brown," "natal brown," "bone brown" where cut or 
bruised, staining more conspicuously downward; flesh 
initially solid, becoming soft and spongy upward as the 
apex enlarges, white to pallid, on exposure staining 
slowly, irregularly brown (6E5-4, 7E5-4) to dark brown 
(7F5-4), "rood's brown," "vandyke brown," "army brown," 
"natal brown," "bone brown". ODOR not distinctive. TASTE 
bitter. 


MACROCHEMICAL REACTIONS: FeCl., FeCl. + EtOH, PCR, 
PHN, PYR, ANO, GUA, SYR = positive; KOH, NH4OH, NAP, TYR = 
negative. 


MYCELIAL HYPHAE 2.5-5 yim diam, branched, clamped, 
uninflated; walls thin or irregularly thickened to 1 gim, 
hyaline in KOH, refractive under phase contrast, smooth, 
echinate or echinulate; projections up to 0.5 gm high, 
hyaline, insoluble in KOH, acyanophilous; clamps 
uninflated or inflated (-12 zm), sometimes medallion or 
ampulliform; hyphal contents amorphous, hyaline in KOH. 
TRAMAL HYPHAE 3-9.5 yim diam, more or less parallel to 
longitudinally interwoven basally, more loosely interwoven 
upward, radially interwoven beneath the subhymenium, 
uninflated, inflated (-12 ,m) or broadly undulate, 
branched, clamped; walls thin or irregularly thickened to 
1 ym, hyaline in KOH, refractive under phase contrast, 
smooth; clamps uninflated or inflated (-16 ym), sometimes 
medallion or ampulliform; hyphal contents amorphous, 
hyaline in KOH. GLOEOPLEROUS HYPHAE 3-5 yim diam, arising 
from generative hyphae at clamp connections, scattered 
throughout the trama, more abundant downward, uninflated, 
inflated (-9.5 ym) or strangulated, branched, clamped; 
walls thin, hyaline in KOH, smooth; clamps uninflated or 
inflated (-8 jm), sometimes medallion or ampulliform; 
hyphal contents subopalescent, yellow in KOH, refractive 
under phase contrast, cyanophilous. SUBHYMENIUM 
rudimentary; hyphae 2.5-5 »m diam, interwoven, branched, 
clamped, uninflated; walls thin, hyaline in KOH, smooth; 
clamps uninflateds; hyphal contents amorphous, hyaline in 
KOH. HYMENIUM thickening, extending from near the base of 
the basidiocarp over the apex, composed of basidia and 
leptocystidia: LEPTOCYSTIDIA 50-80 x 2.5-4 yim, scattered 
among and scarcely projecting beyond the basidia, 
cylindric, subcylindric or strangulated, inflated apically 


168 


at maturity, then narrowly clavate, at times apically or 
subapically branched, clamped; walls thin, hyaline in KOH, 
smooth; clamps uninflated; hyphal contents amorphous, pale 
yellow in KOH, acyanophilous: BASIDIA 80-105 x 8-11 gm, 
narrowly clavate to clavate, inflated apically at 
maturity, then broadly clavate, clamped; walls thin or 
irregularly thickened, hyaline in KOH, smooth; clamps~ 
uninflated; basidial contents multiguttulate and 
refringent to aguttulate and amorphous, pale yellow in 
KOH; sterigmata (1,2,3) -4, 8-11 ym long, broadest 
basally, narrowed to obtuse apex, incurved. BASIDIOSPORES 
white in deposit, 7-10.5 x 4.5-5.5 pm (L = 8.4 pm; 

We = 5.0 jms) E = 1,4-2.0; En = 1.7); broadly ovate to 
amygdaliform; walls thin, hyaline in KOH; contents 
multiguttulate and refringent to aguttulate and amorphous, 
pale yellow in KOH; acyanophilous, inamyloid; hilar 
appendage oblique with an obtuse apex. 


HABIT, HABITAT, AND DISTRIBUTION: Solitary or 
scattered; terrestrial; duff; mixed deciduous-coniferous 
forests under Larix, Picea, Betula, Populus, Tilia, and 
Pinus; Massachusetts. 


SPECIMENS EXAMINED: UNITED STATES. MASSACHUSETTS. 
Berkshire Co.: Williamstown Reservoir, 14 Aug 1986, 
Methven No. 4714 (TENN 47401) [HOLOTYPE]. 


The hymenium extending from near the base of the 
basidiocarp over the apex, mycelial hyphae in which the 
walls are echinate or echinulate, and the broadly ovate to 
amygdaliform basidiospores with an Em €2.0 indicate that 
C. flavidus is best placed in subg. Clavariadelphus sect. 
Clavariadelphus. In North America there are several taxa 
within sect. Clavariadelphus with which C. flavidus might 
be confused, including C. occidentalis, C. pistillaris, 
and C. americanus. Although Clavariadelphus americanus 
shares similar habitats with C. flavidus in mixed 
coniferous-deciduous forests in eastern North America, its 
larger, more robust, pale ochraceous basidiocarps which 
darken to cinnamon brown in age, and longer, broader 
basidiospores (L_ = 9.6 ym; W_ = 5.2 ym) are distinctive. 
Clavariadelphus pistillaris differs in its more robust, 
clavate, pale ochraceous basidiocarps which darken to 
cinnamon brown or fawn brown in age, larger basidiospores 
(10.5-14 x 6-7.5 ym), and habitat in deciduous forests in 
association with Fagus. Clavariadelphus occidentalis, 
which, as far as is known, is restricted to western North 


169 


America, exhibits more robust, pale yellow basidiocarps 
which darken to greyish orange in age, larger 
basidiospores (9-13.5 x 5.5-7 pm), and mild tasting flesh. 

Among the European taxa included in sect. 
Clavariadelphus, C. flavo-immaturus Petersen and C. 
xanthocephalus Rahm & Schild are similiar to, yet distinct 
from, C. flavidus. Clavariadelphus flavo-—immaturus 
features more robust, clavate basidiocarps, larger 
basidiospores (11-13.5 x 5.5-7 ym), and an orange staining 
reaction with KOH. Clavariadelphus xanthocephalus 
exhibits larger, more robust, turbinate or capitate 
basidiocarps, larger basidiospores (9.5-12 x 4.5-6 jim), 
and is not reactive with KOH. 

In the field, C. flavidus might also be confused with 
C. ligula and C. sachalinensis (Imai) Corner, both of 
whose basidiocarps are infrequently bright yellow in 
color. These taxa can readily be segregated 
microscopically, however, by basidiospore size and shape. 
The basidiospores of C. flavidus are broadly ovate to 
amygdaliform in profile, with an Em €2.0, while those of 
C. ligula and C. sachalinensis are narrowly ellipsoid to 
boletoid in profile with an Em> 3.0. 


Clavariadelphus occidentalis Methven, sp. nov. 
PLR ile 


Receptaculum 10-20 cm altum, apice 10-30 mm latun, 
simplex, clavatum; apex obtusus, laevis dein rugosus; 
hymenium laeve dein rugosum, luteolum vel ochraceum; caro 
spongiosa, albida, brunnescens; odor et sapor nullus; 
sporae 9.5-13 x 5.5-7 gm (L_ = 11.1 pm; W_ = 6.3 pm; 

Reegie 4 2.03)-E = 1.7): late ovatae vel amygdaliformes, 
laeves, albae; hyphae monomiticae, intertextae, fibulatae. 
Sparsa vel gregaria ad terram in sylvis frondosis-— 
coniferis Americae borealis occidentalis. 


HOLOTYPUS: TENN: United States, California, Yuba 
Co., Bullard's Bar Recreation Area, Schoolhouse 
Campground, 4 Jan 1987, legit A. S. Methven No. 5068 
(TENN 47402). 


BASIDIOCARP 5-20 cm high, 5-12 mm diam basally, 10-30 
mm diam apically, simple, initially subcylindric to 
subfusiform, enlarged upward in age, then clavate to 
broadly clavate, finally irregularly, laterally 
compressed; mycelial hyphae interwoven or aggregated into 


170 


rhizomorphic strands up to 1 mm diam, white to pallid; 
base terete, smooth, pruinose, white to pallid where 
covered, otherwise yellowish white (3A2) to pale yellow 
(4A3), “Ivory yellow," "cartridge, buff," “lightebuGaen 
"pale ochraceous-buff"; hymenium initially smooth, 
longitudinally rugose to rugulose in age, initially light 
yellow (4A5-4) to orange white (5A2), "light buff," "pale 
ochraceous-buff,'"' "pale ochraceous-salmon," "warm buff," 
"chamois," "light ochraceous-buff," "cream buff," finally 
light orange (5A4-3) to greyish orange (5B4-3, 6B4-3), 
"capucine buff,'' "pale yellow-orange," "cinnamon-buff ," 
"pinkish cinnamon"; apex initially subacute, then obtuse 
or broadly rounded, smooth, concolorous with the hymenium; 
surface staining slowly, irregularly brown (6E6-4, 7E6-4) 
to dark brown (7F6-4), 'verona brown," "mikado brown," 
"russet,'' "mar's brown" where cut or bruised, staining 
more conspicuously downward; flesh initially solid, 
becoming soft and spongy upward as the apex enlarges, 
white to pallid, on exposure staining slowly, irregularly 
brown (6E6-4, 7E6-4) to dark brown (7F6-4), "verona 
brown," "mikado brown," "russet,” "mar's brown". ODOR AND 
TASTE not distinctive. 


MACROCHEMICAL REACTIONS: FeCl., FeCl. + EtOH, ANO, 
PHN, PYR, PCR, GUA, SYR = positive; KOH, NH4OH, NAP, TYR = 
negative. 


MYCELIAL HYPHAE 2.5-4 jim diam, branched, clamped, 
uninflated; walls thin or irregularly thickened to 1 pn, 
hyaline in KOH, refractive under phase contrast, smooth, 
echinate or echinulate; projections up to 1 pm high, 
hyaline, insoluble in KOH, refractive under phase 
contrast, acyanophilous; clamps uninflated or inflated 
(-9.5 pm), sometimes medallion or ampulliforms; hyphal 
contents amorphous, hyaline in KOH. TRAMAL HYPHAE 4-12 pm 
diam, more or less parallel to longitudinally interwoven 
basally, more loosely interwoven upward, radially 
interwoven beneath the subhymenium, uninflated, inflated 
(-13.5 wm) or broadly undulate, branched, clamped; walls 
thin or irregularly thickened to 1 ym, hyaline in KOH, 
refractive under phase contrast, smooth; clamps uninflated 
or inflated (-16 jm), sometimes medallion or ampulliforn; 
hyphal contents amorphous, hyaline in KOH. GLOEOPLEROUS 
HYPHAE 2.5-5 yim diam, arising from generative hyphae at 
clamp connections, scattered throughout the trama, more 
abundant downward, uninflated, inflated (-8 jm) or 
strangulated, branched, clamped; walls thin, hyaline in 


171 


Figs. 9-11. Micromorphological characters of 
Clavariadelphus pallido-incarnatus (TENN 47404, Holotype). 
9. Apical pellis. 10. Mycelial hyphae. 

11. Basidiospores. Scale bars = 10 ym for apical pellis 
and mycelial hyphae; 5 pm for basidiospores. 


172 


KOH, smooth; clamps uninflated or inflated (-12 pm), 
sometimes medallion or ampulliform; hyphal contents 
subopalescent, yellow in KOH, refractive under phase 
contrast, cyanophilous. SUBHYMENIUM rudimentary; hyphae 
2.5-5 pm diam, interwoven, branched, clamped, uninflated; 
walls thin, hyaline in KOH, smooth; clamps uninflated; 
hyphal contents amorphous, hyaline in KOH. HYMENIUM - 
thickening, extending from near the base of the 
basidiocarp over the apex, composed of basidia and 
leptocystidia: LEPTOCYSTIDIA 55-85 x 2.5-5 pm, scattered 
among and scarcely projecting beyond the basidia, 
cylindric, subcylindric or strangulated, inflated apically 
at maturity, then narrowly clavate, at times apically or 
subapically branched, clamped; walls thin, hyaline in KOH, 
smooth; clamps uninflated; hyphal contents amorphous, pale 
yellow in KOH, acyanophilous: BASIDIA 70-125 x 9.5-14 pm, 
narrowly clavate to clavate, inflated apically at 
maturity, then broadly clavate, clamped; walls thin or 
irregularly thickened, hyaline in KOH, smooth; clamps 
uninflated; basidial contents multiguttulate and 
refringent to aguttulate and amorphous, pale yellow in 
KOH; sterigmata (2) -4, 8-12 ym long, broadest basally, 
narrowed to an obtuse apex, incurved. BASIDIOSPORES white 
in deposit, 9.5-13 x 5.5-7 pm (L_ = 11.1 pm; W_ = 6.3 pom; 
E = 1.4-2.0; E. = 1.7)3 broadly ovate to amygdaliform; 
walls thin, hyaline in KOH, smooth; contents 
multiguttulate and refringent to aguttulate and amorphous, 
pale yellow in KOH; acyanophilous, inamyloid; hilar 
appendage oblique with an obtuse apex. 


HABIT, HABITAT, AND DISTRIBUTION: Scattered to 
gregarious, infrequently in caespitose clusters; 
terrestrial; duff; coniferous or mixed coniferous- 
deciduous forests: Alaska, Arizona, California, Idaho. 


SPECIMENS EXAMINED: UNITED STATES. ALASKA. 
Gustavus, Rink River, Johnson Homestead, 6 Sep 1981, Cooke 
No. 59896 (WBC; TENN 47397); Fairbanks, Harding Lake 
Campground, 26 Aug 1965, Wells and Kempton Nos. 
8/26/65-14, 8/26/65-15 (ut Clavariadelphus pistillaris) 
(WK 1984, 1985); Fairbanks, Livingood Road, 15 Aug 1964, 
Wells and Kempton No. 8/15/64-10 (ut Clavariadelphus 
pistillaris) (MICH); Glacier Bay National Monument, Water 
Pumping Station, 6 Sep 1979, Cooke No. 57176 (WBC; TENN 
44854); Glacier Bay National Monument, Bartlett Cove, 

3 Sep 1979, Cooke No. 57002 (WBC; TENN 44855); Glacier Bay 
National Monument, Water Pumping Station, 7 Sep 1979, 


173 


Cooke No. 57231 (WBC; TENN 44852). ARIZONA. Graham Co.: 
Coronado National Forest, 29 Aug 1958, Lowe No. 9425 (TENN 
32040). CALIFORNIA. Humboldt Co.: Willow Creek, no 
date, Largent No. 8158 (HSC). Marin Co.: Samuel P. 
Taylor State Park, 26 Dec 1985, Methven No. 4409 (TENN); 
Alpine Lake, 8 Dec 1984, Thiers No. 48393 (ut 
Clavariadelphus pistillaris) (SFSU). San Mateo Co.: San 
Francisco Watershed, 12 Jan 1986, Saylor No. 3041 (SFSU). 
Santa Barbara Co.: 17 Feb 1940, Rea No. 370 (ut Clavaria 
Pistillaris) (MICH). Trinity Co.: Brizard Ranch, east) of 
Salyer, Denning Road, 6 Dec 1982, Lanphere No. 12/6/62 (ut 
Clavariadelphus pistillaris) (MICH). Tuolumne Co.: Hwy 
120, Moccasin Creek Recreation Area, 27 Jan 1979, Thiers 
No. 39368 (ut Clavariadelphus pistillaris) (SFSU). Yuba 
Co.: Bullard's Bar Recreation Area, Schoolhouse 
Campground, 15 Dec 1983, Methven No. 2682 (TENN); 
Bullard's Bar Recreation Area, Schoolhouse Campground, 

4 Jan 1987, Methven No. 5068 (TENN 47402) [HOLOTYPE]; 
Bullard's Bar Recreation Area, Schoolhouse Campground, 

4 Jan 1987, Methven No. 5069 (TENN). IDAHO. Bonner Co.: 
Priest Lake, 29 Sep 1984, Methven Nos. 3320, 3323 (TENN); 
Nordman, Granite Creek, 12 Oct 1956, Smith No. 54506 (ut 
Clavariadelphus pistillaris) (MICH); Upper Priest River, 
21 Sep 1968, Smith No. 76553 (ut Clavariadelphus 
pistillaris) (MICH); Priest Lake, Binarch Creek, 22 Sep 
1968, Smith No. 76574 (MICH); Priest Lake, 29 Sep 1968, 
Smith No. 76770 (ut Clavariadelphus pistillaris) (MICH); 
Priest Lake, Binarch Creek, 8 Oct 1968, Smith No. 77019 
(MICH); Priest River, 21 Sep 1966, No. 32771 (ut 
Clavariadelphus pistillaris) (TENN); Priest River, 6 Oct 
1966, No. 32241 (ut Clavariadelphus pistillaris) (TENN); 
Priest River, 21 Sep 1968, No. 33997 (TENN); Priest River, 
22 Sep 1968, No. 34017 (TENN); Priest Lake, 24 Sep 1968, 
No. 34041 (TENN); Priest Lake, 23 Sep 1968, No. 34076 (ut 
Clavariadelphus pistillaris) (TENN); Priest River, 1 Oct 
1968, No. 34261 (ut Clavariadelphus pistillaris) (TENN); 
Priest River, 7 Oct 1968, No. 34340 (TENN); Priest River 
3 Oct 1968, No. 34708 (TENN). Kootenai Co.: Couer 
d'Alene, 30 Sep 1966, No. 32206 (ut Clavariadelphus 
pistillaris) (TENN); Couer d'Alene, 7 Oct 1970, No. 34737 
(ut Clavariadelphus pistillaris) (TENN). Shoshone Co.: 
Devil's Elbow, Couer d'Alene River, 24 Oct 1972, Smith No. 
83190 (ut Clavariadelphus pistillaris). 


Clavariadelphus occidentalis is one of several taxa 
in sect. Clavariadelphus passing under the name C. 


pistillaris sensu lato. Clavariadelphus occidentalis 


174 


differs from the latter by its light yellow basidiocarps 
which darken to greyish orange in age, habitat in 
coniferous or mixed coniferous-deciduous forests, and 
smaller basidiospores (10.5-14 x 6-7.5 pm for Cc. 
pistillaris). Other taxa within sect. Clavariadelphus 
with which C. occidentalis might be confused include C. 
subfastigiatus, C. caespitosus, and Cc. fasciculatus. ~ 
Clavariadelphus subfastigiatus is distinct from C. 
occidentalis in its pallid flesh-color to light cinnamon 
basidiocarps, forest green staining reaction with KOH, and 
smaller basidiospores (8-10.5 x 5-6 pm). Clavariadelphus 
caespitosus differs from C. occidentalis by its slender, 
narrowly clavate, greyish red to dull red basidiocarps 
with subacute apices, caespitose growth habit, and smaller 
basidiospores (8-11 x 4.5-6.5 pm). Clavariadelphus 
fasciculatus, known at this writing only from Mexico, is 
unique in its strongly fasciculate growth habit, pinkish 
orange basidiocarps, larger basidiospores (11-14 x 5-7 
jim), and acerose- to acicular-shaped crystals encrusting 
the walls of the mycelial hyphae. 

Small, immature basidiocarps of C. occidentalis might 
also be confused in the field with C. ligula and C. 
sachalinensis. These taxa can readily be segregated 
microscopically, however, by size and shape of 
basidiospores. Basidiospores of C. occidentalis are 
broadly ovate to amygdaliform in profile with an Fm<2.0, 
while those of C. ligula and C. sachalinensis are narrowly 
ellipsoid to boletoid in profile with an Em> 3.0. 


Clavariadelphus pallido-incarnatus Methven, sp. nov. 
Figs. 9—DPiGsks 


Receptaculum 7-15 cm altum, apice 10-15 mm latun, 
simplex, clavatum vel turbinatum; apex obtusus dein 
truncatus, rugosus, sterilis; hymenium laeve dein rugosvum, 
pallido-incarnatum, in KOH luteum; caro spongiosa, albida, 
brunnescens; odor nullus, sapor dulcis; sporae 9-11.5 x 
6-7 pm (L_ = 10.3 pms; W_ = 6.3 yim; E = 1.4-1.83 E_ = 1.6); 
late ovatae vel amygdall formes, laeves, albae; hyphae 
monomiticae, intertextae, fibulatae. Sparsa vel gregaria 
ad terram sub Sequoia sempervirenti, Picea sitchensi, et 
Alno spp. in America boreali occidentali. 


175 


Figs. 12-13. 12. Basidiocarps of Clavariadelphus 
occidentalis (Methven 5067). x 1/2. 13. Basidiocarps of 


Clavariadelphus pallido-incarnatus (TENN 47404, Holotype). 
6 ey as 


176 


HOLOTYPUS: TENN: United States, California, 
Humboldt Co., Prairie Creek Redwoods State Park, Davidson 
Road, 15 Oct 1986, legit A. S. Methven No. 5003 (TENN 
47404). 


BASIDIOCARP 7-15 ecm high, 5-10 mm diam basally, 10-15 
mm diam apically, simple, initially subcylindric, enlarged 
upward in age, then clavate to narrowly turbinate; 
mycelial hyphae interwoven or aggregated into rhizomorphic 
strands up to 1 mm diam, off-white to pallid; base terete, 
smooth, pruinose, yellowish white (4A2) to orange white 
(5A2), "pale ochraceous-buff," "light buff," "cartridge 
buff"; hymenium smooth, longitudinally rugose to rugulose 
in age, initially pale orange (5A3) to greyish orange 
(5B3), "seashell pink," "pale ochraceous-salmon," "pale 
pinkish cinnamon,'' finally flesh (6B3) to brownish orange 
(604-3), "light ochraceous-salmon," "light pinkish 
cinnamon," "light vinaceous-cinnamon"; apex initially 
obtuse to broadly rounded, more or less truncate in age, 
smooth, more or less concolorous with the hymenium; disc 
plane to depressed, not perforate or excavate; margin 
broadly rounded; surface staining slowly, irregularly 
brown (6D6-5, 6E7-6), “ochraceous-tawny," "buckthorn 
brown,'' "sayal brown," "tawny-olive" where cut or bruised, 
staining more conspicuously downward; flesh initially 
solid, becoming soft and spongy upward as the apex 
enlarges, white to pallid, on exposure staining slowly, 
irregularly brown (6E6-5, 7E6-5), "russet,"” "snuff brown," 
"rood's brown". ODOR not distinctive. TASTE slightly 
sweet. 


MACROCHEMICAL REACTIONS: NAP, PYR, GUA, FeCl.,, FeCl 
+ EtOH, PCR, PHN, ANO, SYR = positives; KOH = yellow; 
NH40H, TYR = negative. 


y 


MYCELIAL HYPHAE 2.5-4 yim diam, branched, clamped, 
uninflated; walls thin or irregularly thickened to 1 pm, 
hyaline in KOH, refractive under phase contrast, walls 
smooth, encrusted with acicular- to acerose-shaped 
crystals; crystals up to 8 x 1 pm diam, hyaline, insoluble 
in KOH, refractive under phase contrast, acyanophilous; 
clamps uninflated or inflated (-9.5 pm), sometimes 
medallion or ampulliform; hyphal contents amorphous, 
hyaline in KOH. TRAMAL HYPHAE 4-14.5 jim diam, more or 
less parallel to longitudinally interwoven basally, more 
loosely interwoven upward, radially interwoven beneath the 
subhymenium, uninflated, inflated (-19.5 pm) or broadly 


177 


undulate, branched, clamped; walls thin or irregularly 
thickened to 1 ym, hyaline in KOH, refractive under phase 
contrast, smooth; clamps uninflated or inflated (-19.5 
jim), sometimes medallion or ampulliform; hyphal contents 
amorphous, hyaline in KOH. GLOEOPLEROUS HYPHAE 5-9.5 jum 
diam, arising from generative hyphae at clamp connections, 
scattered throughout the trama, more abundant downward, 
uninflated or inflated (-12 pm), branched, clamped; walls 
thin, hyaline in KOH, smooth; clamps uninflated or 
inflated (-12 ym), sometimes medallion or ampulliform; 
hyphal contents subopalescent, yellow in KOH, refractive 
under phase contrast, cyanophilous. SUBHYMENIUM 
rudimentary; hyphae 2.5-5 pm diam, interwoven, branched, 
clamped, uninflated; walls thin, hyaline in KOH, smooth; 
clamps uninflated; hyphal contents amorphous, hyaline in 
KOH. HYMENIUM thickening, limited to the sides of the 
basidiocarp, composed of basidia and leptocystidia: 
LEPTOCYSTIDIA 60-85 x 2.5-6 jim, scattered among and 
scarcely projecting beyond the basidia, cylindric, 
subcylindric or strangulated, inflated apically at 
maturity, then narrowly clavate, at times apically or 
subapically branched, clamped; walls thin, hyaline in KOH, 
smooth; clamps uninflated; hyphal contents amorphous pale 
yellow in KOH, acyanophilous: BASIDIA 65-90 x 6-9.5 pm, 
narrowly clavate to clavate, inflated apically at 
maturity, then broadly clavate, clamped; walls thin, 
hyaline in KOH, smooth; clamps uninflated; basidial 
contents multiguttulate and refringent to aguttulate and 
amorphous, pale yellow in KOH; sterigmata (2) -4, 6-9.5 jim 
long, broadest basally, narrowed to an obtuse apex, 
incurved. APEX sterile: apical pellis a palisade of 
scattered basidia and sterile elements: STERILE ELEMENTS 
20-40 x 5-10 jim, fusoid-ventricose to ventricose, clamped; 
walls thin, hyaline in KOH, smooth; clamps uninflated; 
contents amorphous, hyaline in KOH, smooth: BASIDIA as 
described above. BASIDIOSPORES white in deposit, 9-11.5 x 
5.5-6-7 pm (L_ = 10.3 ppm; W_ = 6.3 pms E = 1.4-1.83 7 
EusrPe).6); broadly ovate to amygdaliforn; walls thin, 
hyaline in KOH, smooths; contents multiguttulate and 
refringent to aguttulate and amorphous, pale yellow in 
KOH; acyanophilous, inamyloid; hilar appendage oblique 
with an obtuse apex. 


HABIT, HABITAT, AND DISTRIBUTION: Scattered to 
gregarious; terrestrial; duff; under Sequoia sempervirens 
(Lamb.) Endl., Picea sitchensis (Bong.) Carr, and Alnus 
spp.; California. 


178 


SPECIMENS EXAMINED: UNITED STATES. CALIFORNIA, 
Humboldt Co.: Prairie Creek Redwoods State Park, Davidson 
Road, 15 Oct 1986, Methven No. 5003 (TENN 47404) 
[HOLOTYPE]; Prairie Creek Redwoods State Park, Rugg Grove, 
8 Nov 1986, Saylor No. 3732 (SFSU). 


The sterile, truncate basidiocarp apex, acicular- to 
acerose-shaped crystals encrusting the mycelial hyphae, 
and broadly ovate basidiospores place this taxon in subg. 
Clavariadelphus sect. Cantharellopsis, quite near C. 
truncatus. The pale coloration of the basidiocarps, lack 
of reactivity with KOH, and habitat in coastal coniferous 
forests of Sequoia sempervirens and Picea sitchensis 
distinguish C. pallido-incarnatus from C. truncatus. 


ACKNOWLEDGMENTS 


I am grateful for the assistance of Mrs. Ellen Thiers 
in preparation of the Latin diagnoses. The following 
people are recognized for assistance provided during my 
field work:- Drs. J. Ammirati; T.,Baronl, R. Bruce: 

J. Haines, R. Halling, D. Largent, R. H. Petersen, and 

H. D. Thiers, Mrs. Kit Scates-Barnhart, Mr. Dennis 
Desjardin, and Mr. Herb Saylor. I wish to thank the 
directors and curators of the herbaria listed under 
"specimens examined" who kindly provided materials for 
study. For helpful comments and suggestions I am indebted 
to DreiR 2H. Petersen: 


LITERATURE CITED 


Corner, E. J. H. 1950. A monograph of Clavaria and 
allied genera. Ann. Bot. Mem. I: 1-740. 

1970. Supplement to, A monograph of 
Clavaria and allied genera. Nova Hedw. Heft 33: 
1-299, 

Donk, M. A. 1933. Revision der Niederlandischen 
Homobasidiomycetae -— Aphyllophoraceae II. Meded. 
Bot. Mus. Rijksuniv. Utrecht. 9: 1-278. 

Holmgren, P.. K., W. Keuken, and E. K. Schofield.) @igei 
Index herbariorum. Part I. The herbaria of the 
world. 7th ed. Regnum Veg. 106. 

Kornerup, A., and J. F. Wanscher. 1978. Methuen handbook 
of colour. 3rd ed. Eyre Methuen, London. 252 pp + 
30 Spi 


179 


Marr, C. D. 1979. lLaccase and tyrosinase oxidation of 
spot test reagents. Mycotaxon 9: 244-276. 

-, and D. E. Stuntz. 1973. Ramaria of western 
Washington. Biblio. Mycol. Band 38. Lehre, 
Germany. 232 pp. 

Petersen, R. H. 1972. Notes on clavarioid fungi. XII. 
Miscellaneous notes on Clavariadelphus, and a new 
segregate genus. Mycologia 64: 137-152. 

sete Ranmeandel.swoCchitd.selo/ 4,8 Notes, on 
clavarioid fungi: Three new species of 
Clavariadelphus. Trans. Brit. Mycol. Soc. 63: 
469-474, 

Ridgway, R. 1912. Color standards and nomenclature. 
Publ. privately, Washington, D. C. 44 pp + 53 pls. 

Singer, R. 1986. The Agaricales in modern taxonomy. 4th 
ed. Koeltz, Koenigstein. 981 pp. 

Smith, A. H. 1971. Some observations on selected species 
of Clavariadelphus. Mycologia 63: 1073-1076. 

Wells, V. L., and P. E. Kempton. 1968. A preliminary 
study of Clavariadelphus in North America. Mich. 
Bote / 2735-574 


mitt 


i An Mies, We, yey p d 
phiee ad 5, he 
AR Aaa abs a 
, ait i ee ih M4 e b a 
wy A EM ie age: % 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 181-196 January 20, 1989 


KEY TO THE SPECIES OF INOCYBE IN CALIFORNIA 


FLORENCE H. NISHIDA 


Botany Section, Natural History Museum 
900 Exposition Blvd., Los Angeles, CA 90007 


SUMMARY 


A key to the species of Inocybe found in 
California is presented, based on a_ recent 
systematic study of the genus in southern 
California. The occurrence of nearly 60 
species is indicated LoL the state, 
augmenting the 16 nominal species previously 
reported in the literature. Several species 
with type localities in California or the 
west coast of the U.S. were re-collected and 
determined for the first time since their 


original description. Distinct Inocybe 
floras appear in northern and southern 
California, associated with different 


forests, mixed conifer vs. live oak, 
respectively. Less than a quarter of the 
total species occur in both regions. A 
checklist is provided of nominal Inocybe 
species of California, based on published 
reports and herbaria records at SFSU and LAM. 


The genus Inocybe has been poorly studied in most of 
North America, some conspicuous exceptions being areas near 
New York (Peck, 1872-1910), Michigan (Kauffman, 1920, 1924, 
1925; A.H. Smith, 1939; Stuntz, 1954), Washington (Stuntz, 
1940, 1965; A.H. Smith & Stuntz, 1950), Florida (Murrill, 
eeiem 1944,/..1945), and Nova Scotia, Canada (Grund & 
Stuntz, 1968-1984). A recent systematic study of Inocybe 
in southern California (Nishida, 1987, and ms. in prep. for 
Contributions in Science, Natural History Museum of Los 
Angeles County) revealed that the genus is surprisingly 
well-represented in semi-arid southern California, in 
relatively moist, coastal woods of live oak (Quercus 
_agrifolia). At least 26 smooth-spored species and 5 
nodulose-spored species are reported from southern 
California. Voucher specimens are deposited in herbarium 
LAM (Natural History Museum of Los’ Angeles County, 
California). It is worth noting that most of these 
nodulose species occur in inland, montane conifer woods of 


182 


Pinus ponderosa, P. jeffreyi, and Abies concolor. 

My study of the herbarium specimens at San Francisco 
State University (SFSU) indicates a northern California 
Inocybe flora of at least 26 smooth-spored species and 13 
nodulose-spored species. Many of the Inocybe collections 
had been annotated by Daniel Stuntz in 1971, but many other 
collections had not been determined, nor examined 
microscopically and none had been reviewed in light of 
recent taxonomic revisions of the genus. Nearly 60 species 
of Inocybe are presently reported from California, in 
contrast to 16 nominal species previously reported in the 
literature (Earle, 1904; Peck, 1909; Kauffman, 1924; 
Murrill, 1916; A.H. Smith, 1939, 1941; Smith & Stuntz, 
1950; Grund & Stuntz, 1970). 

Based on collections in the herbarium at SFSU there is 
evidence of a contrasting Inocybe flora in northern 
California. This abundant collection represents’ the 
mycological activities of Harry D. Thiers and his students 
over many years in the mixed conifer and conifer-hardwood 
(Pseudotuga menziesii, Abies grandis, Picea sitchensis, 
Quercus chrysolepis, Q. kelloggii) forests of northern 
California. Although the two regions (northern and 
southern California) appear to have the same number of 
species of Inocybe, a closer examination shows that there 
are two distinct floras, with some overlapping of species. 
In general, nodulose species are more to be _ found 
associated with conifers. The northern California 
collection, mostly of the mixed conifer region, has 13 
nodulose spored species, compared to 5 in southern 
California, where most of my collecting has been in the 
live oak woods. Nearly half (48%) of the total species 
occur only in northern California; less than a third (30%) 
occur only in southern California. Less than a quarter 
(23%) of the species are found in both regions. Among 
these latter are species of wide distribution in the world 
(I. fraudans, I. geophylla, I. leptocystis, I. rimosa, I. 
mixtilis, and I . variabillima). 

California is an area of great floristic, topographic, 
and climatic diversity. Sampling of the live oak woods of 
the southern coast and the pine and fir woods of the 
northern coast has yielded nearly 60 species of Inocybe. 
Large areas, potentially rich in Inocybe species, remain 
unstudied, especially the stands of Quercus kelloggii, and 
Q. chrysolepis on the lower slopes of the Sierra Nevada 
range, and of Q. douglasii surrounding the central valley, 
as well as much of the conifer forest extending along the 
Sierra Nevadas. 

Recent taxonomic studies by Thomas W. Kuyper, including 
a revision of European smooth-spored species (1986), have 
provided useful information on some particularly difficult 
"species complexes," as well as updating nomenclature. My 
own recent systematic study of the southern California 
species of Inocybe has resulted in some taxonomic 
revision. Both these works should be consulted for 
complete descriptions and synonymy of species, as well 
as Nishida (1988) for new species, reported in this key to 
species of Inocybe in California. 


183 


Key to Subgenera, Sections, and Subsections of Inocybe * 


1. 


Pleurocystidia absent; cheilocystidia thin-walled, no 
crystals at apex; spores alwayS SMOOth..cccccccccccseed 


2. 


3. 


Sis 


Cheilocystidia originating from the hymenophoral 
trama; basidia with necropigment; stipe usually 
Shorter than diameter of plleusii.. cise vec cccosccee 
Cece ccccvcccccccccccccccccceee SULGEN. Mallocybe 


Cheilocystidia originating as modified (sub)hymeni- 
al elements; basidia without necropigment; stipe 

usually longer than diameter of pileusS......ccccenee 
cece cere cer ere cccreccccccccveceesecoubgen. Inosperma 


Pileus squamulose to squarrulose; context often 
reddening; basidia slender; odor often distinct... 
eeeccovcccsicvccesesovocsceses eect. Cervicolores 
Pileus radially rimose or rimulose, not squamu- 
lose or squarrulose; context reddening or not; 
basidia broader; odor mild or spermatic....ccceee 
Slee vice cc es ecco scoccccsocecceevcscs eSect. Rimosae 


Pleurocystidia present, almost always thick-walled and 
with crystals at apex; cheilocystidia of two kinds, 
pleurocystidioid and clavate to (sub)globcse para- 
cystidia; spores smooth, angular, or nodulose in out- 
TINE]. cc cccccccccccccccveccccccccccescessoubgen. Inocybe 


4. 


Spores smooth in outline (subangular at times in 

I. lacera): ovate, ellipsoid, (sub)amygdali- 

EOrm me CSS, OCTON@ (SUD) FENLLOLM ic stciele's%s ae ore lee ec ecco 6 
Cec cc ccc cc ccccccccccccvccecccceenect. Inocybium 


5. Caulocystidia present over entire length of 
stipe (occasionally scarce in lower 1/3, or 
difficult to find or interpret) as thin- or 
thick-walled metuloids or thin-walled clavate, 
cylindrical cells of various sizes, or combi- 
nation of these; cortina absent in young stage 
Wise ote clelcisis ciclo sicls o cts vols pDUDSECL. Holoconiatae 

5. Caulocystidia present at apex of stipe only, 
occasionally rare or absent (sometimes with 
cystidioid terminal cells at the base, but not 
considered true cystidia); cortina often 
present in young SPECIMENS... ccrcccccccccscvcceres 
Sieleleiclclsisleieisicisicie ete sleleiciese DUDBECU ns ACrOCOnNiaAtae 


Spores generally angular, or coarsely to definitely 
nodulose to stellate in outline. .cccscocccciscccsecer 
slatateta clove Glela eo atévavele 's) clelele¥evelelelio etelsteusistemeClus Inocybe 


* The infrageneric classification follows Kuyper (1986). I 
concur 
smooth-spored and nodulose-spored species. 


that this is a phylogenetically valid grouping of 


184 


6. Caulocystidia at apex of stipe only, or absent; 
cortina present in young Stage...eccccccccccece 
aje'eeje.0 cleleeiela ele slelensisielneie « 6UUBG Chaat sauna 

6. Caulocystidia over entire length (including 
lower 1/3); cortina absent in young stage...... 
ces ces cwices clea secccce cies  eOUDSECEs MALMAIe une 


Key to Species of Subgenus Mallocybe 


Pleurocystidia absent; cheilocystidia thin-walled, no 
crystals at apex; spores always smooth; cheilocystidia 
originating from hymenophoral trama; basidia often with 
necropigment. 


1. Small basidioma, pileus 10-25 mm, wooly-tomentose to 
coarsely scaly-squarrose, color fulvous to cinnamon- 
brown, context pallid to amber, purplish then brown in 
KOH; lamellae (sub)distant, margin pallid or concolored; 
cortina sparse; stipe sparsely to obviously fibrillose; 
cheilocystidia articulated, thin-walled, clavate; 
spores ovate to subreniform with distinct suprahilar 
depression, 10-l1l x 5-6.5 pm; under Salix in wet 
PLACES. ce ccccvccccccccccvcccevcccvvcccvccele Aulcamara 


1. Larger basidioma, pileus 20-60 mm, appressed-fibrillose 
but not scaly or wooly-tomentose, ochre to brownish 
ochre; lamellae pallid at first, often becoming con- 
colorous; stipe fibrillose peronate with distinct, 
thick cortina; cheilocystidia thin-walled, clavate; 
spores elliptical to subreniform, 8-10 x 4.5-5.5 jam; 
UNder CONLLEFS.ccccccccccccccccccvccvcveeeele agardhii 


Key to Species of Subgenus Inosperma 


Pleurocystidia absent; cheilocystidia originating as modi- 
fied hymenial elements; basidia without necropigment. 


1. Pileus diffracted scaly on disc, brown, isabella to 
umber, squarrulose to squamulose in margin; stipe long 
(25-80 mm), pale brown at apex, darker at base, often 
with grey-green, blue-green to olivaceous blackish 
stains, surface coarsely squamulose; context reddening; 
odor sweetish, like Pelargonium, or fishy.....cccccoee 
(Sect. Cervicolores, one species).....1. calamistrata 


1. Pileus and stipe not scaly, squarrulose or squamulose... 
(Sect. Rimosae) dja je ee jes 0.0 6 610,06 0.6 0 8 0.01410 of0ie.a lee lereniene 


2. All parts of basidioma, including context, changing to 
vinaceous, pinkish red, or vinaceous brown; distinctive 
odor lacking; spores 10-12.5(-15) x 6-7(-7.5) pm..... 
Cocco rece r sc cesecccreseceseceseccseccoeseeele adaequata 

2. Basidioma not changing as above.....«.+«+esces acini 

3. Pileus buff, ochraceous, yellow-brown to fulvous.....4 
4. Stipe with blue-violet, gray-blue tints.....ccecccceeD 
5. Spores (sub)reniformy) > 6-pm broad. «.'ss «sole sal aiaenie 


185 


Cec occescesccevesccccccescceseevele vinosistipitata 


5. Spores elliptical, < 6 pm broad; stipe gen. pale 


yellow, occ. violet..ccccccccceccccoeele Quietiodor 


4. Stipe white, pale yellow, becoming more yellow....... 
6. Pileus often pallid overall at first, due to heavy 


covering of white velipellis (particularly in 
southern California husky, inodorous forms), in time 
becoming rimose and also yellow, yellow-brown to 
fulvous; odor absent, spermatic, or of fresh corn; 
spores 10-13 x 6-7 pm, (Sub)reniform....secccccccces 


co cecrcrcacvccecsercecccevscvsesceccccesoele rimosa 


6. Pileus with conspicuous central white patch of veli- 


3. 


pellis, drying entire, rarely, in squamules; umbo 
very obtuse or absent; stipe wax colored, smooth, 
equal or swollen, never emarginate-bulbous; spores 
9-10 x 5-5.5 pm, ellisoid, fewer subreniform; odor 
lacking or faintly rancid-mealy......I1. brunnescens 
Pileus brown, reddish brown, chestnut, russet, to 


umbrinous, lacking conspicuous velipellis; silky 


smooth; odor mild or faintly spermatic; spores 9-10 x 
5-6 pm, mostly ellipsoid with obtuse apex..cccccccvce 


Dialavsls oleic seis cele p elec se clscieseecciosecceec eel fastigiella 


Key 


to Species of Subgenus Inocybe, Section Inocybium 


Spores smooth (ovate, ellipsoid, subamygdaliform, sub- 
reniform; pleurocystidia present. 


Key to Species of Subsection Holoconiatae 


Caulocystidia found along entire length of stipe. 


1. Entire basidioma reddening with age and/or on bruising, 
pure white at first; stipe base bulbous, sub-emarginate; 
odor spermatic; spores (10-)11-12.5(-15) x 6-7.5 pm, 
subamygdaliform to sublimoniform, apex subconical....... 


plaints is leie © © 6 ele cle. 0 s 0) 06 oe 0 em0lee'e e see 0 ces 06 0106 0 0 eke godeyi 


l. Basidioma not reddening with age...ccrccccccvcccccccseed 
2. Stipe incarnate (some shade of reddish to orange, 
salmon, pinkish tan, reddish brown, reddish orange, or 
orange ochraceous), at least at the apex..ccrccccescvved 


3. 
4. 


OACOr Of bitter AaALlMONG. i.e sece cle sees cccesecceccescnseeS 
Odor of bitter almond in lamellae, spermatic in stipe; 
pileus ochraceous to ochraceous brown, radially 
fibrillose, sometimes becoming scaly; stipe orange 
ochraceous, not distinctly reddish; pleurocystidia 
cylindrical-clavate, to fusiform, moderately yellow 
GN"KOH. sess ccc cseccoeeeols Nirtella var. hirtella 
Odor weakly of almond, or else spicy as Pelargonium, 
frequently spermatic; pileus ochraceous brown to 
fulvous, tending to become squarrose-scaly around the 
disc; stipe often short, distinctly reddish brown at 
apex, or becoming purplish; pleurocystidia with 
bright yellow walls in KOH, fusiform to sublageniform 
slo ewecsicescvesesotsceoevvsevecevescssecelse Muricellata 


Sr eWLLNOUcTGLStiNnct Odoreoh bitter Amon ile ccc cD 


186 


5. Pileus bright colored (orange-brown, "rusty," tawny, 


Ful Vous ) os.c ce c'0 vle.0 ce 0 6.006 010 6.0 016 ¥ 0 bale 016s 6) ate Meare ane nee 


6. Pileus often bicolorous, bright, tawny brown to red- 


brown on disc and ochraceous to honey in margin; 
stipe salmon-incarnate, context incarnate or pinkish 
orange; pileus and stipe often with tawny to orange 
fibrils; pleurocystidia extremely abundant, with 
very thick walls (4.5-5 pm); spores 9-11 x 5.5-6 pm 


Coe rcccccccccccceecccccccccecccsccceccsoole laetior 


6. Pileus unicolorous, bright orange-brown to fulvous 


orange; stipe pale orange-incarnate, context white; 
spores 8.5-9.5 x 5.0-5.5 pm... ececcceceeele Vaccina 


5. Pileus not brightly colored, definitely brown.......7 


7. Pileus tending to become squarrose-scaly around 


disc, without obvious velipellis, ochraceous brown 
to fulvous; stipe distinctly reddish brown, some- 
times becoming purplish; pleurocystidia bright 
yellow in KOH, thick-walled; odor spermatic, 

rarely, faintly spicy or of almond...ccccccccvscceccs 
weccccnecoscceceesecoesevescesceveceee Le MUL tCeLLaue 


7. Pileus. smooth, uniformly brown....ccccsccscceevscves& 


8. Pileus ochraceous brown, chestnut-brown, sometimes 
greyish on disc from velipellis; stipe reddish 
brown to orange ochraceous, with marginate bulb; 
pleurocystidia thin-walled, hyaline; spores rather 
small (7.5-9.5 x 4.5-5.5 ym), mostly ellipsoid..... 
ccc r cece cc cece cccccvecccccoccccccsecele amblyspora 

8. Pileus dark reddish brown to mahogony brown, silky 
smooth, greasy appearing, velipellis absent or 
scanty; stipe incarnate, pinkish buff to fleshy 
ochracaceous, without bulbous base; pleurocystidia 
thick-walled; spores larger (8.5-10 [-12] x 5.5- 
Go5 WM) ceccveccccccvcccccsccescvceeceele Leiocephala 


2. Stipe white, pallid, or variously colored, sometimes 


9. 


9. 


distinctly darkening, but nowhere incarnate.....ccccee9 


Spores generally >11 pm long (10-12 [-16] um), 
ellipsoid to subamygdaliform; robust habit; pileus 
buff, tan, dull ochraceous to ochraceous brown, paler 
in center from thick velipellis; husky stipe (30-80 
mm long), with subbulbous to emarginate bulbous base, 
often with grey-green stains; pleurocystidia variable, 
often subcapitate; thin-walled clavate caulocystidia 
sometimes present at DbaSe...ccceccecceeeele bakeri 
Spores generally <1ll pm long. ..ccccccccccccccvcccceslD 


10. Pileus light (cream, isabella brown, buff, tan, dull 


ochraceous, ochraceous brown); stipe not distinctly 
Garkening...c«ccce cece s clvcccecs ese s ce 0.0 6c nlemieninunneee 
ll. Pileus bright, yellow to golden with fulvous disc 
when fresh, becoming entirely orange brown when 
dry; velipellis absent; stipe pale yellow; 
pleurocystidia fusiform with very thick (4 um), 
pale to moderately yellow wall; spores amygdali- 
form to sublimoniform 10-11 x 5.5-6 Mim... ccccccecs 
cece ccereccccccscccsccecscceeceeele Chrysocephala 
11. Pileus pallid, not bright; stipe long, pallid...12 
12. Habit robust (pil. 10-60 mm); pileus cream to 
isabelline, ochraceous, fibrillose-scaly at 


187 


disc; stipe long (40-80 x 4-10 mm), even to 
clavate at base; spores 8-10 x 5-6 pm, sub- 
amygdaliform; pleurocystidia lageniform to 
ruUsBOLG—Ventricose, sabundants's ss «sisies cles cicleeis sc 0 
Corre e recor ccceccccccscecevccvcceeele Kauffmanii 
12. Habit regular (pil. 38 mm); pileus white to buff 
silky-smooth; context light brown under pellis; 
acrid taste; spores 9-10 x 5-5.5 pm, elliptical 
to subreniform; pleurocystidia short, very 
rare, Clavate to subfusiform, with extremely 
EVLCKe WALL Se 4-55.57 UM), Valine. o-alciele cletele.cia ess 
ec erececcccccccecesccvcccccecvelte menthi-gustans 
10. Pileus definitely brown, sometimes appearing bi- 
colorous; stipe darkening in lower half; odor of 
Pelargonium or of Amanita phalloides, or else 
MOGI LUC siete ts ererstalatere! dlelsierers/tie'e ele elite lelatete ts cisiele csc alelete oLic 
13. Habit robust (pil. to 70 mm); stipe distinctly 
darkening to dark orange-brown, dark reddish brown 
or blackish brown; pileal disc light (yellowish 
brown), margin dark (cinnamon-brown to dark red- 
brown), uniformly brown when dry; spores 
ellipsoid, subamygdaliform, 8.5-9 x 5.0-6.0 pm.... 
. secre crs ccccccccescccecccccecscoseveele Splendens 
13. Habit slight (pil. to 55 mm); stipe not distinctly 
darkening, but discoloring with handling (tan to 
orange brown); disc vinaceous to chestnut-brown, 
margin reddish brown, uniformly russet when dry; 
spores 9-10 x 5.5-6.0 mm, amygdaliform, conical 
APCKececcccccccsecccccccccesccccssveceolts Phaecoleuca 


Key to Species of Subsection Acroconiatae 
Caulocystidia found only in upper 1/3 to 1/2 of stipe. 


1. Basidioma generally white to cream and remaining so or | 
Meromingetinted: with Other COLOrS...cecc ce cieiceecsiccsceed 
2. Slender habit (cap:stipe = <0.60); pileus 10-35 mn, 
silky-smooth to fibrillose, conical or convex, umbonate 
sometimes subviscid; odor spermatic...ccccccccccccccoed 
3. Staining orange-red with time and/or damage, lubricous 
when wet, Shiny.-.ccccccsccccecvceel. whitei f. whitei 
SENOS Ca LN 1G FOLAaNngGe—Le selec sisleis cies le cielslelcleléieleiaieue s siclclee 4 
4. Pileus with violet or lilac colors, especially in 
young specimens......eeeeeele geophylla var. lilacina 
Aeeeseusewiathout violet on lilac colors}. .< <ecsiecececisD 
5. Remaining more or less silky white, or becoming buff 
iN AGE. erececececveveeele. Geophylla var. geophylla 
5. Not remaining white; disc of pileus fuscous, margin 
paler, and base of stipe also fuSCOUS...ccccccceeces 
cece crc cccveresvccsccccccccccccescsecel. fuscodisca 
2. Stocky habit (cap:stipe = or >0.60), resembles Hebeloma 
pileus 20-50 mm, convex, exumbonate; dry or subviscid 
in wet weather; white to dingy buff, becoming matte 
buff to tan, never silky-fibrillose; odor lacking...... 
wee cece cr ccc ccecccsecccccccccccccccccccccele insinuata 


188 


1. Basidioma colored from the first (ochre, yellow-brown, 
fulvous, reddish brown, umbrinous, €tc..)% 0. ses asleeeee 
6. Basidioma having a distinct sweet, aromatic, or fruity 
OGOL sd. bes. 6 6:b'e.0 6 eee ehoin (6 6ie.e a #0616 aie. 6.6 loleue. sais tae 
7. Pileal disc, and/or some part of the stipe with green 
(greyish green to dull olive-green), pileus often 
scaly; context whitish, not reddening; strong odor of 
"Peruvian balsam" or cinnamaldehyde..ccccccccccccecces 
diel cole a s cle 0's suis o els eins sicdie COLVORLINA VAL oo am mmanneee 
7. Pileus smooth, color pale ochraceous, pale fulvous, or 
mixture of yellow, red, dark russet, umbrinous, 
vinaceous brown; context pallid, reddening upon 
exposure; stipe becoming brick red then vinaceous; 
strong odor, variously described as methyl cinnamate, 
"Peruvian balsam," or "matsutake".........-l. fraudans 
6. Basidioma lacking a distinct aromatic, sweet odor, or 
else with a spermatic! Odor. «2 .csccceccse cele cipie eiuialeatetng 
8. Violet or lilac shades in some or all parts: pileus, 
lamellae;: Stipe. oe oceecjs0 aces eves ces cee cs 0 sieislsieewiees 
9. Pileus small, 10-35 mm, ochraceous brown, smooth, 
in age becoming diffracted-scaly in margin; lamellae 
briefly violaceous; stipe equal to subbulbous, often 
violaceous in lower portion; pleurocystidia fusoid- 
ventricose, hyaline; spores subamygdaliform, 
765-1005 x) 4.55.5 pum. ne nec cle cece cn ccle sisi aan Due 
9. Pileus large, 15-50 mm, dark brown, umbrinous to 
reddish brown, with violaceous tinges from underlying 
context, distinctly scaly to tomentose-squamulose; 
stipe equal with brown fibrillose punctae, streaks, 
or squamules, especially toward base; pleurocystidia 
slenderly fusiform, with thick, bright yellow walls; 
spores amygdaliform with subconical to papillate apex 
8.5-10 x 5-5.5 pm..eeeeeeeeele phaeocomis var. major 
8. Violet or lilac shades absent and pileus pallid, 
brightly colored or Drown. 2... ses ce ce asi e ciue 6s ute tsterenealt 
10. Pileal colors pallid, ochraceous, to ochraceous 
brown or brightly colored, not definitely brown..11l 
11. Spores = or > 11 wm (10.5-13 x 5.5-6.0 wm) subamyg- 
daliform; pleurocystidia broad, utriform to fusoid, 
thick walled, hyaline; pileus 25-55 mm, pale ochra- 
ceous to ochraceous brown, smooth, often with dirt 
or litter adhering; stipe white, husky (50-80 x 5-15 
mm) with bulbous to emarginate base; differentiated 
caulocystidioid hairs sometimes found on base of 
stipe; odor lacking. .cccccccccccccccccceslte bakeri 
11. Spores <LU pm wecccceccccccsccccsvesce oe sisinia cere 
12. Pleurocystidia long and narrow (60-90 mm), thick 
walled, pale to bright yellow; spores 7.5-9 x 4.5 
-5.0 pm, subamygdaliform; pileus isabella, greyish 
ochraceous to ochraceous brown; odor spermatic and 
FALINACEOUS. .crccceccccvccceececsesele SUDOCHLTaCea 
12. Pleurocystidia short (40-60 um), fusiform to 
utriform, with pale to bright yellow wall; spores 
8.3-9 x 4.5-5.5 pm, subamygdaliform; pileus yel- 
low, pale cream to ochraceous yellow ..cccccevccce 
Cece err ccc ccc vce rccccccccccvcccccsccsele aUricoma 
10. Pileus definitely brown; basidiocarp rarely with 


189 


violaceous colors ehetetelele =: akelals lalerslsletulelele siateleteies 4 e/eleetlis 


US eS 


vege 


Basidioma with striking lamellar colors (bright 
orange to ochraceous tawny); also pileus orange to 
cinnamon at first, becoming cinnamon to "Sayal 
brown," context orange-cinnamon; stipe with heavy 
covering of cinnamon: colored fibrils, context and 
surface cinnamon, becoming brownish, spores sub- 
amygdaliform; pleurocystidia fusoid-ventricose.... 
cece ccc c cere cccrccsccvecceveccesecevels Cinnamomea 
Basidioma without striking lamellar color, but 
stipe may be incarnate (with pinkish, pinkish 
brown, reddish orange, reddish brown) or not....14 


ai, Stipe PCAY NAL Ge Gicki ic chalesure rote elals eieloierslalvveletohe wie re ete LS 
15. Pleurocystidia uniquely thin-walled, without a 


double membrane, clavate to utriform with wavy 
Margin, apex obtuse or subcapitate; spores 
subamygdaliform, 8-9 x 5-6 pm; pileus red-brown, 
"bay," sometimes fading to "tawny," at center or 
margin; odor faint, raphanoid to farinaceous... 


Bele lsiecdlolsielel's taialeiaie Sielele efajere alelelelevals e.elkve leptocystis 


15. Pleurocystidia definitely thick-walled........16 


14. 


POemODOLeS Generally ao) lpm liciiisletete egiesmreisieleie sie L 7 
17. Spores elongate to slightly angular, 11-14 x 
4.5-5.5 pm; pileus coarsely fibrillose, 
tomentose, squamulose to squarrose; stipe 
buff at apex, sometimes with pinkish tinge; 
brownish toward base, fibrillose..I. lacera 
17. Spores never angular, but broad, 11.5-13.5 x 
6-7 um; pileus orange-brown to cinnamon - 
brown with conspicuous and persistent white 
patch of velipellis in center; smooth to 
slightly diffracted scaly on disc; stipe 
light brown with reddish to orange tinges, 
with submarginate bulbouS baSe..cccccccccoes 
cece ccc cecccccccccssscccesccvecle Hemileuca 
POSE ODOLCS in < Lie AiM ls ia lalate, ole aieteleleie | ele tele le ete.elelsibinie levels « LS 
18. Pileus with disc distinctly darker than 
margin (disc umber to fuscous, sometimes 
ochraceous brown, outer whitish to cream); 
surface smooth to appressed-scaly to 
diffracted scaly; stipe densely fibrillose, 
reddish brown context and surface, caulo- 
cystidia at apex very rare; spores 
9-10 x 5-5.5 pm; pleurocystidia subcylin- 
drical to subfusiform, walls moderately 
thick (1.5-2 pm)....ececcceeeeste phaeodisca 
18. Pileus unicolorous, dark brown, rufous to 
umbrinous, margin sometimes tawny olive; 
pleurocystidia ventricose, very thick-walled 
(3.5 pm); spores 10 x 5-5.5 mm, subamyg- 
GALLIOLN te OGOTNSDEXMALI Coie ss le eleit ios lavelalelelelielet ails 
eer ccccccccrccceccecccveveele Subdestricta 
Stipe not incarnate; pileus pale brown, ochraceous 
brown, fibrillose-squamulose especially over disc, 
often recurvately squamulose-squarrose; stipe 
white, becoming pale ochraceous or pale brownish; 
spores 8.5-9.5 x 5-6 pm, subamygdaliform; pleuro- 


190 


cystidia fusoid-ventricose, with thick (3.5 pm), 

pale to bright yellow wall; caulocystidia at stipe 
apex sometimes rare; odor spermatic-acidulous..... 
ceesevedcceeseesoceceseseseeee sce cele smh Ce mune 


Key to Subgenus Inocybe, Section Inocybe 


Spores angular, angular-nodulose, or obtusely to definitely 
nodulose to stellate in outline; pleurocystidia usually 
present. 


Key to Species of Subsection Marginatae 


Caulocystidia along entire stipe; stipe pruinose overall; 
cortina lacking or attached to base of stipe which is 
frequently bulbous. 


1. Basidioma reddening and with pleasant, aromatic, fruity 
or sweet odor; pileus fulvous to yellowish brown; spores 
7-9 x 5-6.5 pm, bluntly nodulose; pleurocystidia fusoid 
ventricose, thick walled........e.eeeeeeeele bresadolae 


1. Basidioma' not reddening .....acesccccscc eee see cleleretatunenae 
2. Stipe incarnate or becoming tinged with pink, orange, 
SALMON « « 0.4 6 0000.00.00 06000 nec ces 505000 ves ss 0 ele a Mamisielalane 
3. Pileus bicolored, white to pale cream on disc, grey- 
vinaceous brown, or pale pinkish brown in margin; 
pellis smooth to minutely squamulose; spores 6.5-8 x 
4.5-6 pm, with 7-9 nodules; pleurocystidia ventricose 
with thick wall (5-6.5 pm); odor spermaticC....ccccceee 
Cece crc c crc cccv eres ccccercccccesecesceeeele Albodisca 
3. Pileus red-brown, cinnamon-brown, glabrous; stipe 
with marginate bulb, color gradually darkening; spores 
9-11 x 5.5-8 pm, with 8-10 large, prominent nodules; 
pleurocystidia broadly ventricose to subglobose....... 
cece ccc cccccccccccccccccccccvescesesele Oblectabilis 
2. Stipe not becoming incarnate...ccccsccescecics cies cenisisnne 
4. Basidioma entirely white, remaining pallid...........5 
5. Pileus 25-50 mm smooth, greasy to subviscid at first, 
in time pale ochre-yellow; stipe base not distinctly 
bulbous; spores 8-10 x 5-6 um, ellipsoid-oblong with 
7-9 coarse nodules; pleurocystidia fusoid ventricose, 
thick walled (2.5-3 pm); strong odor of Cortinarius 
PUFPUFASCENS ceoccecccccccccccccccvevsccevveele Olida 
5. Pileus 15-30 mm, dry, fibrillose, becoming 
obscurely scaly, white becoming dingy to ochraceous; 
Stipe base bulbous to submarginate; spores 7-10 x 5- 
6 pm, polygonal, 4-9 nodules, or merely angular; 
pleurocystidia distinctly thick walled (4-5.5 mm), 
short (30-55 pm), fusoid to clavate. .\.. «os ss smwitanenee 
Cer cece reer ever ccrcrosccocecesevccceseele umbratica 
4, Basidioma not white, but variously colored.........+-6 
6. Stipe noticeably darkening or blackening....-.cccccee? 
7. Pileus 20-35 mm, straw-yellow to brownish yellow, 
often paler in margin or disc from velipellis, moist 


191 


to subviscid; stipe pale dingy straw color, 
darkening to gray or black on drying; spores 9-12 
x 7-9 pm, with 9-12 large, prominent nodules; 
pleurocystidia lageniform, ventricose, with thick, 
bright yellow wall ....ccecccccccceeeele xanthomelas 
7. Pileus 15-60 mm, pale brown to grayish brown, 
surface becoming appressed-fibrillose scaly; stipe 
pale yellowish, blackening on drying; spores 9-10 x 
7-9 um, oblong, with 8-12 large, prominent nodules; 
pleurocystidia slenderly ventricose with long neck, 
PaCheWwawLeGs (3 60-> ELM) sen VALLNG /elstetciece se sicicicelcle's #166 
cere cree crccccccceccecccccccceecceceele Nigrescens 


6. Stipe not noticeably darkening or blackening .......8 


8. Pileus straw, ochraceous brown to fulvous in margin, 
with yellow disc, 15-40 mm, silky to lubricous; 
lamellae grayish pallid; stipe pallid, densely 
pruinose, with marginate bulbous base; spores 8-10 
SD. DO um, with 9-10 prominent nodules; 
pleurocystidia 40-55 pm long, broadly ventricose, 
Very thick walled (2-5 jim), yellow. .cecscccccccevcce 
Coe ercecercercceesceccceccsesccecccecceele mixtilis 

8. Pileus ochraceous brown to cinnamon-ochre, covered 

~ with greyish to grey-brown veil, smooth or greyish 
brown, scaly, 45-65 mm, umbonate, subviscid, dirt 
often adhering; stipe pallid, becoming yellowish 
brown; spores very large, 9-13 (-15) um, scarcely 
nodulose, polygonal; pleurocystidia ventricose to 
clavate, thick walled (3 um) .........-L. decipiens 


Key to Species of Subsection Cortinatae 


Caulocystidia lacking on base of stipe; stipe not pruinose 


all 


over; cortina present in young specimens; stipe base 


bulbous or not. 


1. Pleurocystidia lacking or extremely rare or difficult to 


WaRRE TP TRMUMETOST Woven Ve". Te cc og 6 oe ois te 6. ele ere eteT el eis.4 eles elere 6 #08 6.6 obo ele e 


2 
3 


Zig 


Pileus brown, tomentose-scaly, to squarrulose-scaly...3 
Pleurocystidia lacking; only thin-walled subglobose 
cheilocystidia; pileus dry, disc densely covered with 
Minute squarrose scales, margin smooth, walnut-brown 
to umber; stipe floccose fibrillose to tomentose, 
concolorous; spores 10-12 x 8-9 pm ellipsoid, with 
12-20 prominent subconical nodules; in damp woods, on 
FOCtEN WOOd..cecreccccvccecvecccevecseeele. Leptophylla 
Pleurocystidia short (25-45 pm), difficult to locate, 
ovate; pileus and stipe dark brown, densely squarrose 
to scaly overall, with pale zone below lamellae; 
spores 8-10 x 5.5-7 um, with 8-10 small nodules; in 
conifer woods and on rotten wood .......1. lanuginosa 

Pileus not strongly squarrulose-scaly; ochraceous, 

fulvous brown, reddish brown, fibrillose to finely 

appressed-fibrillose scaly; stipe pallid to tan, 
glabrous; spores 9-11.5 x 5.5-6.5 mm, oblong, 
notable by their extreme variability, from merely 
angular to angular-nodulose, to nodulose, but with low 


192 


nodules; pleurocystidia rare, short, 40-55 x 18- 30 pm, 
with distinctive, ovate or utriform shape, often with 
mucronate apex, wall not thick (1.5-2 pm) .cccccccccce 
wie elec cccccieccescescces ceecieisies¢cs 6.c cle ole 6) VEE at hs sm eie 
1. Pleurocystidia present and ObDViOUS...ccccccccccccvcccee4d 
4, Pleurocystidia unique, truly thin-walled (reminiscent 
of I. leptocystis), 55-72 x 15-22 pm, clavate, 
ventricose, subcapitate; spores 9-10 x 6.5 pm, oblong, 
with 10-14 prominent nodules; pileus 10-50 mm, dark 
umber-brown to "snuff-brown" on margin, with large, 
erect scales on disc, squarrose on margin; stipe 
pallid, darkening to brownish in age, pinkish buff at 
apex, concolorous with pileus below ececccccccece 
Cece rece cc cc ecc cee rsescceccccsecesvesecede JONGLCYEELS 
4. Pleurocystidia normal.cccercccsccvcccscccccssecccasesed 
5. Basidioma very small, pileus 5-15 mm, pale grey-brown, 
disc darker, greyish brown to chestnut-brown, with 
greyish white margin; spores 7-9 x 4.5-6 um, obscurely 
nodulose; pleurocystidia with bright yellow wall...... 
Coc c ccc rece c cc cc ev eccccccccccccccccccveel.e petiginosa 
5. Pileus 20-40 mm, vinaceous brown to tawny olive on 
Margin with cinnamon-buff disc, silky smooth at first 
from velipellis, becoming fibrillose, subrimose; 
stipe white to incarnate; spores unique, 5.5-8 x 13- 
19 pm, bullet shaped, with 1-5 large, obtuse nodules, 
or merely angular; pleurocystidia fusiform to clavate, 
thin-walled. cccccccccccccccccsvccevecele Chelanenesis 


Checklist of Species of Inocybe in California 


This is a checklist of nominal Inocybe species which either 
have appeared in the literature as occurring in California, 
or are associated with collections in the herbarium, SFSU. 
Some names, therefore, are synonyms or have been 
misapplied. Recent taxonomic studies of European inocybes 
by Kuyper (1986) and southern California inocybes by 
Nishida (1987, in prep.) should be consulted for 
discussions of species and complete synonymies. The columns 
to the right provide a quick ‘reference to reports of 
Inocybe species in California. Headings are: (R), reports 


in literature; (N), collection from northern California 
in herbarium SFSU, only those studied and annotated by D.E. 
Stuntz and/or Nishida; (S), collection of southern 
California in LAM determined by Nishida; (P), associated 
plants: "Q" = Quercus, "C" = conifer. In columns, "T" 
indicates type collection made in California, eh Seater 
present, "0" = absent. 

Smooth-spored Species R N S P 
I. adaegquata (Britz.) Sacc. 0) Xe 
I. agardhii (Lund) Orton 0 X 0 C 
as maalael ats Peck = I. whitei 0 Xo NO Fees 
i. amblyspora Kuhn. 0 0. eee 
I. anomala Murr. ae 0 0 Cc 
I. auricoma (Batsch) J. Lange 0. X (tie 


JR] 
e e 


JH 
e 


[HH 
e e 


PHIHI RII HII IRI 


Ta) La 


PH] Hy] a Hy] a] a] 4 
e e@ e @ ® e e e @ 6 e 


ea) Da) 
6 e e 


PHIR HHI eH 


es) Lt aed (un! (onc! (at (ue! 
e 


bakeri Peck 
brunnescens Earle 

non I. brunnescens Atk. 
bulbosa Peck 

(probably = I. bakeri) 
calamistrata (Fr.) Quel. 
cincinnatula Kuhn. 


= I. phaeocomis var. major 
cinnamomea A.H. Smith 


corydalina Quel. 


dulcamara (Alb. et Schw.: Fr.) Kumn. 


fastigiata (Fr.) Quel. = I. rimosa 
fastigiella Atk. 

flocculosa (Berk.) Sacc. 

fraudans (Britz.) Sacc. 
fuscodisca (Pk.) Massee 

geophylia — (Fr.:Fr.) Kummer 
geophylla var. lilacina (Pk.)Gill. 


geophylla f. perplexa K. Kauff. 
= I. whitei 


godeyi Gill. 
hirsuta var. maxima 
= I. calamistrata 
Aivteiia Bres. 
insinuata Kauff. 
jurana sensu auct. = I. adaequata 
kauffmanii Smith 
lacera (Fr.:Fr.) Kumn. 
laetior Stz. 
leiocephalia Stz. 


leptocystis Atk. 
jilacina (Pk.) Kauff. 


= I. geophylla var. lilacina 


muricellata Bres. 


olympiana Olympiana (A.H. Smith) 


phaeocomis var. major (S. Petersen) Kuyp. 


phaeodisca Kuhn. 


phaeoleuca Kuhn. 
pudica Kuhn. = I. whitei 


pusio P. Karst. 


pyriodora sensu auct. = I. fraudans 


gquietiodor M. Bon 


retipes Atk. 
rimosa (Bull.:Fr.) Kumn. 


serotina Peck 


(report is based upon a misapplied nam 


sororia Kauff. = I. rimosa 


splendens Heim 

subbrunnea Kuhn. = I. leiocephala 
subdestricta Kauff. 

subochracea (Pk.) Pk. 

vaccina Kthn. 

vinosistipitata Grnd & Stz. 
whitei (B. & Br.) Sacc. 


FHN #2487 
FHN #3976 


eae. 


ba 


ocooMxMOOCONOoOOy 


xO 


xMoOoooOoNK OH oS 


ooooooocooooco M~ooOonMnOoOCOoOCcS 


—! 


oO <~ oO 


a a a a ot a a oa 


< © 


CON K OM OKOK KM MO MK KOM KK MO 


COKOK KM NOM 


~ MM 


(aay piel Yeeys (ee) (any bee ee LE o (as) (any ed Fam) evel Ces yee feral rd ee) Com: Sic 


(=) 


ON OK MM OKN CO 


Seal Cea el Sel Ces) Cosy ped ee) bel Ce) 


ers 


moNQANM ANMNHMHONQ QAO AWDOVANODODWO NRO ANHMWMOOANOCHOHnOA O©Hn YP WY 


. 
QD 


) 


OF OF OF COR el Oe OF 2) 
QD 


194 


Q 


FHN #498 0 X xX 


ve) 
Zz 
~” 


Nodulose-spored Species 


[HH 
e 
a 
i=) 
=) 


acystidiosa Kauff. = 

= I. variabillima 
albodisca Peck 
bresadolae Mass. 
californica Kauff. 
chelanensis Stz. 
decipiens Bres. 
decipientoides Peck 

= I. variabillima 
fallax Peck 
lanuginosa (Fr.) Kummer 
leptophylla Atk. 
longicystis Atk. 


mixtilis (Britz.) Sacc. 
nigrescens Atk. 
oblectabilis Britz. 

olida Maire 

petiginosa (Fr.) Gill. 
umbratica Quel. 
variabillima Speg. 
xanthomelas Bours. & Kuhn. 


J] a] Hy] A] Ha 
2 © @ @ 8 @ 
ooowHo°o 
= = 
Q QD 


QD Q 


AWD AAW ANN APAAQ NANMAANRD AQ Ww 
Q 


Sa be bala Te ECU bt pen oe Dc Lacan D6 (bd DG) esa ns 
(ee eae Cam hey pony pope amy hile Pee, ob oo ore 


! Lama al Land Lond Ld Ln nd Can cD 
COoCCC COC COCCSO 


ACKNOWLEDGEMENTS 


This manuscript is based in part on a thesis submitted in 
partial fulfillment of the requirements for the Master of 
Science degree at California State University at Los 


Angeles. I thank the chairman, Daniel P. Mahoney and 
members of my committee, Harry D. Thiers and Brian Capon 
for their guidance and assistance. I thank Don R. 


Reynolds, Curator of Botany at the Natural History Museum, 
and Dr. Thiers for providing facilities and collections 
used in this research. Further, I wish to thank Darryl W. 
Grund, Harry Thiers and Don Reynolds for helpful comments 
and suggestions which improved this contribution. 


LITERATURE CITED 


Earle, F. S. 1904. Mycological studies II. Bull. New York 
Bot. Gard. 3: 289-312. 
Grund, D. W. & D. E. Stuntz. 1968. Nova Scotian Inocybes I. 
Mycologia 60: 406-425. 
& meme - 1970. Nova Scotian Inocybes II. Mycologia 
62: 925-939. 
-—---- & -----. 1975. Nova Scotian Inocybes III. Mycologia 


“= & -----. 1977. Nova Scotian Inocybes IV. Mycologia 
69: 392-408. 
--e— & -----. 1980. Nova Scotian Inocybes V. Mycologia 72: 


-—--- & -----. 1981. Nova Scotian Inocybes VI. Mycologia 


195 


733 655-674. 


et em & -----. 1983. Nova Scotian Inocybes VII. Mycologia 
Ba ce2oin 270. 
----- & -----. 1984. Nova Scotian Inocybes VIII. Mycologia 


763 733-740. 

Kauffman, C. H. 1920. Studies in the genus Inocybe. New 
York State Mus. Bul. (Rept. State Bot. 1919). 233/234: 
43-60. 

eae or om - 1924. Inocybe. North American Flora. 10(4): 227-260. 

a an Sn oe - 1925. The fungus flora of Mt. Hood, with some new 
species. Pap. Mich. Acad. Sci., Arts, and Letters 5: 
115-148. 

Kuyper, T. W. 1986. Revision of the genus Inocybe in 
Europe. I. Subgenus Inosperma of the smooth-spored 
species of subgenus Inocybe. Persoonia, Suppl. 
331-247. 

Murrill, W. A. 1916. Notes and brief articles. Mycologia 8: 
312. 

ois etoy - 1941. Some Florida novelties. Mycologia 33: 279-287. 

lad lag - 1944. New Florida fungi. Proc. Florida Acad. Sci. 
weeei—123. 

wep Sor - 1945. New Florida fungi. Quart. Journ. Florida Acad. 
Sci. 8:2 186-191. 

Nishida, F. H. 1987. The genus Inocybe in southern 
California (Agaricales, Basidiomycetes). M.S. Thesis. 
California State University, Los Angeles. 

ccm eee ° 1988. New species of Inocybe from _ southern 
California. Mycotaxon 33: 213-222. 

Peck, C. H. 1872. New York State Mus. Bul. (Rept. State 
Bot.). No. 23: 95-96. 

----- - 1874. New York State Mus. Bul. (Rept. State Bot.). 
NoOw1203 57. 

----— - 1875. New York State Mus. Bul. (Rept. State Bot.). 
Novw2.3) 95's 

----- - 1879. New York State Mus. Bul. (Rept. State Bot.). 
NOcwoe: 28-29. 

eaters - 1888. New York State Mus. Bul. (Rept. State Bot.). 
No. 41: 65-67. 

----- - 1898. New York State Mus. Bul. (Rept. State Bot.). 
NOs 22 289-290. 

-———= - 1904. New York State Mus. Bul. (Rept. State Bot.). 
No. 75: 17. 

a ne are - 1909. New species of Inocybe. Bull. Torr. Bot. Club 
662332-333. 

a - 1910. New York species of Inocybe. New York State 
Mus. Bul. (Rept. State Bot.). No. 139: 48-66. 

Smith, A. H. 1939. Certain species of Inocybe in the 
Herbarium of the Univ. of Michigan. Pap. Mich. Acad. 
Sci., Arts and Letters 24: 93-105. 

sete teat - 1941. New and unusual Agarics from North America. 
II. Mycologia 60: 406-425. 

a cs aie - & D. E. Stuntz. 1968. New or noteworthy fungi from 
Mt. Rainier National Park. Mycologia 42: 80-134. 
Stuntz, D. E. 1940. Inocybes of Western Washington. Ph. D. 

Dissertation. Yale University. 

=e aoghetys - 1954. Studies on the genus Inocybe II: new and 

noteworthy species from Michigan. Pap. Mich. Acad. 


196 


— = oe oe 


Sci., Arts and Letters 39: 53-84. 


- 1965. Inocybe (Fr.) Fr., subgenus Inocybium (Earle) 
Singer: section Inocybium Stuntz, section Rimosae 
(Fr.) Quel., section Dulcamarae Heim. University of 

Washington, Seattle (unpublished manuscript). 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 197-216 January 20, 1989 


FOLIICOLOUS FUNGI 8: CAPNODIUM IN CALIFORNIA* 


DON R. REYNOLDS 


Natural History Museum 
900 Exposition Boulevard 
Los Angeles, California 90007 


One species OL Capnodium occurs in 
California, which is redescribed with darkly 
pigmented and transverse to muriform septate 
ascospores. Capnodium dematum (V. M. Miller 
Seebonar) Reynolds nov. comb. ‘is culturally 
found, as predicted in the literature, to 
have Phaeoxyphiella fisheri Batista in 
Paweocal & /Ciferri, as its sanamorph.'\ Other 
California fungi identified, misdescribed, 
and misidentified as Capnodium are 
annotated. 


The systematic stability of the genus Capnodium 
sensu lato is representative of other ascomycete 
genera in which the sexual state has been 
historically presumed to be associated with one 
or more conidium-producing states. The first 
concept of the genus followed the spore 
morphology system of P. A. Saccardo and 
attributed darkly pigmented, muriform ascospores 
to the teleomorph and usually hyaline, 
unicellular or dark muriform pycnidial characters 
to deuteromycetes presumed as the anamorphs. 
Taxonomy of the California fungi called Capnodium 
is a microcosm of mycological progress from a 
Saccardian view of taxa delimitation based on 
character categorization to one of the "whole 
fungus" based on predictable patterns of 
biologically linked teleomorphs and anamorphs. 


* The previous paper in this series was published 
in Mycotaxon 27:377-403. 


198 


Attention was first drawn to California 
capnodiaceous fungi in the San Francisco Bay area 
by Cooke and Harkness (1881, 1884) in their 
series on California fungi. J. B. Ellis@aricees 
M. Everhart (1893-1899) included W. C. Blasdale, 
A. J. McClatchie and W. H. Harkness collections 
from the area in their exsiccati series, Fungi 
Columbiani. Miller and Bonar (1941) described new 
species and misidentified several collections 
from similar material from the same area. 

Batista and Ciferri (1963a, 1963b) incorporated 
these and other specimens in their monographic 
revision of the genus and its supposed anamorphs. 
Records of sooty molds have been made in other 
areas of California (Farlow, 1876; Millspaugh and 
Nuttatiive 1023e tRevnolasig 9387 ie 


CALIFORNIA CAPNODIUM 


1. Capnodium dematum (V. M. Miller & Bonar) 
Reynolds nov. comb. Miller and Bonar 
L941 3471. 
=Phaeosaccardinula dematum V. M. Miller & 
Bonar. Miller and Bonar 1941:411-412. 
=Capnodium baccharidis Batista & Ciferri. 
Batista and Ciferri 1963a:90-92. 
=Leptocapnodium krameri Batista & Ciferri. 
Batista and Ciferri, 1963a:123~12573giPme 
pantere Reynolds Lojane Sik Sen 


Type: USA, California, San Mateo County, Moss 
Beach, H. E. Parks’ 2139,’ 17 April 1924,on 
Bacchanis@pimluliard aye CGi oe 28. 


ASCOSPORES 5-6 celled, usually forming 
longitudinal septa in one or more cells at 
perpendicular to oblique angles to the cross 
septa; fusiform to elliptical in shape; the 
mature ascospore is usually larger at one end 
than the other because of by a smaller septal 
diameter at or near the median lengthoften giving 
the effect of a constricted spore; range in 
length is 25 to 40H; the forward cell of the two 
celled immature ascospore formed by the initial 
transverse septum enlarges more than the lower 
cell; pigmentation begins to be visible at this 
stage, as well as a distinct echinulation formed 
on the outer wall surface; the two-celled 


199 


ascospore measures 25, in length and 8 across 
the widest cell; ascospore size may remain 
constant or increase up to 40u in length and 12u 
across the wider cell as the number of septa 
increase; additional cross septa form in each of 
the two cells; the ascospore may remain 
transversely septate, or characteristic 
longitudinal septa will develop in one or more of 
the ascospore cells. ASCUS eight-spored, with 
thickened wall at apex and lateral areas; apical 
chamber with distinct nasse apicale; in ascospore 
dispersal the inner wall layer extends well 
beyond outer wall layer in the bitunicate 
fashion; fasciculate, maturing sequentially, 
obtuse, ventricose; measuring 10-25u at widest 
diameter, 65-100u in length. HYMENIUM basal, 
asci serially maturing; aparaphysate. ASCOCARP 
WALL multilayered, comprised of textura angularis 
tissue, the cells larger and lighter pigmented in 
the wider median area; outer layer externally 
aparaphysate, with deep brown melanoid wall 
pigment; inner wall layers producing elongate 
periphysoids projecting into centrum cavity that 
seem to be the source of a hygroscopic gelatinous 
matrix filling the ascus chamber. ASCOCARP 
100-150u , ostiolate, becoming collabent at 
times, occurring singly or amassed in groups of 
up to 15; subtended by a sterile tissue stalk of 
almost imperceptible length to one cm in length. 


ANAMORPH 


Phaeoxyphiella fisheri Batista in Batista & 
Ciferri. Batista and Ciferri 1963b:148-149. 

=Phaeoxyphiella morototoni Batista & Ciferri 
miuBatisctal svCiferris:: “Batista and’ Ciferri 
1963b:149-150. 

=Phaeoxyphiella walteri Batista, Nascimento & 
Giterrivin Batistavand Ciferri. " Batista and 
errerra 9690: 152-1535 

[=Hendersoniella sp. described by Fisher 
Neste LO LO Ore | 


CONIDIOSPORES formed in basal hymenial layer; 
linearly dispersed in dry catenate extrusion, 
with the acuminate straight to slightly curved 
and rounded pigmentless apex of one roughened 
conidium loosely adhering to the truncate base of 


200 


——s a 


— _ 
t —T J 


= nae = —— 
cr —= =f = 2 = 
ee eee - 


ee c 
\<EJE= — 
YSC 
Se 

pS 
x2 


Wie 
Ne 
LP 
[Vig ‘ 
Kat \ 
j U \" 
\\ a 


: 
A 


FIGURE 1. Diagramatic representations of the 
teleomorphic and anamorphic fruitbodies of 
Capnodium dematum (V. M. Miller & Bonar) 
Reynolds. A. Ascoma: Two asci extend among 
periphysoids from the apex of the broken asco- 
Carp wall. Ascospores are shown as two-celled 
initials, and with transeptate to muriform 
septation. B. Conidioma: The pycnidium is 
shown as a longtudinal section. The basal 
conidiogenous cells produce conidia which become 
loosely attached, end on end, as they move 
through and beyond the pycnidial neck. Bar=20uU . 


201 


the darkly pigmented conidium in line above it; 
length 40-75u; 4-8yu width at widest diameter; 
with 8 to 15 cross septa. CONIDIOGENOUS CELLS 
unicellular, hyaline, phialidic, solitary to 
multiconidiogenic, integrated into wall of 
conidiomata. PYCNIDIUM (conidium producing) 
with melanoid pigment in one or two outer layers 
of wall, comprised of textura globulosa tissue in 
the lower wall surrounding the conidiogenous 
centrum changing to textura prismatica tissue 
from the base of the neck onward to the ostiole; 
the innermost wall cells lining the conidiogenous 
centrum are hyaline; neck with central canal for 
spore dispersal 15-25u in width, extending 45-80U 
in length; lower area containing conidiogenous 
centrum 40-60,u in diameter and 65-85uU in height 
to base of neck. 


Pure culture isolates were made from single and 
multiple conidia of P. fisheri, and ascospores of 
Ceeeeuacum On half—strength YMA agar (2g 

yeast, 5g malt extract, 2g glucose [or 4g 
sucrose], 15g agar, 1000ml distilled water) and 
Nitrogen Base Medium (10ml of a concentrate made 
from 5g KCl, 5g MgSO4.7H20, 0.1lg FeSo4.7H20 added 
PomeienziPO4, Sg NaNO3, 20g sucrose,. 15g agar, 
1000ml distilled water). 


Germinating spores were observed in material 
collected from nature, as well as on the surface 
of the agar plates. On 2% plain agar, the hyphae 
from a germinating conidium were observed to grow 
downward through the agar with few hyphae 
developing near the agar surface. 


The colonies on YMA agar grew to a diameter of 
one cm in 14 days. The three-dimensional 
mycelium remained tightly formed into a pulvinate 
mass, with no perceptible individual hyphae 
extending into the agar at any measurable 
distance. The mycelial mass was transferred to 
nitrogen base medium without agar and maintained 
on a shake table at approximately 850 
revolutions/minute resulted in no discernable 
increase in mycelial mass nor change in colony 
morpholgy after two weeks. 


202 


Ascomata and conidiomata development was 
initiated through the cell division of a hyphal 
Cellmorerrom aycellyvotean 

ascospore or a conidium 


The type specimen of Phaeosaccardinula dematia, 
H. E. Parks 2139, was also cited as the type of 
Naetrocymbe scoriadea Batista & Ciferri, and 
mentioned as an additional specimen of 
Leptocapnodium kramerii Batista & Ciferri by 
Batista and Ciferri (1963a) (with the collector 
misprinted as H. E. Parker). 


Hughes (1976) predicted three Coelomycete taxa as 
the anamorphic component of Capnodium species, 
i.e. Fumagospora, Phaeoxyphiella and 
Scolecoxyphium. The proposed pleomorphy was 
based on concomitant occurrence in nature and the 
work of Fisher (1933, 1939) and Fraser (1934, 
1935a, 1935b). Representatives of these asexual 
taxa occur in association with naturally 
occurring C. dematum. The conidial state later 
described as Phaeoxyphiella fisheri was suggested 
as the anamorph of C. dematum (Miller and Bonar, 
1941), and has been culturally confirmed here. 
Fisher (1933), working with Australian species, 
failed to demonstrate pleomorphy between 
Phaeoxyphiella fisheri (as Hendersoniella sp.) 
and any ascosporic state; her spore-derived pure 
culture proof for the link between Capnodium 
salicinum Montagne and its anamorph in the same 
study is unquestionable. 


The handwritten label on P. A. Saccardo specimen 
2507, from Upper Yarra, Australia (LAM 202028), 
lists the names C. pelliculosum and C. ellisii 
Saccardo under C. walteri Saccardo. A 
coelomycete fitting the description of P. fisheri 
is present in the collection. I believe that the 
Saccardo species is based on heterogeneous 
material. The genus Capnodium is defined as 
having muriform ascospores (Reynolds, 1978). C. 
dematum produces ascospores with variable 
septation from transverse to muriform septation. 
Often the entire spore complement of an ascus 
will have only the transversely septate 
ascospores. 


203 


The other Capnodium species that has been 
suggested as having Phaeoxyphiella fisheri as its 
anamorph, C. walteri (Hughes, 1976), was described 
as having only transverse septate ascospores. 
Acceptance of this species into Capnodium as 
Hughes advocated (1976) raises a major question. 
A species with transverse to muriformseptate 
ascospores is acceptable into Capnodium as it now 
stands. To admit C. walteri as Fraser (1935b) 
described the species with only transversely 
septate ascospores requires an emendation of the 
genus. Or, the Phaeoxyphiella link must be 
accepted as a character uniting the variable 
spore septation pattern. Fraser (1935b) emended 
the Saccardo description of C. walteri without 
indicating that the type was reexamined; only 
material from the type locality was mentioned. 
None of the cited Fraser material that I examined 
had ascocarps of Capnodium; only Phaeoxyphiella 
fruitbodies were present. Obviously ascocarps 
were seen by her in some specimens that were the 
basis of the emendation. No designation was made 
in the list of examined specimens cited to 
indicate whether there was merely a presumption 
of pleomorphy and the presence of the supposed 
anamorphic state was taken as evidence of a 


Capnodium species. 


The proof of pleomorphy accepted by Fraser 
(1935a, 1935b) was indirect, based on cultural 
characters of the hyphae, such as measurement of 
the diameter of the colonies. (In some cases 
only comparative hyphal morphology was used as 
pleomorphic proof rather than the unquestionable 
demonstration via production of reproductive 
structures from single spore isolates.) 

Fraser's approach to the demonstration of 
pleomorphy is also evident in her description 
finaser, §1935a) lof ajvariety of C:, salicinum 
based on supposed differences in pycnidial 
characters, including the septation of the 
conidium. Fisher (1933) had earlier culturally 
demonstrated a varability in the number of 
conidial septa in the anamorph of Capnodium 
salicinum from an "abnormal" single septum to 
conidia with "increased septation in a muriform 
conidium." Based on study of the California and 
Australian material of C. dematum I consider the 


204 


pleomorphic link of Capnodium species with an 
anamorphic state other than Phaeoxyphiella to be 
tenuous until further data are available. 


CALIFORNIA FUNGI MISDESCRIBED AS CAPNODIUM 


1. Capnodium spongiosum Barr 


=Capnophaeum spongiosum (Barr) Batista & 
Ciferri. Batista and Ciferri 1963a:108 


=Morfea alaskensis (Saccardo & Scalia) 


OC AE ETAT OHO PT 


Batista & Ciferri. Batista and Ciferrigi96ca-1425) 


=Limacinia alaskensis Saccardo & Scalia 
Emended Barr. Barr 1955:501 


The concept of Capnodium used by Barr (1955) was 
"Species of sooty molds with muriform spores." 
The genus Capnodium sensu stricto of Hughes 
(1976) was characterized by sessile or shortly 
stalked ostiolate ascocarps without appendages; 
the ascospores were defined as transverseto 
muriform septate. Barr characterized the 
ascocarps of C. spongiosum as being appendaged 
and sessile, but occurring in or on a mycelial 
mat comprised of loosely tangled hyphae. Barr 
observed no longitudinal septa in the ascospores 
of the California material, although recent 
collections (LAM300844, LAM300845) demonstrate 
the variable septation of the ascospore in the 
California population of this species. The 
association of anamorphic states predicted for 
this species have yet to be validated (Barr, 
1955; Hughes, 1974, 1976). 


The Morfea alaskensis concept of Batista and 
Ciferri (1963a) was based on California material 
cited by Barr (1955) in her description of 
Capnodium spongiosum. The type material of 
Limacinia alaskensis, the supposed basionym of M. 
alaskensis, contains only immature ascocarps. 
Hughes (1974) provisionally included M. 
alaskensis as a synonym of Euantennaria 
rhododendri (Woronochin) Hughes. 


Our collections of Capnodium spongiosum 
(LAM300846) are mixed with deuteromycete species 


in Antennariella and Antennatula. The logic of 
the Hughes's (1976) hyphal morphology system that 
redefines the capnodiaceous sooty molds would 


205 


place Barr's California C. spongiosum in either 
one of two families. The pycnidium would 
indicate the ascomycete genus Antennulariella 
(allowing for muriform ascospores) of the 
Antennulariellaceae; the blastospore suggests the 
genus Euantennaria of the Euantennariaceae. 
Similar discrepancies can be noted (Hughes, 1968, 
tees O76) *for<another California sooty mold, 
Strigopodia batistae Hughes, which has conidia 
attributed to it from his Euantennariaceae 
(Antennatula) and from his Metacapnodiaceae 
[=Capnodiaceae sensu Reynolds, 1985] 


(Capnophialophora) 
2. Capnodium rhamni Cooke & Harkness. 


The original description of this species by Cook 
and Harkness (1884) makes this name an anamorphic 
one. The emended description by Batista and 
Ciferri (1963a) adds a teleomorph in such a way 
that the the holomorphn becomes a nomen 
confusum. 


Cooke and Harkness (1884) described the conidia 
as uniseptate, darkly pigmented, 18-20 x 9 and 
associated with a sexual state. No asci were 
described in their material (Harkness #2482). 


Hiebacista and Ciferri (1963a), the C. rhamni 
type was not declared as having been seen. 
Instead, California material (URM2380=IMUR5249 ex 
BPI) and another California collection (URM2807 
=EMUKooGS ex UC) collected by §H. E.- Parks from 
Marin county were examined instead. 


In their discussion of the URM2380 specimen under 
Capnodium rhamni Batista and Ciferri (1963a) 
mentioned that the material came to them from 
Herbarium BPI labeled as Capnodium rhamni on 
Rhamnus californicus; they found "only one 
species of Phaeoxyphiella and another one of 
Asbolisia." Batista and Ciferri's (1963b) 
description of Phaeoxyphiella fisheri 

Batista, Ciferri & Nascimento cites a Herbarium 
BPI collection with similar data as its type; no 
IMUR number was given. The specimen was 
collected "on leaves of Rhamnus, March 1931. 
Collected near Mirabel mine, Lake Co. Associated 


206 


with Asbolisia ampullula Speg. In the same 
material we were not able to find Capnodium 
rhamni Cooke and Hark. as determined by H. E. and 
S. T. ‘Parks... Miller-and,Bonar +(1941)sciteme 
specimen under their discussion of C. rhamni 
collected, "near Middletown, Lake Co., March, 
1931, H.. E. Parks 3581:'. Mirabell Minesre 
located four miles south of Middletown in Lake 
County, California (Yates and Hilpert, 1946). 


Clearly, the material collected by H. E. Parks 
from Lake County was used for two species 
descriptions. The redescription of Capnodium 
rhamni, was based on a portion which was kept in 
Herbarium URM as IMUR 5249 (Batista and Ciferri, 
1963a). The description of Phaeoxyphiella 
fisheri (Batista and Ciferri, 19963b) was based 
on the portion of the material returned to 
Herbarium BPI. 


My examination of URM2380 (= IMUR 5249) revealed 
P. fisheri, Asbolisia sp., and sporulating 
Hormonema ?prunorum (Dennis & Buhagiar) 
Hermanides-Nijhof. Specimen URM2807 was cited by 
Batista and Ciferri (1963a) as a gift from Lee 
Bonar. The data given were "on Rhamnus purshiana 


D.' Cc. », from Marin Co. California s\ucStAteae = 
Herb. Univ. California, Leg. H. E. Parksveepeoms 
19, 1931." The fungus present in the collection 


is Vertixore atronitidum V. M. Miller & Bonar. 


Batista and Ciferri (1963a) redescribed C. rhamni 
"based on this very poor material." The apparent 
reference is to the two specimens cited as 
IMUR5249 and IMUR566N. The asci were said to be 
commonly "early evanescent", but were described 
anyway. I believe that the Batista and Ciferri 
redescription of C. rhamni is based on characters 
from unrelated species present in the collections 
they examined. 


ty Capnodium heteromeles Cooke & Harkness. 


This species is a nomen confusum. The 
description of the spores was tentative (Cooke 
and Harkness, 1884), made from "free sporidia and 
hence uncertain." The morphology of the spores 
was proposed as triseptate, muriform, darkly 


207 


pigmented, 18 x 8u. The material in the type 
specimen, now located in Herbarium BPI rather 
than Herbarium CAS as stated in Miller and Bonar 
(1941), does not provide additional information 
about C. heteromeles. Batista and Ciferri (1963a) 
cited the binomial under "Excluded species of 
Capnodium" saying that "The spores are not surely 
recognizable." 


Two concepts are outstanding for the California 
material. The W. C. Blasdale collections, LAM 
200614, 200643, 200648, 202012 (= Ellis and 
Everhart, Fungi Columbiana 114 and North American 
Fungi 2918), made at the time that W. H. Harkness 
obtained his California material (LAM 200847), 
and a few years earlier than M. A. Howe collected 
material (LAM 200613) have the fruitbodies of the 
"Capnodium citri" pycnidium. Chaetasbolisia 
californiana Batista, Ciferri & Nascimento was 
described from Fungi Columbiana 114 (UC151179) 
(Batista and Ciferri, 1963b). The interpretation 
DvuevemMeeMiller and Ls Bonar} £xYfty years later, 
is supportive of the type description in 
septation, pigmentation and size of the supposed 
eonrarin, LAM 202010), 1202068) (Ex¥California® Fungi 
603, H. E. Parks collection 3621) Phaeochaetia 
arbutifoliae Batista, Ciferri & Nascimento was 
described from California Fungi 603 (Ex UC756704) 
(Batista and Ciferri, 1962). Reynolds (1982) 
considered P. arbutifoliae and the California 
species, Vertixore atronitidum Miller & Bonar to 
be conspecific. The description of the latter 
species (Miller and Bonar, 1941) cited the H. E. 
Parks collection 3621 and C. heteromeles was 
mentioned as associated with it. Recent 
collections from the Berkeley area (LAM 300843, 
300844, 300845) have conidiomata of these 
species. 


Other taxa have been described from the type 
material of C. heteromeles. Cooke and Harkness 
(1884) described Meliolopsis heteromeles Cooke & 
Harkness and Batista and Ciferri (1963a) found 
Naetrocymbe stevensonii Batista & Ciferri on the 
Harkness collection of Heteromeles leaves. 
Reynolds (1971b) transferred both M. heteromeles 
and N. stevensonii to Limacinula as synonyms of 
L. anomala (Cooke & Harkness) Reynolds. 


208 


MISIDENTIFIED CALIFORNIA CAPNODIUM 


1. Several collections labeled as Capnodium 
salicinum are mentioned in the literature. The 
California "Capnodium" ascospores in specimens 
associated with these reports that have been 
reexamined are too large for C. salicinum fide 
the Reynolds's (1978) restudy of the type. They 
are Capnodium dematum. 


Batista and Ciferri (1963a) did not state that 
they examined the Montagne type of C. salicinum 
for their monograph of the Capnodiaceae; rather 
they cited a material from Contra Costa County 
collected in 1931 by Lee Bonar. This collection 
is one of Capnodium dematum, UC966391, that A. C. 
Batista (unpublished data from herbarium label) 
also listed as having Capnodium salicinunm, 
Phaeoxyphiella morototoni, Fumagospora 
gaultheriae Batista & Vital, as well as Morfea 
hendrickxii (Hansford) Batista in the same 
collection. Collections from California were 
distributed as Capnodium salicinum Montagne in 
several exsiccati, including those of W. C. 
Blasdale in Ellis and Everhart's as Fungi 
Columbiani #112 and North America Fungi #2917, 
and that of H. E. Parks» as California»Fungieyooee 
Miller and Bonar (1941) cite a questioned C. 
salicinum determination on #112 as a specimen of 
Phaeosaccardinula dematum. Batista and Ciferri 
(1963a) cite the 2917 from P. A. Saccardo's 
herbarium as the type of Capnodium baccharidis. 


Miller and Bonar (1941) refer a California 
collection to Pleosphaeria salicina (Mont.) 
Arnaud (Arnaud, 1911). They confusingly 
described a setose, globose fruitbody anda 
glabrous Capnodium -like fruitbody "as described 
by Arnaud" from collections found on the bark of 


Baccharis pilularis DC. subspecies consanguinea 
(DC.) Wolf from Marin and Santa Clara Counties. 


Batista and Ciferri (1963b) cited a specimen of 
Fumagospora gaultheriae Batista, Vital & Ciferri 
associated with Capnodium salicinum from 
California on Baccharis pilularis, UC756704; 
this same collection (= LAM200744) serves as the 


209 


type specimen of Phaeochaetia arbutifoliae 
Batista, Ciferri & Nascimento (Batista and 
Geferra ,"/1962).. 


2. Several taxa have been described with darkly 
pigmented, muriform conidia. Most likely these 
fungi are related to the species cluster of 
Fumagospora capnodioides Arnaud, F. cistophila 
Batista & Ciferri and F. gaultheriae (Arnaud, 
Poe BaAtLSta, andy Ciferri,) .1963b),: 


a. Capnodium elongatum Berkeley & Desmazieres 


A California specimen named C. elongatun, 
collection by Marshall A. Howe, 6 June (error for 
January, according to L. Bonar [unpublished 
data]) 1894, was cited under Fumagospora 
gaultheriae in Batista and Ciferri (1963b). 
Miller and Bonar (1941) mention a W. C. Blasdale 
specimen of C. elongatum from Berkeley. 


| ely: Capnodium caespitosum Ellis & Everhart 


This species was described from A. J. McClatchie 
collection 748, July 1894, Pasadena California. 
The pycnidia were described as subulate- 
cylindrical, collected into hemispherical 
clusters, subtended by sparse moniliform hyphae. 
The conidia were described as muriforn, 
dark-brown, 12-20 x 7-10U. I have not seen the 
type specimen. 


c. Capnodium araucariae Thumen 


Thumen (1879) gave no description of the fruit- 
body and detailed the dark muriform spores | 
Waite ndtcating their,onigin.m j\Saccardo, \(1882) 
listed this species under "asci ignoti." 


This species was reported from Berkeley by Miller 
and Bonar (1941) as ?Polychaetella araucariae 
(Thumen) Spegazzini. They note the paucity of 
details in accounts of the species and the 
absence of authentic determined material for use 
in the determination. The larger spores they 
utilized were "seen out of the perithecia;" 
smaller spores were observed in the ascocarps. 


210 


3.) "Capnodiumici tri comprexs 


Several species in the sooty mold family 
Asbolisiaceae of Batista and Ciferri (1963b) have 
been referred to since the turn of the century in 
plant pathological literature as Capnodium citri 
(Farlow, 1876). This group of deuteromycetes, 
characterized by unicellular, hyaline conidia 
produced in pycnidia, are species in 


Leptoxyphium, Podoxyphium, and Polychaeton 
(unpublished data). 


a. Capnodium citri Berkeley & Desmaziéres. 


This name was used by Farlow (1876) fora 
California deuteromycete found on citrus and 
olive from San Diego and Santa Barbara. 

Batista and Ciferri (1963b) reinterpreted 
collections from Berkeley (W. C. Blasdale on 
citrus, March 1893) and Los Angeles (D. 
Coquillett on citrus, August 1889) as Podoxyphium 
yuccae Batista, Nascimento & Ciferri. 


b. Capnodium coffeae Patouillard 


Miller and Bonar (1941) uncertainly described 
under this name pycnidia under this name with 
hyaline conidia from Inverness Ridge in Marin 
County = 


c. Capnodium footii Berkeley & Desmaziéres 


Miller and Bonar (1941), citing a specimen 
(uncertain determination) from Berkeley, noted 
that they had only a pycnidiaceous sooty mold 
with hyaline, unicellular conidia. This is the 
same taxon that Batista and Ciferri cited from 
Moss (erroneously listed as Moon [L. Bonar, 
unpublished data]) Beach as Scolecoxyphium 
(Microxyphium) americanum Batista. I. Tavares 
(personal communication) noted a discrepancy 
between the conidial size reported for the 
species by Miller and Bonar (1941), 4-5 x 2-2.5H, 
and Batista and Ciferri (1963a) for the generic 
CESCrETD ELON ges are ar ee Leet Lt 


211 


"6 i Capnodium tuba Cooke & Harkness. 


Cooke and Harkness (1884) did not see asci in 
their specimen #2395 from Palo Alto. The 
ascocarps were described as cylindrical and the 
questioned conidia were characterized as oval, 
nonseptate, and hyaline. Batista and Ciferri 
(1963a) transferred the Cooke and Harkness 
species to Morfea without seeing the type. They 
based the concept of their new combination ona 
another Herbarium BPI collection by J. P. Thom. 
According to them, "It is possible that this 
material, collected in the same state and on the 
same genus of the host plant, would be referable 
to Capnodium tuba, agreeing with the description 
of Cooke and Harkness of Capnodium tuba for 
perithecia , since the ascospores were not 
described." 


Miller and Bonar (1941) cite specimens of C. tuba 
from Sausalito and Ross in Marin County, and from 
Wildcat Canyon in Contra Costa County consisting 
Onryeot pycnidia. 


SPECIMENS EXAMINED 


The specimen data are listed, if known, in the 
following order: country; state and county, or 
Gonparable “political division of a country; 
specific locality; collector and field number; 
date of collection; most original label 
determination (det.) and latest redetermination 
(redet.); associated plant; and curation history 
are given. The acronyms are those listed in the 
1981, Edition 7, of the Index Herbariorum.—_All 
specimens examined in the course of this study 
are listed, whether referable to the species 
discussed or not. 


Portions of a borrowed foliicolous fungal 
collection are many times retained by an 
investigator and added to an associated 
herbarium. This standard practice has, in my 
experience, resulted in multiple names for the 
same fungus, especially in the work of A. C. 
Batista and R. Ciferri. Knowing the collector of 
the specimen and the curation history as well are 


212 


often a valuable aid in the interpretation of a 
species, especially where the material is scanty 
or no longer in good condition. Therefore, the 
herbaria where the specimen at hand has been 
curated as part of a larger portion are listed as 
part of the specimen data when these data are_ 
known. 


AUSTRALIA. New South Wales: Ullaula, L. R. 
Fraser, April 1934, det. Capnodium walteri, on 
Casuarina glauca, LAM2009918 (Ex BPI, ex K), 
LAM200977 (Ex IMI26088). Victoria: Upper Yarra, 
Walter, det. Capnodium walteri and Capnodium 
pelliculosum and Capnodium ellisii, on Bursaria 
spinosa LAM200791 (Ex PAD2507, ex herbarium of P. 
Aw sSaACCardo) a WYONG ilies Sh. LE GaSe sb DOs 
January 1935, det. Capnodium walteri, LAM200750 
(EX DAR12717=DAOM110692). National Park: L. R. 
Fraser A93, June 1933, det. Capnodium walteri, on 
Dodonsea sp., LAM200752 (Ex DAR14686). FRANCE. 
Eure-et-Loir. Dreux, Gallet and Feulleaubus 571, 
September 1883, det. Capnodium footi and Fumago 
fagi, LAM201523 (Ex ILL, ex C. Roumeguere Fungi 
Selecti Gallici Exsiccati 3096). ITALY. Palermo: 
V. Beltrami, October 1883, det. Capnodium 
araucariae, LAM200662 (Ex ILL, ex Rabenhorst 
Fungi Selecti Gallici 3051). Parma: G. 
Passerini, 1875, det. Capnodium australe, redet. 
Fumagospora gaultheriae, on Alnus glutinosa, 
LAM201241 (Ex URM4806=IMUR10905, ex P. A. 
Saccardo herbarium, ex Thtimen Mycotheca 
Universalis 1350), LAM201242 (Ex URM4808 
=IMUR10907, ex Thumen Herbarium Mycologicum 
Oeconomicum 433). Verona: 1876, det. Fumagospora 
gaultheriae, on Alnus glutinosa, LAM201232 (Ex 
URM4518=IMUR10616, ex P. A. Saccardo herbarium). 
PORTUGAL. Coimbra: A. F. Moller; Auguste sae 
det. Capnodium araucariae, LAM201495 (Ex ILL, ex 
de Thtimen, Mycotheca Universalis 1737). A. F. 
Moller, det.Capnodium araucariae, LAM201529 (Ex 
ILL, ex C. Roumeguere Fungi Selecti Gallici 
Exsiccat 1e503i jae USA. California: Alameda 
County, Sunol, W.-H. Harkness 2425, AprilbMeee 
det. Meliopsis heteromeles and Capnodium 
heteromeles, on Heteromeles, LAM200847 (Ex BPI, 
ex CAS1961). Alameda County, Berkeley, M. A. 
Howe, 15 June 1892, det. Capnodium heteromeles, 
on Heteromeles arbutifolia, LAM200613 (Ex 


213 


ILL6642, ex California Fungi). Alameda County, 
Berkeley, W. C. Blasdale, March 1893, det. 
Capnodium heteromeles, LAM200643 (Ex ILL, ex 
Ellis and Everhart, North American Fungi 2918) 
LAM200648 (Ex ILL, ex Ellis and Everhart Fungi 
Columbiani), LAM 202012 (Ex FLASf1870). Alameda 
County, Berkeley, University of California 
campus, Robert Ornduff, 5 February 1988, det. 
Capnodium dematum, on Baccharus pilulans 
LAM300844. Alameda County, Berkeley, University 
of California campus, Don R. Reynolds, November 
1987, det. Capnodium dematum, on Baccharus 
pilulans LAM300843. Alameda County, Berkeley, 
University of California campus, Don R. Reynolds, 
30 March 1988, det. Capnodium dematum, on 
Baccharus pilulans LAM300845. Contra Costa 
County, Mt. Diablo, W. H. Harkness 2482, May 
1881?, det. Capnodium rhamni, LAM200874 (Ex 
URM2380=IMUR5249, ex BPI, ex CAS). Contra Costa 
County, north end of Wildcat Canyon, Lee Bonar, 
21 April 1931, det. Phaeosaccardinula dematia, on 
Baccharus pilulans, LAM201029 (Ex UC966391, ex 
California Fungi 796), LAM205515 (Ex ILL,). 
Tilden Regional Park, Ron Rossa, 1969, det. 
Phaeosaccardinula dematia, on Baccharus pilulans, 
LAM205443 (Ex IMI142282). Del Norte County, South 
Fork of the Smith River, Rock Creek Lodge, W. B. 
Cooke’vand V.. G. Cooke 25536,:.16 August 1949, det. 
Capnodium spongiosum, on Chamaecyparis 
lawsoniana, UC915700 (Ex Mycobiota of North 
America 400). Del Norte County, Smith River 
Lodge, H. E. Parks 5174, September 1934, det. 
Capnodium tuba "conidial stage", on Umbellaria 
californica, LAM201226 (Ex URM2806=IMUR5668, ex 
UC, ex Fungi of California). Del Norte County, 
SmrcoupRiver; HH. Eo) Parks5936,) May 1937, ‘det. 


Arthroboytrum spongiosum, redet. Capnodium 


spongiosum, redet. Capnophaeum spongiosum, on 
Libocedrus decurrens, LAM205509 (Ex UC692997), 


LAM205504 (Ex UC681382, ex California Fungiwi4 18% 
Del Norte County, Smith River, Lee Bonar, 13 July 
1933, det. Arthrobotryum spongiosum, redet. 
Limacinia multiseptata, on Libocedrus decurrens, 
LAM205507 (Ex UC960424). Marin County, H. E. 
Parks, 19 February 1931, det. Capnodium rhamni, 
LAM200875 (EX URM2807=IMUR5669, EX UC). Marin 
County, Inverness, H. E Parks, det. Capnodium 
salicinum plus Chaetasbolisia, redet. 


214 


Phaeosaccardinula dematia, on Baccharis 
pilularis, LAM201031 (Ex UC4394878, ex California 
Fungi 257). Marin County, Mt. Tamalpais #ih wee 
Parks 3065, det. Aithalomyces rhododendri, on 
Arctostaphlys sensitiva, LAM200583 (Ex ILL, ex 
UC21436).° Marin County, Mt. Tamalpais Fives 
Tradl, “Hover Parks) 218: Marchi 1926, dete 
Aithalomyces rhododendri, redet. Limacinia 
multiseptatum, redet. Ophiocapnocoma 
multiseptatum, on Manzanita, UC276037 (misprinted 
in) Batista and Ciferri, 1963aas;/UC2750g 7 

Marin County. Marin County, Mt. Tamalpais, H. E. 
Parks 3406, April 1929, Aithaloderma 
rhododendri, on Arctostaphylos sensitiva, 
LAM200888 (EX UC371934). Marin County, Mt. 
Tamalpais, Vera Mentzer, March 1933, det. 
Aithalomyces rhododendri, redet. Limacinia 
multiseptata, on Arctostaphylos sensitiva, 
LAM205503 (Ex UC498813, ex California Fungi 301). 
Marin County, Mt. Tamalpais, Mill Valley, 
southside, Cascade Trail, W. B. Cooke 16789, 23 
August 1942, det. Aithalomyces rhododendri, on 
Arctostaphylos sensitiva, LAM205508 (Ex 
UC681322). Marin County, east of Point Reyes 
lighthouse, Lee Bonar, 14 July 1939, det. 
Phaeosaccardinula dematia, on Baccharus pilulans, 
LAM201030 (Ex UC620826), LAM200620 (Ex#iListoegs 
San Mateo County, Moss Beach, H. E. Parks 2139, 
17 April 1924, det. Phaeosaccardinula dematia, on 
Baccharis pilularis, LAM200885 (EX UC617322), 
LAM200622 (Ex ILL31509), LAM200504 (Ex URM9128 
=IMUR13161, ex BPI). Sierra County, Sierra City, 
Don R. Reynolds, 9 April 1988, det. Capnodium 
spongiosum, on Libocedrus decurrens, LAM300846. 
Sonoma County, 2 miles east of Camp Meeker, M. A. 
Nobs and A. R. Kruckeberg, 28 September 1947, 
det. Arthrobotryum spongiosum, on Cupressus 
Sargentii, LAM200483 (Ex UC960406). Sonoma 
County, between Knights and Alexander Valley, H. 
FE. sParks3.622)9 20 May 1937 (det. Capnodium 
heteromeles , redet. Phaeosaccardinula anomala, 
on Photenia arbutifolia , LAM200550 (Ex 
URM12490=IMUR5276), LAM200615 (Ex ILL6644), 
LAM200744 (Ex UC75f6704, ex California fungi 
603), LAM201512 (Ex FLAS£41882). Washington: ~ H. 
E. Bailey,, March 1935, Mt. Ranier National Park, 
Adelaide Lake, det. Phaeosaccardinula dematia, on 
Abies amabilis, LAM200481 (Ex UC620741). 


215 


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Hist Publ. 212 7" Bot. "ser: 53 l—4935 (313 Sea 
fung1).VREYNOLDS)) (Dee R VN Loy lal NOLES 
capnodiaceous fungi II: Leptocapnodium. Bull. 
Torrey Bot. Club 98:151-154. REYNOLDS] (Deana 
1971b. The sooty mold ascomycete genus 
Limacinula. ‘Mycologia’ 63#1173-1209.." REYNOEDS au. 
R. 1978. Foliicolous ascomycetes 2: Capnodium 
salicinum Montagne emend. Mycotaxon 7:501-507. 
REYNOLDS, D. R. 1982. Foliicolous ascomycetes: 
4. The capnodiaceous genus Trichomerium 
Spegazzini emend. Mycotaxon 14:189-220 REYNOLDS, 
D. R. 1985. Foliicolous ascomycetes. 6. The 
capnodiaceous genus Limacinia. Mycotaxon 
2331531682 REYNOLDS, D.’ Ro) 1987." Provimomeees 
anamorphic connection. Chapter 8. pp 157-180. 
in J. Sugiyama (editor), Pleomorphic fungi: the 
diversity and its taxonomic implications. 
Kodansha;,'"*Ltd; Tokyo. \SACCARDO; ||P. "A. sie aan 
Sylloge fungorum omnium hucusque cognitorum. 
13:2-766"'(p.4 75) Padua. THUMEN,. “Fi Von eee 
Contributiones ad floram mycologicam lusitanicam. 
seri-II. Inst. Revista Sci. Litt. Coimbraweewee 
ser.)'s16-26. (YATES Ro 'G. ) and eS 0 Hae 
1946. Quicksilver deposits of eastern Mayacmas 
District, Lake and Napa Counties, California. 
Calif. J. Mines Geol. 42:231-286. 


ACKNOWLEDGEMENTS 


I thank Dr. Isabelle Tavares for the review of 
this paper and for making available the notes of 
Lee Bonar. Dr. Robert Ornduff kindly provided 
fresh study material from Alameda and Marin 
Counties. The field work was supported in part by 
the Los Angeles County Museum of Natural History 
Foundation. 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 217-220 January 20, 1989 


A NEW SPECIES OF GYMNOPILUS FROM NORTHERN CALIFORNIA 


Michelle T. Seidl/ 


Department of Biological Sciences, 
San Francisco State University, 
pan Francisco, California 94132 


This highly distinctive species of Gymnopilus, 
growing on fallen pine logs, was collected in Wunderlich 
Park in San Mateo County, California. The description 
which follows is taken from a thesis submitted to San 
Francisco State University in partial fulfillment of the 
Master of Arts degree in Biology (Seidl 1987). 


- The colors cited in the following description are from 
Kornerup and Wanscher (1978). 


Gymropilus thiersii Seidl, sp. nov. Fig.,1. 


Pileus (1-2-)3-15 cm latus, primo convexus, deinde 
planus, postremo subdepressus, siccus, fibrillosus vel 
Squamulosus discum versus, margine appendiculatus, initio 
omnino purpureo-brunneus, dein subflavidus et squamis sub- 
aurantio-brunneis vel rubello-brunneis tectus; vestigia 
veli partialis brunneolo-aurantia. Lamellae luteae vel 
subaurantio-brunneae, adnatae vel annexae, confertae. 
Stipes 3-10 cm longus x 0.7-3.2 cm crassus, aequalis vel 
attenuatus sursum, solidus, subfibrillosus, dilute aurant- 
iacus, supra annulum vulgo purpureus; apex dense pruinosus, 
pruina e vinaceo albescens. Annulus apicalis, submembran- 
aceus. Sporae 6-7.8 x 4.2-4.8(-5) um, ellipsoideae, 
asperae, in cumulo aurantiaco-brunneae. Cheilocystidia 
lecythiformia, hyalina. 


Basidiocarpia solitaria vel gregaria, ad lignum arborum 
coniferarum. Holotypus: Wunderlich Park, San Mateo 
Counvy, California, 13 March 1987, leg. M. TT. Seidl 2291 
(SFSU). 


Pileus (1.2-)3-15 cm broad, convex expanding to plane 
or slightly depressed on disc, irregular in age, purple 
brown to reddish brown (8E7), margin inrolled, appendicu- 
late with mandarin orange (65B8) to brownish orange (6C8) 
veil remnants, even, sometimes slightly hygrophanous, 


Ipresent address: University Herbarium, University 
of California, Berkeley, California 94720 


218 


Fig..1. Gymnopilus thiersii, holotype. 
Basidiomes ca. X .5. 


always fibrillose; surface dry, fibrillose, becoming scaly 
towards the disc, scales becoming erect, rimose-areolate in 
mature specimens, light orange (5A4, 5A5) to reddish brown 
(8E7), background pale yellow (3A3) or pastel yellow (3A4), 
developing vinaceous tints or, rarely becoming watersoaked 
where injured; context up to 2 cm thick on disc, white to 
pallid becoming purplish red (14A8) to deep Magenta (14E8) 
above lamellae and yellowish above stipe. Taste slight to 
very acrid. 


Lamellae adnate to adnexed, some with a decurrent 
tooth, crowded, edges becoming eroded in age, pale yellow 
(3A3), butter yellow (4A5), maize (4A6), sunflower (4A7), 
or deep yellow (4A8), becoming orange (6A8), dark orange or 
light brown (6D6) in age, bruising orange brown to brown 
where handled. 


Stipe 53-10 cm long, apex 0.7-5.2 cm thick, equal@or 
tapering upward, some flattened due to growth habit, at- 
tachment central to slightly eccentric, solid, surface 
streaked-fibrillose, pale orange (6A3) to greyish orange 
(6B3) background, purple (15C8) to dark purple (14F8) above 
annulus, sometimes with slight vinaceous tints below an- 
nulus, whitish tomentum at base, staining brownish orange 
(6C8) where handled, apex densely pruinose, pruina dark 
purple at first, becoming white when mature; annulus a- 
pical, submembranous, may appear densely fibrillose, usual- 
ly persistent; context white becoming yellowish in center 
and vinaceous at margins. 


2D 


Spores 6-7.8 x 4.2-4.8 um, ellipsoid, roughened, 
nondextrinoid, without germ pore, brown to rusty brown in 
Melzer's reagent; orange brown or reddish golden (6B8, 6C7, 
6C8, 6D7) in mass. Basidia 27-40(-43.5) x 6-10 un, 
clavate, hyaline, guttulate, (two)four-sterigmate. 
Pleurocystidia lacking. Cheilocystidia lecythiforn, 
hyaline, capitate or non-capitate, thin-walled, 34-38 x 
(3-4.5 at head) 8-143 um. Caulocystidia present. Lamellar 
trama of subparallel and slightly interwoven hyphae, rusty 
brown oleiferous hyphae apparent when mounted in 3% KOH. 
Hymenium orange brown, subhymenium light orange. 


Cuticle of light brown, repent, cylindric to slightly 
inflated, thin-walled hyphae, 7.5-15 um in diam, with en- 
crusting pigments and fascicles of hyphae; clavate en- 
crusted pileocystidia present; pileus context of inter- 
woven, cylindric to filamentous hyphae, slightly darker 
than cuticle with scattered orange brown oleiferous hyphae, 
3-5.5 um in diam. Clamp connections present. 


Habit, habitat and distribution. Solitary to gregari- 
ous. On fallen Pinus radiata D. Don. (Monterey pine), 
erumpent between scales of outer bark. January to April. 
California. 


Material examined. California: M. T. Seidl 2291 
MAOROUYPE: oFfoU), M. T. Seidl 2305 (ISOTYPE: SFSU), 
Wunderlich Park, San Mateo County, California. 


The burgundy, vinaceous or purplish tones present on 
the young unexpanded pileus and stipe make this species 
quite distinct. Singer (1986) refers to the following 
species with lilac or violet tones: G. janthinosarx 
(Sing.)Sing., G. calobasis, Pyrrhoglossum lilacipes 
Sing., and P. lilacinum Horak. Gymnopilus janthinosarx 
is from coniferous trees in Central Asia, has a lilac- 


violet context and a slightly viscid pileus (Singer 1949). 
Gymnopilus calobasis is an unpublished species from 
Amazonia, and no data are available concerning its char- 
acteristics. The two species of Pyrrhoglossum are char- 
acterized by an eccentric or lateral stipe, and are readily 
distinguished from G. thiersii. Gymnopilus luteofolius 
(Pk.) Singer has a similarly colored pileus, but the con- 
textual colors are different and the spores are dextrinoid 
(Hesler 1969). A related species, G. viridans Murr., has 
nondextrinoid spores and lacks pleurocystidia, plus many 
other differences such as an umbonate pileus and in the 
colors of the basidiome. Flammula purpurata Cke. et Mass. 
(now a Gymnopilus), described from Australia (Cleland 
1976), appears to be the most closely related species, 
however, the basidiome is much smaller, may possess green- 
ish tones on the pileus, lacks purple or vinaceous tones 
above the annulus, the spores are oblique and not el- 
lipsoid, and cystidia are lacking. 


220 


ACKNOWLEDGEMENTS 


The author would like to thank Dr. Roy E. Halling for 
technical assistance. I am grateful to Mr. David Christy 
for granting annual collection permits. For assistance. 
with the Latin diagnosis, I am indebted to Ellen Thiers. 


LITERATURE CITED 


Cleland, J. B. 1976. Toadstools and Mushrooms and other 
Larger Fungi of) South) Australia. Parts /I and@im 
1934-1935. A. B. James, Government Printer, South 
Australia.) 462). 

Hesler, L. R. 1969. North American Species of Gymnopilus. 
Hafner Publishing Company, New York. 117 p. 

Kornerup, A. and J. H. Wanscher. 1978. Methuen Handbook 
of Color. 3rd Ed. Eyre Methuen, London. 

Seidl, M. T. 1987. The Higher Fungi of Wunderlich Park, 
san Mateo County, California. San Francisco State 
University) SanlNrancitscos, 4 4551p ome) ne sien 

Singer, R.' 1949. ‘The Agaricales? Lilloa 22:562= 5008 

Singer, R. 1986. The Agaricales in Modern Taxonomy. 
4th Ed. Koeltz Scientific Books, Koenigstein, Germany. 


MYCOTAXON 


nas ee ees YIU eh YS NNO pi RoR IOL AD ie Werest (atl Hao 
Vol. XXXIV, No. 1, pp. 221-238 January 20, 1989 


THE LICHENS OF SERPENTINE ROCKS AND SOILS IN 
CALIFORNIA 


Lorene L. Sigal 
Environmental Sciences Division 
Oak Ridge National Laboratory 
Oak Ridge, IN 37831-6038 


SUMMARY 


Seventy-six species of lichens in 

33 genera were collected from serpentine 
rocks and soils at five locations in 
central California. This number was 
approximately double that of the genera 
and species reported for serpentine areas 
in Bosnia, Yugoslavia. None of the 
serpentine-endemic lichens reported for 
Bosnia were found in California; however, 
the prominent species on serpentine rocks 
were similar for both areas. Fourteen 
percent of the lichens found on California 
serpentine were cyanophilic, and 

11 percent were endolithic; 12 percent of 
the lichens collected on the high- 
magnesium serpentine were identified by 
various authorities as calcareous species. 


INTRODUCTION 


The unusual floras of serpentine soils have 
been of interest to botanists around the world for 
hundreds of years. The paucity of species, 
dwarfism, xeromorphism, and endemism of these floras 
have been attributed to various chemical and 
physical’ factors! (Brooks, 1987) including the 
following: 


222 


low levels of plant nutrient in the soils; 
excesses of magnesium over calcium; 


e high concentrations of phytotoxic chromiun, 
nickel and cobalt; and 

e the well-drained character of the 
shallow soils. 


It has also been suggested that the growth of 
endemic plants on serpentine soils is related to the 
freedom from competition provided by the substrate 
rather than to specific nutritional requirements. 
The general consensus at present is that no single 
factor is responsible for the serpentine vegetation 
but that a combination of edaphic and biotic factors 
results in the unusual serpentine floras (Brooks, 
Lo Saja 

The serpentine areas in the central coastal 
ranges of California have been studied by scientists 
of many disciplines (Whittaker, 1954, 1960; Barbour 
& Major, 1977; Kruckeberg, 1984a, 1984b, 1984c). 
Their reports contain examples of serpentine-endemic 
vascular plants, disjunct populations, and ecotypic 
variations; however, virtually no mention is made of 
the lichen or moss floras and their relationship to 
the serpentine substrate. The only field study to 
describe the lichen flora of serpentine substrates 
was performed by Ritter-Studnicka and Klement 
(1968) in the Republic of Bosnia in Yugoslavia. The 
purpose of this study was to increase our knowledge 
of lichen floras on serpentine substrates by 
collecting, identifying, and comparing the lichens 
from five serpentine sites in California; in 
addition, comparisons were made between the lichen 
floras on serpentine substrates in Bosnia and in 
California. 


METHODS 


Five areas of serpentine substrate were located 
on the geological maps of California with the 
assistance of staff at the California Division of 
Mines and Geology (Fig. 1). The study sites chosen: 


223 


(1) include serpentine exposures distributed over 
the central portion of the state, (2) represent 
typical serpentine conditions, (3) have published 
analyses of the soil and rock substrates, and 

(4) have vascular plant species characteristic of 
and/or endemic to serpentine. 

To ensure a complete sampling of the lichen 
flora, the author and a fellow botanist made at 
least two collection trips to each site (except for 
the Monkey Rock site which was visited only once by 
three botanists). The descriptive features of the 
study areas are given in Table 1; additional site 
information is found in Sigal (1975). 

Lichens were identified using the most suitable 
keys and descriptions available for California's 
saxicolous lichen flora, which has been largely 
neglected except by Hasse (1913) and Herre (1910), 
In places where these sources were incomplete, it 
was necessary to use the more general works of Fink 
(1935), Howard (1950), Nearing (1947), and Duncan 
(1959). Saxicolous species were also studied by 
Wetmore (1968) and Brodo (1968). Although these 
latter manuscripts describe floras of eastern 
sections of the United States, they were helpful in 
some determinations. 

Specimens in question were compared with known 
specimens in the herbaria at San Francisco State 
University and the University of California at 
Berkeley. Synonymy was a recurring problem, as were 
recent changes in nomenclature. The problem of 
comparing serpentine lichens in Bosnia with those in 
California was best expressed by Herre (1910): 
"Perhaps no more important work could be done than 
the careful overhauling of the synonymy of American 
lichens by someone who has access to the published 
exsiccata of Europe and America as well as the 
literature of the subject." 

Collections that were particularly difficult to 
identify and that were not represented by specimens 
in the local herbaria were mailed to specialists for 


224 


LITTLE RED 
MOUNTAIN 
MENDOCINO _e 
MONKEY ROCK 
COMPLEXION 
SPRINGS 


REDWOOD CITY Y 


SAN MATEO 


SAN LUIS OBISPO 


CUESTA RIDGE 


CALIFORNIA 


Fig. 1. Map of collection sites, 


225 


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226 


confirmation or identification. Dr. John W. Thomson 
assisted with Verrucaria, Physcia, Dermatocarpon, 
and Aspicilia. A specimen and chromatogram of 
Xanthoparmelia cumberlandia (Gyeln.) Hale was sent 
to Dr. Mason E. Hale, Jr., for confirmation wags 
single collection of an unknown "Bacidia" from 
Complexion Springs was sent to Dr. Brian J. Coppins. 
He identified it as Ramonia gyalectiformis (Zahlbr.) 
Vézda and mailed the collection to Dr. Vézda for his 
comments. Dr. Coppins stated that as far as he 
knew, the species was known only from its type 
locality at Palm Springs (Hasse 1913). 

Loans of material, particularly Lecidea, were 
made from Dr. Hannes Hertel, Dr. Chris Davidson, 
Dr. Howard Crum, Dr. John Thomas, and 
Dr. Isabelle Tavares. Species of the genus 
Aspicilia were a problem, and Dr. William Weber was 
consulted regarding this perplexing group. 
Collections were deposited at San Francisco State 
University. Nomenclature in this report follows 
thateot Evans Gl o7n 


RESULTS AND DISCUSSION 


Seventy-six species of lichens in 33 genera 
were collected from the five serpentine sites 
(Table 2). Compared to the study by Ritter- 
Studnicka and Klement (1968), which reported 
39 species of lichens in 16 genera on serpentine in 
Bosnia, this study indicates the richness of the 
lichen flora of California serpentine. 

For several reasons, the largest number of 
different lichens (42 species) was collected at 
Complexion Springs. This site had very large areas 
of weathered and unweathered serpentine as well as a 
constantly running stream that created yet another 
ecological niche. Although Little Red Mountain also 
had a constant water supply, fewer species (31) were 
collected at this site. The substrate was a fairly 


224 


homogeneous dunite type of serpentine, and this 
characteristic probably explains the more limited 
number of species. 

Both Redwood City and Cuesta Ridge lacked 
stream water, but the Redwood City site with 
24 species consistently had more in common with 
Complexion Springs than with Cuesta Ridge 
(24 species). Species at Redwood City were similar 
to those at Complexion Springs, probably because the 
blue-green type of serpentine was common to both of 
these sites, whereas Cuesta Ridge consisted largely 
of outcroppings of a weathered, brownish peridotite 
type of serpentine. 

Cuesta Ridge had 11 species that were found 
only at this southernmost site. Since neither 
climate nor substrate were substantial variables, 
one must consider other factors to explain the 
variation. Observers of the vascular plant flora 
indicate that the area of San Luis Obispo is unique 
for its number of endemic species. The flora of the 
Santa Lucia Mountains (Cuesta Ridge) contains many 
vascular plant species that have widely or markedly 
distinctive distributions due to isolation from 
connecting ranges. It appears that topography may 
influence also the lichen flora at Cuesta Ridge. Of 
the species found only at this site, Thelidium is 
considered a southern California lichen, and Peltula 
omphaliza is a distinctly southwestern species. 
Lecanora argopholis and three species of Caloplaca 
were unique to the southern collection site. In 
particular, Caloplaca bolanderi, with its 
outstanding iridescent red apothecia, was commonly 
present. The other species at Cuesta Ridge were 
also found at Complexion Springs and Little Red 
Mountain. 

There were 15 species of lichens collected at 
Monkey Rock, eight of which were not found 
elsewhere. These eight were lichen species expected 
at higher altitudes. 


228 


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231 


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232 


Of particular interest were the well-developed 
colonies of Rhizocarpon geographicum. Kruckeberg 
(1971) discusses R. geographicum in relation to 
serpentine: "Little is known yet about the edaphic 
preferences of bryophytes and lichens in the Pacific 
Northwest. One most striking case is the near 
absence on serpentine of the crustose lichen 
R. geographicum. It is plentiful on circumjacent 
nonserpentine rocks." In California I found 
R. geographicum on serpentine only at high-elevation 
sites. 

There was another yellow Rhizocarpon on the 
serpentine at mid-elevation sites that at first was 
mistaken for a depauperate R. geographicum but was 
identified as R. viridiatrum on closer inspection. 
It was listed only for the Complexion Springs site, 
but I have also found thriving colonies at other 
serpentine sites, particularly the Elwell Ranch 
property at Cedar Mountain, Alameda County, 
California, and at Goodyear’s Bar, Sierra County, 
California. Hasse (1913) cited a collection of R. 
viridiatrum on "trap" rock, and Herre (1910) noted a 
collection on sandstone, adding that R. viridiatrum 
was "a distinct species, very rare with us." It is 
possible that R. viridiatrum is a serpentine species 
since it is not commonly found elsewhere. Two 
collections were made of another lichen that, 
according to Herre (1910), is abundant on 
serpentine. Rinodina tephraspis was found at 
Complexion Springs and at Cuesta Ridge on the 
brownish, weathered peridotite. 

When the identifications were completed, some 
general floristic patterns were detected. Whether 
these patterns are related to the chemical or 
physical factors of serpentine substrates has not 
been determined because the source of nutrients and 
elements and the effects of substrate on saxicolous 
lichen species are largely unknown. Nevertheless, 
14 percent of the lichens found at the serpentine 


233 


sites were cyanolichens compared to an estimated 
8 percent of lichens worldwide (Hitch and Stewart, 
1973). Galun et al. (1982) suggest that 
cyanophilous lichens adapt well to extreme physical 
conditions; they may also be tolerant of extreme 
chemical conditions. Endolithic lichens represent 
11 percent of the species found on the various types 
of serpentine rock. Apothecia on the rock surfaces 
were the only clues to the presence of these 
lichens. At Monkey Rock, there were many well- 
developed apothecia of an unknown species of Lecidea 
on the slick, blue-green type of serpentine. 
Several species of Caloplaca were commonly 
endolithic, including: C. bolanderi, C. laeta, and 
C. "festiva" (identification uncertain). Lecanora 
polytropa was also present, recognizable by its 
light-brown, solitary apothecia scattered over the 
flat surfaces of the rocks as well as grouped in the 
crevices. There were several species of Verrucaria, 
their presence evident only by the partially innate 
perithecia, V. margacea and V. aethiobola in 
particular. Although these species were noted by 
Fink (1935) as having thalli that "sometimes 
disappear," the serpentine substrate appears to 
intensify the endolithic characteristic of the 
lichens for unknown reasons. A study by Wilson 
et al. (1981) on the weathering of serpentine by 
Lecanora atra suggested that lichens can sequester 
harmful ions in an amorphous silica gel that is 
formed from the oxalic acid secreted by the 
mycobiont and thus can tolerate high concentrations 
of phytotoxic elements. It is not known if the same 
strategy applies to endolithic lichen species on 
serpentine. | 

Twelve percent of the lichens collected on the 
high-magnesium serpentine were identified by various 
authorities as calcareous species. An outstanding 
example, the soil lichen Catapyrenium cinereum, was 
very abundant and obvious in the spring when mats of 


234 


the squamulose thallus were still green and turgid 
from the rains. During the summer, the lichen was 
harder to find, because the thallus dried brown 
and blended with the soil. According to 

Dr. John W. Thomson, C. cinereum is distinctly a 
calciphilous lichen, yet, predictably, it is found 
on the soil in serpentine areas. Ritter-Studnicka 
and Klement (1968) briefly mention "vague soil 
genera which also settle on chalk." They included 
in this group Dermatocarpon miniatum, Candelariella 
vitellina, Physica caesia (Hoffm.) Furnr., and 
Placodium saxicolum (Poll.) Kbr. [syn: Lecanora 
muralis (Schreber) Rabenh.]. The species of 
Dermatocarpon and Candelariella listed above were 
found at the California study sites but not 
necessarily on soil. Physcia caesia was found on 
serpentine but not at the study sites; L. muralis 
was not found. 

Hannes Hertel’s monograph on Lecidea (1967), in 
which he describes calciphilous species in the Alps, 
mentions two species of Lecidea that were commonly 
found on the California serpentine: Lecidea 
carpathica (syn: Lecidella carpathica) and Lecidea 
stigmatea (syn: Lecidella stigmatea). Lecidella 
carpathica was also found in Bosnia, but L. 
stigmatea was not mentioned by Ritter-Studnicka and 
Klement (1968). 

When the lichens collected at each site were 
compared, a group of species was found occupying 
the serpentine rock at most of the sites. (These 
lichens presumably had similar gross ecological 
requirements.) They comprise the "lichen 
association" of the selected serpentine sites in 
California. Included in the group are Aspicilia 
caesiocinerea, Lecidella carpathica, Rhizocarpon 
bolanderi, Caloplaca squamosa, Candelariella 
vitellina, and Acarospora fuscata. Ritter-Studnicka 
and Klement (1968) concluded that most of the 
lichens found on serpentine in Bosnia were members 
of the Aspicilictum ochraceae, a primary 
association. According to them, the lichens of this 
initial association occurred on slightly weathered 


235 


serpentine along a watercourse, above the high-water 
mark. In abundantly settled lichen colonies on 
serpentine, they found that Aspicilia caesiocinerea 
and Candelariella vitellina were the most 
conspicuous species of the association. The same 
species were also among the dominant members of the 
lichen association on the California serpentine--an 
interesting parallel to report. 

Ritter-Studnicka and Klement consider the 
following four lichens to be found only on 
serpentine: Rhizocarpon sphaericum (Schaer.) Mig., 
Aspicilia polychroma (Anzi) Nyl. var. ochraea Anzi, 
Aspicilia serpentinicola (Suza) Ras., and Aspicilia 
crusii Klem. Specimens of A. serpentinicola were 
borrowed from Dr. Hannes Hertel in Munich because 
this species was not found on the sites studied in 
California. At the time of this study, the other 
two species of Aspicilia were considered European 
because they were not listed in Hale’s checklist of 
North American lichens (1970); however, A. 
polychroma is listed in Egan’s 1987 checklist. 
Rhizocarpon spahaericum (Schaer.) Mig. is synonymous 
Witlionwe clLiriguratum (Anzi) Th. Fr., which is not 
considered to be serpentinicolous (W. Weber, 
personal communication). R. sphaericum has a bright 
yellow to whitish-yellow thallus and so differs 
markedly from the brown thallus of R. bolanderi, 
which is the Rhizocarpon found so commonly on the 
California serpentine. Because R. sphaericum has 
uniseptate spores and R. viridiatrum (also a yellow 
thallus) has multiseptate spores, these lichens are 
not likely to be confused. None of the European 
serpentine-endemic lichens were found at the study 
sites in California. 

According to Ritter-Studnicka and Klement 
(1968), the community changes as the serpentine 
weathers. They found that the Aspicilictum 
ochraceae association gave way to the more widely 
spread association of Aspicilictum cinereae, which 
is common throughout middle and southern Europe and 


236 


generally occurs on silicate rocks. The 
Aspicilictum cinereae association differs from the 
Aspicilictum ochraceae in the absence of Aspicilia 
polychroma var. ochracea and A. serpentinicola. The 
"silica-genera" found on the serpentine in Bosnia, 
which were considered part of the Aspicilictum 
cinereae association, were remarkably similar to 
those found in California. The species common to 
both continents included Lecidea carpatica KOorb.. 
(syn: Lecidella carpathica), Rhizocarpon 
geographicum, R. viridiatrum, and Aspicilia cinerea. 
In Europe the association of Aspicilictum cinereae 
is succeeded by that of Parmelietum molliusculae. 

I found no evidence on the sites studied of such a 
foliose association becoming the climax vegetation. 
The most common foliose lichen found at the study 
areas was Xanthoparmelia cumberlandia, generally 
considered a pioneer species. 

Discussion of the lichen flora of California 
serpentine is not complete until taxonomic 
uncertainties with the prominent Aspicilia cf. 
caesiocinerea Nyl. are described. The lichen is 
characterized by a variable rimose-areolate thallus 
with sunken apothecia and K-, C-, and I- thallus 
reactions; the exciple and hymenium of the apothecia 
have a wine-red I+ reaction. Multiple samples sent 
to two taxonomists were identified as A. 
caesiocinerea by one and A. cf. calcarea by the 
other. Comparison with the type specimen was not 
possible, but acceptance of the A. caesiocinerea 
determination further enhances the similarity 
between the lichen species found on serpentine in 
Bosnia and California. 


257 


ACKNOWLEDGEMENTS 


This field study was completed at San Francisco 
State University under the guidance of 
Dr. Harry D. Thiers, mentor and friend, without whom 
all that followed would not have been possible. 
Thanks, Harry. 

Oak Ridge National Laboratory is operated by 
Martin Marietta Energy Systems, Inc., under contract 
DE-ACO5-840R21400 with the U.S. Department of 
Energy. 


LITERATURE CITED 


Barbour, M. G. & J. Major. 1977. Terrestrial Vegetation of 
California. 1002 p. Wiley, New York. 

Brodo, I. M. 1968. The Lichens of Long Island, New York: a 
Vegetational and Floristic Analysis. New York State Museum and 
Science Service, Bulletin 410. 330 p. Albany, New York. 

Brooks, R. R. 1987. Serpentine and its Vegetation, a 
Multidisciplinary Approach. 300 p. Dioscorides Press, 
Portland, Oregon. 

Duncan, U. K. 1959. A Guide to the Study of Lichens. 164 p. 
Scholar’s Library, New York. 

Egan, R. S. 1987. A fifth checklist of the lichen-forming, 
lichenocolous and allied fungi of the continental United States 
and Canada. Bryologist 90:77-173. 

Fink, B. 1935. The Lichen Flora of the United States. 426 p. 
Reprint. University of Michigan Press, Ann Arbor, 1969. 

Galun, M., P. Bubrick & J. Garty. 1982. Structural and metabolic 
diversity of two desert-lichen populations. J. Hattori Bot. 
Lab. 53:321-324. 

Hale, M. E., Jr. & W. L. Culberson. 1970. A fourth checklist of the 
lichens of the continental United States and Canada. Bryologist 
73:499-543. 

Hasse, H. E. 1913. The lichen flora of southern California. 
ConurioonU.o..Nab., Herb, /) 1721-132, 

Herre, A. W. C. T. 1910. The lichen flora of the Santa Cruz 
Peninsula, California. Proc. Wash. Acad. Sci. 12:27-269. 

Hertel, H. 1967. Revision Einiger Calciphiler Formenkreise der 
Flechtengattung Lecidea. 155 p. J. Cramer, Germany. 

Hitch, C. J. B. & W. D. P. Stewart. 1973. Nitrogen fixation by 

. lichens in Scotland. New Phytol. 72:509-524. 

Howard, G. E. 1950. Lichens of the State of Washington. 191 p. 

University of Washington Press, Seattle. 


238 


Kruckeberg, A. R. 1971. Plant life of serpentine and other 

ultramafic rocks in northwestern North America. 
Syesis 2:15-114. 

1984a. California’s serpentine. Fremontia: 
LEGA) ke) 14 

1984b. The flora of California’s serpentine. 
Fremontia 11(5):3-10. 

1984c. California Serpentines: Flora, Vegetation, 
Geology, Soils and Management Problems. 180 p. University of 
California Press, Berkeley. 

Nearing, G. G. 1947. The Lichen Book. 648 p. Reprint. Ashton, 
Maryland, 1962. 

Ritter-Studnitka, H. & O. Klement. 1968. Uber flechtenarten und 
deren gesellschaften auf serpentine in Bosnien. Usterr. Bot. Z. 
115293-99,, 

Sigal, L. L. 1975. Lichens and Mosses of California Serpentine. 
M.A. thesis. San Francisco State University, California. 

U.S. Weather Bureau. 1963-1972. Climatological Data: 

California Summary. Vols. 67-76. Dept. of Commerce, 
Washington, DC. 

Wetmore, C. M. 1968. Lichens of the Black Hills of South Dakota and 
Wyoming. Publications of the Museum, Michigan State University. 
Biological Series 3(4):209-464. 

Whittaker, R. H. 1954. The ecology of serpentine soils. IV. The 
vegetation response to serpentine soils. Ecology 35:275-288. 

1960. Vegetation of the Siskiyou Mountains, Oregon and 
California: Ecol. Monogr. 30:270-338: 

Wilson, M. J., D. Jones & W. J. McHardy. 1981. The weathering of 

serpentinite by Lecanora atra. Lichenologist 13:167-176. 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 239-248 January 20, 1989 


LEPIOTA SENSU LATO IN CALIFORNIA. III. SPECIES WITH A 
HYMENIFORM PILEIPELLIS. 


WALTER J. SUNDBERG 


Department of Botany 
Southern Illinots University 
Carbondale, Illinois 62901 


SUMMARY 


_ Four species of Leptota, each with a hymeniform 
pileipellis, are noted from California. Three are 
described and illustrated--Leptota cristata (Bolt.: 
Fr.) Kummer, LZ. neophana Morgan, and L. thtersit. 
Leptota thtersit has ovoid spores and agrees with 
the original diagnosis of L. castanetdisca Murrill. 
Because Murrill's species is actually a synonym of 
L. cristata, L. thiersit is described as new. 


To my knowledge, the California mushroom flora has only 
four medium- to small-sized species of Leptota S. F. Gray 
sensu lato that exhibit a hymeniform pileipellis. In addi- 
tion to pileipellis anatomy and basidiome size, they share 
small spores (length = 8 ym or less) with walls inamyloid 
and weakly to not dextrinoid and an initially smooth, non- 
fibrillose pileipellis that in some species diffracts into 
more or less concentrically arranged scales. One species, 
Leptota Luteophylla Sundberg, was described previously 
(Sundberg 1971) and has since been reported from Michigan 
(Smith, Smith, and Weber 1979). The others are described 
below. With the exception of Lepiota cristata (Bolt.: Fr.) 
Kummer, these species are poorly known; one is described as 
new. All would be placed in section Cristatae Ktihner fol- 
lowing Singer (1975), but because of diversity in spore 
features, some workers (e.g., Bon 1981) align them in dif- 
ferent sections. 


In the descriptions below, colors in quotation marks are 
from Ridgway (1912). Herbarium abbreviations are according 


240 


to Holmgren and Keuken (1974). 


LEPIOTA CRISTATA (Bolt.: Fr.) Kummer. Der Fihrer in die 
Pilzkunde. p. 137. 1871. Figs. 1-2, 10-12. 
PILEUS 1-7 cm broad, almost conic at first, then convex 

to broadly convex, finally plano-convex in age; slightly to 

distinctly umbonate on expansion; margin at first incurved, 
appendiculate, and exceeding the lamellae, then decurved, 
plane to uplifted in age, entire, becoming rimose; surface 
dry; pileipellis continuous and glabrous at first, remain- 
ing so on the disc, becoming somewhat concentrically dif- 
fracted and appressed-scaly exposing the radially fibril- 
lose flesh toward the margin, infrequently splitting radi- 
ally, scales close to widely spaced, often evanescent near 
the margin; disc dark reddish brown ("carob brown", "choco- 
late", "chestnut", "bay", “walnut brown", or "burnt umber") 
to reddish brown ("cameo brown" to "warm sepia" to "verona 
brown"), sometimes as light as "russet" to pale "tawny" to 

"ochraceous-tawny", paler toward the margin, scales grading 

from concolorous near the disc to brownish pink ("fawn col- 

or" to "avellaneous") to "buff-pink" or sometimes "light 
vinaceous-cinnamon" to "light pinkish cinnamon" to "pinkish 
buff" toward the margin; exposed flesh white to "pale 
ochraceous-buff". 

FLESH 1-3 mm thick at the disc, solid, fragile, white 
and unchanging. Taste farinaceous (disagreeable), rarely 
indistinct. Odor sweet and cinnamon-like to farinaceous, 
rarely mild. 

LAMELLAE free, approximate; white, becoming "cartridge 
buff", "pale pinkish buff" to pale "warm buff"; close to 
subdistant; thin and fragile; somewhat ventricose; up to 
4.5 mm broad; margins smooth, concolorous. Lamellulae in 
one to three tiers. 

STIPE 3-12 cm long, 2-6 mm broad at the apex, equal to 
subbulbous; surface dry, almost glabrous at first, then in- 
nately fibrillose to appressed fibrillose, fibrils becoming 
loose in age; white to "pale pinkish buff" to "light pink- 
ish cinnamon", often tinged "vinaceous" near the base, be- 
coming "pale vinaceous" to "light russet vinaceous" to 
"snuff brown" to "sSayal brown", at times almost gray when 
handled or in age, base clothed with white, rhizomorphic 
hairs; stuffed, becoming hollow; pith fibrils white, cortex 
white at first, toward the base becoming "vinaceous-fawn" 
to "vinaceous-brown". 

ANNULUS occasionally persistent, submembranous, and 


241 


Figs. 1-2. Leptota cristata. 1. Sundberg 474. X 1. 


flaring, more frequently collapsed against the stipe or re- 
maining only as scattered patches of evanescent, tomentose 
to submembranous material, most often vanishing entirely; 
median to superior and white to "pale pinkish buff" when 
present. 

SPORES pale “cream"’ in mass, 6.3-7.9 X 3-4.7 jam; bullet- 
or wedge-shaped and spurred opposite the hilar appendix, 
tapered toward the rounded apex, inequilateral, smooth, 
wall slightly thickened, apical pore absent, most uniguttu- 
late, hyaline in KOH, inamyloid, not dextrinoid, pale to 
dark yellow in Melzer's reagent. BASIDIA 24-33 X 5-10 ym, 
clavate, 4-sterigmate, hyaline in KOH and Melzer's reagent. 
PLEUROCYSTIDIA lacking. CHEILOCYSTIDIA abundant, 22-45 X 
10-14 pum; clavate to sphaeropedunculate, bases often long, 
narrow, and with flexous walls, smooth, hyaline in KOH and 
Melzer's reagent. LAMELLAR TRAMA parallel, hyaline in KOH 
and Melzer's reagent. PILEAL TRAMA interwoven, hyaline in 
KOH and Melzer's reagent. PILEIPELLIS a hymeniform layer 
of elongate-clavate to pyriform elements, surface covered 
with a thin amorphous layer; elements 34-74 X 9-16 ym, some 
rostrate or branched, dull rusty brown in mass and hyaline 
to pale brown when viewed singly in KOH and Melzer's rea- 
gent. Subcuticular layer interwoven, hyphae hyaline to 
rusty brown in KOH. CLAMP CONNECTIONS present. 


242 


HABIT AND HABITAT: Solitary to scattered in humus under 
conifers, hardwoods, and mixed woods. November to March. 

COLLECTIONS EXAMINED: Alameda Co.: Thiers 7292 (SFSU). 
Butte Co.: Kowalski 960 (SFSU). Contra Costa Co.: Sundberg 
ITI-7-81-1 (SIU). Marin Co.: Sundberg 1732, 1777, 4068 (all 
SIU); Thiers 7326, 8517 (both SFSU). Mendocino Co.: Thiers 
8187, 8500 (both SFSU). Monterey Co.: Thiers 11811 (SFSU). 
San Francisco Co.: Sundberg 1004; Thiers 8581 (both SFSU). 
San Mateo Co.: Sundberg 474, 1116 (both SFSU). Santa Bar- 
bara Co.: Sundberg I-7-83-5 (SIU). Santa Cruz Co.: Sund- 
berg 1017 (SFSU). Solano Co.: Sundberg 1557 (SIU). Sonoma 
Co.: Sundberg XI-8-86-B-3 (SIU). Trinity Co.: Sundberg 
1144 (SIU). 

DISCUSSION: Leptota crtstata is widely distributed in 
California. It differs from other species in section 
Cristatae included here by formation of spurred spores-- 
spores that are bullet- or wedge-shaped (more or less trun- 
cate) and spurred at the base opposite the hilar appendix. 
Because of this feature, some workers (e.g., Bon 1981) put 
L. crtstata in section Stenosporae (Lange) Kuhner. 


LEPIOTA THIERSII Sundberg, sp. nov. Figs. 3, 5-64 


Pileus 0.8-4 cm latus, convexus demum plano-convexus, 
Siccus, glaber, ad marginum demum squamosus, cinnamomeo- 
brunneus, ad marginum pallide cinnamomeus vel subroso- 
bubalinus. Caro albus. Lamellae liberae, confertae, fere 
albus. Stipes 3-6 cm longus, 2.5-5 mm latus, glaber vel 
obscure fibrillosus, albus, basi cinnamonmeus vel pallide 
brunneus. Annulus evanescens. Sporae 4.7-6.3 X 3-3.9 ym, 
Ovoideae vel breve ellipsoideae, non dextrinoideae vel 
amyloideae. Cheilocystidia nulla.  Cuticula paliformis. 
Typus: United States, California, San Mateo County, San 
Francisco Water Department Property, February 10, 1970, 
Sundberg II-19-87-A-2 (Holotypus: SIU; Isotypus: SFSU). 


PILEUS 0.8-4 cm broad, convex when young, then broadly 
convex, finally plane, often with a slight to broad, convex 
to flat umbo; margin incurved, appendiculate, and exceeding 
the lamellae at first, then decurved and entire, becoming 
plane, eroded to lacerate in age, appendiculate material 
thin, cottony to submembranous, fugacious; surface dry; 
pileipellis continuous and glabrous throughout at first, 
sometimes remaining so, usually becoming more or less con- 
centrically diffracted and appressed-scaly exposing the 
radially fibrillose flesh toward the margin; scales elon- 


243 


Fig. 3. Leptota thtersit. Sundberg II-18-87-A-2. X 2/3. 


gated along the pileus circumference near the disc, irregu- 
lar in outline, smaller and more equal-sided toward the 
margin; flesh fibrillose and radially arranged between the 
scales; disc brown ("verona brown") to "cinnamon brown" to 
“clay color" to "cinnamon" to "pinkish cinnamon", rarely 
"warm buff" to "light buff", unbroken surface and scales 
grading from concolorous to "pale pinkish buff" near the 
margin, exposed flesh white to "pale pinkish buff". 

FLESH 1-5 mm thick at the disc (at stipe edge), solid, 
white and unchanging. Taste disagreeable, farinaceous, 
rarely mild. Odor sweet-farinaceous to farinaceous, rarely 
indistinct. 

LAMELLAE free, close to the stipe, infrequently forking 
and anastomosing near the stipe; white to off-white (pale 
"cream buff" to "pale pinkish buff"); close; thin and frag- 
ile; up to 5 mm broad, margin even, smooth, concolorous. 
Lamellulae in one to three tiers, free ends abruptly taper- 
ed but not truncate. 

STIPE 3-6 cm long, 2.5-5 mm broad at the apex, equal; 
surface dry, innately fibrillose, white at first, becoming 
tinged "cinnamon-buff" to "cinnamon" to dark cinnamon-brown 
("Ssayal brown" to "mikado brown") to pale brown toward the 
base in age or on handling, covered at the base by a white 
tomentum; stuffed, becoming hollow, pith fibrils white, 
cortex concolorous with the surface. 

ANNULUS when present superior, submembranous, appressed 
to slightly flaring, very fragile, off-white, usually eva- 
nescent on most specimens, often leaving only a fibrillose 
zone or adhering to the pileus margin. 


244 


SPORES white (prints thin), 4.7-6.3 X 3.1-3.9 yam; ovoid 
to short ellipsoid, slightly inequilateral, smooth, apicu- 
late, thin-walled, apical pore absent, uni- to biguttulate, 
hyaline in KOH, inamyloid, not dextrinoid, pale yellow in 
Melzer's reagent. BASIDIA 21-29 X 5-8 ym, clavate, 4-ster- 
igmate, hyaline in KOH, hyaline to pale pinkish in Melzer's 
reagent. PLEUROCYSTIDIA lacking. LAMELLAR TRAMA subparal- 
lel to parallel, hyaline in KOH, hyaline to pale pinkish in 
Melzer's reagent; subhymenium cellular, concolorous. PILE- 
AL TRAMA interwoven, hyaline in KOH, hyaline to pale pink 
(more reddish in mass) in Melzer's reagent. PILEIPELLIS an 
irregular and tightly arranged palisade of more or less 
clavate elements (often appearing somewhat cellular in 
thicker sections), surface covered with an amorphous mat- 
erial; elements 27-49(-90) X 9-20 pm, clavate, sometimes 
irregularly shaped, rarely branched, bases at times long 
and narrow, walls unevenly thickened, hyaline to pale yel- 
lowish brown in mass, and hyaline or with a yellowish brown 
content when viewed singly in KOH, more intense yellowish 
brown in Melzer's reagent; some subcuticular elements en- 
crusted. CLAMP CONNECTIONS present at the base of the 
pileipellis elements and in the trama. 

A pale yellow pigment, soluble in water but not alcohol, 
diffuses from the tissue during rehydration for sectioning. 

HABIT AND HABITAT: Scattered to gregarious in humus un- 
der Cupressus macrocarpa.s November through April. 

COLLECTIONS EXAMINED: San Mateo Co.: Sundberg II-18-87- 
A-2 (Holotype, SIU; Isotype, SFSU); Sundberg 1125 (SFSU), 
1842, 4153, 4192, II-20-87-6, II-20-87-8, I1I-20-87-9, 
II-20-87-11 (all SIU); Thiers 11934, 18832 23195) 520 7ie5 
248113, 330700 (al NSESU)S 

DISCUSSION: The specific epithet honors, with deepest 
gratitude, Dr. Harry D. Thiers who first collected the 
species and who introduced me to mycology. 

This species agrees with the original description of 
Leptota castaneidisea Murrill (1912). However, Murrill 
(1914) subsequently listed L. castanetdisca as synonomous 
with L. cristata. Type studies of L. castaneidtsca by 
He. Smith (1966) and myself both confirm Murrill's conclu- 
Sion. Therefore, this species is described here as new. 

Leptota thtersit is morphologically similiar to and usu- 
ally exhibits the odor of LZ. cristata, but differs from it 
by usually lacking the reddish component in the pileus disc 
color and always having ovate rather than spurred spores. 
The formation of scales by the pileipellis on pileus expan- 
sion and a lighter colored stipe as well as the lack of 
both orange in the pileus color and grayish pileal trama 


245 


Fig. 4. Leptota neophana. Sundberg II-20-87-A-1. X 1. 


distinguish L. thiersii from L. cristatoides Einhellinger. 
In Bon's (1981) system, L. thtersti would belong in 
section [tlaceae Bon. 


LEPIOTA NEOPHANA Morgan. J. Mycol. 12: 248. 1906. 
Figs. 4, 7-9. 


PILEUS 1.8-3.7 cm broad, broadly convex to plane, often 
with a low broad umbo; margin appendiculate, exceeding the 
lamellae and incurved at first, becoming decurved to plane, 
entire to rimose, frequently undulating; surface moist to 
dry; pileipellis continuous, glabrous, and sometimes mi- 
nutely rugulose on the disc, becoming more or less concen- 
trically diffracted and appressed-scaly exposing the flesh 
toward the margin; scales irregular in outline, elongated 
along the pileus circumference near the disc and more uni- 
formly equal-sided toward the margin; disc blackish brown 
to very dark brown ("fuscous" to "mummy brown" to "mars 
brown"), scales concolorous but most lacking black tinges 
to somewhat paler ("clove brown"), exposed flesh white at 
first, becoming very pale grayish olive ("olive-buff" to 
"pale olive-buff"). 

FLESH 1-3 mm thick at the disc (at stipe edge), solid, 
fragile, white, tinged "pale pinkish buff" under the disc, 
unchanging. Taste mild. Odor slightly pungent. 

LAMELLAE free, approximate at first, but receding early, 
separated from the stipe by a collar-like band; white near 
the stipe, off-white ("cartridge buff") to "light buff" to 
"warm buff" elsewhere, unchanging; close to subdistant; 
thin and fragile; margins even, minutely fimbriate when 


246 


Figs. 5-12. Anatomical features of Leptota thtersit, Le 
neophana, and L. cristata. 5-6. JL. thtersii. 5. Basidio- 
spores. 6. Elements of pileipellis. 7-9. JL. neophana. 
7. Basidiospores. 8. Cheilocystidia. 9. Elements of 
pileipellis. 10-12. L. ertstata. 10. Cheilocystidias 
11. Basidiospore. 12. Elements of the pileipellis. 

Scale lines = 10 pm. 


viewed with a hand lens, concolorous. Lamellulae in one to 
two tiers; free ends abruptly tapered but not truncate, 
sometimes anastomosing with longer lamellae. 

STIPE 2.5-7.5 cm long, 2.5-6 mm broad at apex, equal; 
surface dry; glabrous to silky appressed fibrillose above, 


247 


fibrillose to obscurely fibrillose scaly toward the base; 
white, becoming "pale pinkish buff" to "pale pinkish cinna- 
mon", sometimes developing "cinnamon" to "“orange-cinnamon" 
or brownish pink ("avellaneous", "vinaceous-fawn", or "fawn 
color") streaks and areas toward the base in age, "vina- 
ceous" when bruised; clothed at the base with numerous, 
white to "pale cinnamon-pink to "light vinaceous-cinnamon" 
mycelial hairs that sheath it where buried in humus; stuff- 
ed becoming hollow, pith fibrils white, cortex white, "pale 
cinnamon-pink" to "light vinaceous-cinnamon" near the base. 

ANNULUS lacking or if present forming only a fibrillose 
to appressed zone near the stipe apex, concolorous with the 
pileus surface. 

SPORES "pale cream" in mass, 4.7-6.3 X 2.7-3.2 jam, ovoid 
to short ellipsoid, inequilateral, smooth, apiculate, thin- 
walled, apical pore lacking, most uniguttulate, hyaline in 
KOH, inamyloid, not dextrinoid, pale yellow in Melzer's re- 
agent. BASIDIA 21-29(-34) X 5-7 um, clavate, 4-sterigmate, 
hyaline in KOH and Melzer's reagent. CHEILOCYSTIDIA 19-48 
X 6-11(-14) ym, abundant; clavate to elongate-clavate, some 
ventricose, infrequently rostrate, base narrow and elongate 
in most; walls thin, often flexuous; hyaline in KOH and 
Melzer's reagent. PLEUROCYSTIDIA absent. LAMELLAR TRAMA 
parallel to subparallel, hyaline in KOH, hyaline to pale 
pink in Melzer's reagent; subhymenium obscure, elements 
small, concolorous with trama.e PILEAL TRAMA interwoven, 
hyaline in KOH, hyaline to pale pink in Melzer's reagent; 
oleiferous hyphae infrequently present. PILEIPELLIS a pal- 
isade of clavate to pyriform, rarely catenulate elements, 
surface covered with a hyaline amorphous material; elements 
25-75(-87) X (11-)14-25 pm, rarely rostrate, some distorted 
and with flexuous walls, bases often narrow and elongate, 
walls thickened, pale yellow to pale yellowish brown in KOH 
and yellowish brown to dark reddish brown in Melzer's rea- 
gent, pigment in the walls. CLAMP CONNECTIONS present in 
the subcuticular layer, pileal trama, and lamellar trama. 

HABIT AND HABITAT: Solitary to scattered in humus under 
Cupressus macrocarpa. December through April. 

COLLECTIONS EXAMINED: Butte Co.: Kowalski 1281 (SFSU). 
San Mateo Co.: Sundberg 1093, 1126 (both SFSU), 1835, II- 
18-87-A1, II-20-87-1 (all SIU). 

DISCUSSION: Known also from Ohio (Morgan 1906) and 
Michigan (Smith 1954), L. neophana is apparently widely 
distributed but is very infrequently reported. The dark 
color of the pileus readily distinguishes this species from 
others noted in this paper. 

In Bon's (1981) treatment, L. neophana would belong in 


248 


section [ilaceae. 


ACKNOWLEDGMENTS 


I wish to thank Dr. Harry D. Thiers of San Francisco~ 
State University for his friendship and guidance, for al- 
lowing use of specimens and photographs in his care, and 
for making my personal field observations possible through 
his collecting permit granted by the San Francisco Water 
Department. I also thank Dr. Barbara Thiers of the New 
York Botanical Garden for providing some needed literature 
and Dr. Mary Palm of the National Fungus Collections for 
her critical and valuable comments. The help of Dr. 
Charles Speck of Southern Illinois University in preparing 
the Latin diagnosis is appreciated. Funding provided for 
some of my field work by the American Philosophical Society 
Grant No. 1388--Johnson Fund is gratefully acknowledged. 


LITERATURE CITED 


Bon, Me 1981. Cle Monographique des "Lepiotes" d'Europe 
(= Agartcaceae, Tribus Leptoteae et Leucocoprineae). 

Docs Mycol. 1is te/7. 

Holmgren, P. Ke, and W. Keuken. 1974. Index Herbariorum, 
Part I. Regnum Veg. 92: 1-397. 

Morgan, A- P. 1906. North American species of Lepiota. 

Je Mycol. 12: 242-248. 

Murrill, W. Ae. 1912. The Agaricaceae of the Pacific Coast 
--II. Mycologia 4: 231-262. 

Murrill, We. Ae 1914. Agaricaceae (Pars). North American 
Flora 10: 1-65. 

Ridgway, Re 1912. Color standards and color nomenclature. 
Washington, D. Ce. 43 pp., 53 plates. 

Singer, Re 1975. The Agaricales in modern taxonomy. J. 
Cramer, Vaduz. 912 pp., 84 plates. 

Smith: A. H., He V- ‘Smith, and N. S. Weber.) 1979.0 show 
know the gilled mushrooms. Wm. C. Brown Company, Pub- 
lishers, Dubuque. 334 pp. 

Smith, H. Ve. 1954. A revision of the Michigan species of 
Leptotae lLloydia 17: 307-328. 

Smith, H. Ve. 1966. Contributions toward a monograph on 
the genus Lepiota, I. Type studies in the genus Lepiota. 
Mycopath. Mycol. Appl. 29: 97-117. 

Sundberg, W. J.« 1971. A new species of Lepiota. Mycol- 
ogia 63: 79-82. 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 249-252 January 20, 1989 


EXAMINATION OF STILBOTHAMNIUM TOGOENSE FOR ASPERGILLUS 
FLAVUS GROUP MYCOTOXINS. 


D.T. WICKLOW, R.F. VESONDER & CESARIA E. MCALPIN 


U.S. Department of Agriculture 
Agricultural Research Service 
Northern Regional Research Center 
1815 North University Street 
Peoria, IL 61604 


R.J. COLE 


National Peanut Research Laboratory 
ARS, USDA 
Dawson, GA 31742 


MARIE-FRANCE ROQUEBERT 


Laboratorie De Cryptogamie 
Museum National D'Histoire 
12, Rue Buffon 
75005 Paris, France 


SUMMARY 


Stilbothamnium togoense (=Aspergillus togoensis), a taxon 
considered by some to be related to the Aspergillus flavus 


group near A. tamarii, produced sclerotia when cultured on 
autoclaved oatmeal, but no A. flavus group mycotoxins (e.g. 
aflatoxins, sterigmatocystin, cyclopiazonic acid, kojic 
acid, dihydroxyaflavinine and aflatrem) were detected in 
extracts of the sclerotia or fermented meal. When cultured 
on autoclaved rice, S. togoense produced substantial 
quantities of sterigmatocystin which is believed to be an 
intermediate in the biosynthesis of aflatoxin. The data 
Supports linking A. togoense to the A. flavus group because 
sterigmatocystin is not produced by species in the A. 
ochraceus group with which A. togoense also shares 
outstanding characteristics. 


Stilbothamnium cogoense Henn. [=Aspergillus togoensis 


(Henn.) Samson and Seifert] occurs in equatorial 
rainforests where it colonizes the seeds of fruit rotting 
on the forest floor. The fungus produces large synnemata 
(6-8 cm tall) with yellow to greenish or brownish, radiate 
conidial heads and is considered by Roquebert and Nicot 
(1985) to be related to the A. flavus group sensu Raper 


250 


and Fennell (1965) near Aspergillus tamarii Kita. 

Roquebert and Nicot noted that the formation of a septum in 
the lower part of the phialide of S. togoense makes it 
different not only from A. tamari but from all other 
aspergilli as well. Samson and Seifert (1985) reduced 
Stilbothamnium to synonymy with Aspergillus retaining the 
species of Stilbothamnium in a separate subgenus. [i.e. — 
Aspergillus subgenus Stilbothamnium (Henn.) Samson and 
Seifert]. They reasoned that there are numerous 
synnematous species in Penicillium, thus the production of 
such structures in connection with the Aspergillus anamorph 
should not serve to exclude those taxa from Aspergillus. 
The suggestion that A. togoense may be an ancestral form of 
the A. flavus group (Samson and Seifert, 1985) prompted us 
to investigate whether S. togoense produces any of the 
known mycotoxins (e.g. aflatoxin, aflatrem, 
dihydroxyaflavinine, kojic acid, cyclopiazonic acid, 
sterigmatocystin, and other aflatoxin-like metabolites) 
associated with different members of the A. flavus group 
[e.g. A. flavus var. flavus Link: Fr., A. flavus var. 
parasiticus (Speare) Kurtzman et al; A. nomius Kurtzman et 
al; and A. tamarii) (Dorner, 1983; Dorner et al., 1964; 
Kurtzman et al., 1987; Wicklow and Cole, 1982; Wicklow and 
Shotwell, 1983). 


We examined two sclerotium producing strains of S. 
togoense: NRRL 13,551 (=LCP 67.3456; CBS 205.75) isolated 
by M.F. Roquebert from a seed of Landolphia sp. 
(Apocyanaceae) collected by R. Heim in 1967, near La 
Maboke, Central African Republic; and NRRL 13,550 (=LCP 
64.1910) isolated by J. Nicot from undetermined seeds 
collected by R. Heim in 1964, near La Maboke, Central 
African Republic. Previous work has shown that substantial 
quantities of A. fiavus group mycotoxins can be extracted 
from A. flavus sclerotia (Wicklow and Cole, 1982; Wicklow 
and Shotwell, 1983; Wicklow et al. 1988). Gram quantities 
of sclerotia were consistently produced by either strain 
when inoculated onto moistened, steam sterilized oatmeal or 
oat seed (D.T. Wicklow and C.E. McAlpin, unpublished). 
Samples of air-dried sclerotia (3 g) produced on oatmeal by 
both NRRL 13,550 and NRRL 13,551 were separately extracted 
and analyzed for A. flavus group mycotoxins (see above) 
according to the method described by Wicklow and Cole 
(1982). These mycotoxins were not detected in the 
sclerotia or in methanol extracts of the fermented oats. 


Strain NRRL 13,550 was inoculated onto steam-sterilized 
rice (30% moisture content) and incubated at 25 C for 6 
weeks. No sclerotia were produced by the fungus on rice. 
However, approximately 600 ug of sterigmatocystin was 
produced per gram of rice, determined by thin-layer 
chromatography as described by Vesonder and Horn (1985). 
Sterigmatocystin was isolated from the methanol extract of 
the rice and crystallized from ethyl acetate as pale yellow 


251 


needies melting at 245 C. Conclusive identification was by 
comparisons of melting point, infrared (IR) and mass 
spectrum with that of a standard. No cyclopiazonic acid or 
aflatoxins were detected in methanol extracts of the 
fermented rice. Sterigmatocystins are produced by several 
species of Aspergillus, including some with the teleomorph 
State Emericella, the dematiaceous hyphomycete oipesaeis 
(=Dreschlera), and certain species of the ascomycet 
Chaetomium (Hamasaki and Hatsuda, 1977; Rabie et iy 1977; 
Udagawa et al. 1979). Sterigmatocystin nas been proposed 
aS an intermediate in aflatoxin biosynthesis and 
14C-labeled sterigmatocystin is converted by Aspergillus 
Parasiticus*into aflatoxin Bl (Hsieh et al., 1973). At the 
Same time, Ssterigmatocystin is not known to be produced by 
any species assigned to the Aspergillus ochraceus group 
(Cole and Cox, 1981) with which S. togoense also shares 
outstanding characteristics (Christensen, 1982). These 
analyses provide additional evidence linking S. togoense to 
the A. flavus group (Samson and Seifert, 1985). 


ACKNOWLEDGEMENT 


We take great pleasure in celebrating the pre-eminent 
mycological career of Professor Harry D. Thiers on his 70th 
birthday. 


LITERATURE CITED 


Christensen, M. 1982. The Aspergillus ochraceus group: two 
new species from western soils and a synoptic key. 


Mycologia 74: 210-225. 
COte,; 7 R.J. and R.H. Cox. 1981.9 Handbook. of Toxic Fungal 


Metabolites, Academic Press, New York. 


Dorner, J.W. 1983. Production of cyclopiazonic acid by 
Aspergillus tamarii Kita. Appl. Environ. Microbiol. 
46: 1435-1437. 


Dorner, J.H., R.J.~ Cole, and U.L. Diener. 1984. The 
relationship of Aspergillus flavus and Aspergillus 
parasiticus with reference to production of aflatoxins 
and cyclopiazonic acid. Mycopathologia 87: 13-15. 


Hamasaki, T. and Y. Hatsuda. 1977. Sterigmatocystin and 
related compounds, pp. 597-607. In: Mycotoxins in 
Human and Animal Health, J.V. Rodricks, C.W. 
Hesseltine and M.A. Mehlman (Eds.) Pathotox 
Publishers, Inc., Park Forest South, Illinois. 807pp. 


Hsien, 7D. P.H., MeTeoLin, and’ R.C. Yao.. ©1973. “Conversion 
of sterigmatocystin to aflatoxin Bl by Aspergillus 


parasiticus. Biochem. Biophys. Res. Commun. 52: 
992-997. 


252 


Kurtzman, C.P., B.W. Horn, and C.W. Hesseltine. 1987. 
Aspergillus nomius, a new aflatoxin-producing species 


related to Aspergillus flavus and Aspergillus 


tamarii. Antonie van Leewenhoek 53: 147-158. 


Rabie, C.J., M. Steyn, and G.C. van Schalkwyk. 1977. New 
species of Aspergillus producing sterigmatocystin. 
Appl. Environ. Microbiol. 33: 1023-1025. 


Raper, K.B. and D.I. Fennell. 1965. The genus 
Aspergillus. Williams and Wilkins, Baltimore. 


Roquebert, M-F. and J. Nicot. 1985. Similarities between 
the genera Stilbothamnium and Aspergillus, pp. 
221-228. In: Advances in Penicillium and Aspergillus 
Systematics, R.A. Samson and J.I. Pitt (eds.) Plenum 
Press, New York, and London. 483 pp. 


Samson, R.A. and K.A. Seifert. 1985. The ascomycete genus 
Penicilliopsis and its anamorphs, pp. 397-426. In: 
Advances in Penicillium and Aspergillus Systematics, 
R.A. Samson and J.I. Pitt (eds.) Plenum Press, New 
York, and London. 483 pp. 


Udagawa, S., T. Muroi, H. Kurata, S. Sekita, K. Yoshalmere: 
and S. Natori. 1979. The production of chaetoglobos- 
ins, sterigmatocystin, O-methylsterigmatocystin, and 
chaetocin by Chaetomium spp. and related fungi. Can. 
Jie Microbiol.) 253) 170-177. 


Vesonder, R.F. and B.W. Horn. 1985. Sterigmatocystin in 
pri iPeca contaminated with Aspergillus versicolor. 
Appl. Environ. Microbiol. 49: 234-235. 


Wicklow, D.T. and R.J. Cole. 1982. MTremorgenic indole 
metabolites and aflatoxins in sclerotia of Aspergillus 
flavus: an evolutionary perspective. Can. J. Bot. 
60: 525-528. ith 


Wicklow, D.T. and O.L. Shotwell. 1983. Intrafungal 
distribution of aflatoxin among conidia and sclerotia 


of Aspergillus flavus and Aspergillus parasiticus. 
Cand  EMIGroObLOINe 2975. 


Wicklow, D.T., P.F. Dowd, M.R. Tepaske, and J.B. Gloer. 
1988. Sclerotial meabelcad of Aspergillus flavus 
toxic to a detritivorous maize insect (Carpophilus | 
hemipterus, Nitidulidae). Trans. Br. Mycol. Soc. 91: 
(in press). 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 253-257 January 20, 1989 


THE OCCURRENCE OF VESICULAR-ARBUSCULAR MYCORRHIZAE IN 
BURNED AREAS OF THE SNAKE RIVER BIRDS OF PREY AREA, IDAHO. 


MARCIA WICKLOW-HOWARD 
Biology Department 
Boise State University 
BOLS ee LUD Ood Lo 


SUMMARY 


Soils from 1981, 1983, and 1985 burn 
sites and an unburned site within the 
Snake River Birds of Prey Area were 
collected. All sites were vegetated 
primarily with Artemisia tridentata 
Nutt. var. wyomingensis (Beetle and 
Young) Walsh (big sage) which com- 
monly forms vesicular-arbuscular 
mycorrhizae (VAM) for its healthy 
growth. These soils were inves- 
tigated using pot culturing methods 
to determine if they contained active 
VAM fungal propagules. The results 
from pot cultures inoculated with the 
1981 and 1983 soils showed that fun- 
gal propagules were present and 
actively infected 100% of the roots. 
With the 1985 burn soils, infection 
was much lower; only 50% of the roots 
from the pot cultures became infec- 
ted. 


Wildfires have beome an increasingly common occur- 
rence within the Snake River Birds of Prey Area. Over 
the past ten years, 37% (90,000 acres) of the sagebrush- 
dominated rangeland has burned; and 80% (72,000 acres) of 
this burned in the past five years. Artemisia tridentata 
Nutt. var. wyomingensis (Beetle and Young) Walsh (big 
sage) is the dominant variety of sagebrush present. 

After a burn, the previously sagebrush-dominated area is 
revegetated by big sage and invading non-native grasses, 


254 


particularly cheat grass (Bromus tectorum L.). After 
each fire more big sage is replaced by cheat grass, and 
eventually the area may be completely overrun by cheat 
grass. Cheat grass burns very quickly and therefore is 
hard to control. Repeated fires within these cheat : 
grassrdominated regions result in a ripple effect. The 
fire burns the cheat grass and a zone of previously 
undisturbed big sage surrounding it. With each consecu- 
tive fire, these areas increase in size and more big sage 
is lost (Winward, 1985). 


The big sage community supports high densities of 
black-tailed jack rabbits (Lepus californicus Gray). 
Jack rabbits are the main prey of golden eagles (Aquila 
chrysaetos L.), and research has shown a strong relation- 
ship between eagle productivity and jack rabbit abundance 
(U.S. Dep. Interior, 1979). The severe decline in jack 
rabbit numbers in recent years is thought to have been 
facilitated by the extensive loss of big sage habitat. 
There are concerns that the large loss of big sage may 
continue to maintain low jack rabbit numbers and result- 
ing golden eagle numbers. 


A positive solution to the problem is to revegetate 
these areas with the big sage soon after a burn, before 
the non-native grasses take a stronghold. The purpose of 
this study was to determine if VAM fungi, which are 
essential for the healthy growth of A. tridentata var. 
wyomingensis are present and remain active in soils after 
experiencing a recent burn. 


MATERIALS AND METHODS 


Soil samples, taken to depths of 15 cm, were col- 
lected from the rhizosphere of Artemesia tridentata var. 
wyomingensis at four different sites. The first collec- 
tion site was from an unburned control area vegetated 
primarily with big sage. The remaining three sites were 
vegetated primarily with the big sage before experiencing 
a burn in 1981, 1983, or 1985. Three plots from each 
Site were randomly selected. 


Soil samples were used as inoculum for pot cultures 
the day after collection. For each plot, two pot cul- 
tures were inoculated, resulting in six pot cultures per 


205 


Site. An equal amount of inoculum soil was mixed with 
500 g of sterilized, sandy garden soil. One control from 
each plot, in which the inoculum soil was sterilized as 
well, was prepared. All were then planted with surface 
sterilized seeds of sudan grass (Sorghum vulgare Pers 
var. sudanese Hitchc.). Pot cultures were watered, and 
received sixteen hours of light, daily. 


The sudan grass was harvested three months after 
planting the seeds. Roots were removed with approxi- 
mately 50 g of soil. This soil was washed from the roots 
using a strong stream of distilled water, and collected 
for wet sieving, following the sucrose centrifugation 
procedure in Daniels & Skipper, 1982. 


Following the procedure of Phillips and Hayman 
(1970), sudan grass roots fixed in FAA were cleared and 
stained with trypan blue. The roots were then examined 
for mycorrhizal propagules. 


RESULTS AND DISCUSSION 


Table 1. shows the incidence of VAM infection within 
sudan grass roots. The results from pot cultures inocu- 
lated with the 1981 and 1983 soils show that fungal 
propagules were present and actively infected 100% of the 
roots. With the 1985 burn soils, infection was much 
lower. Only 504 of the sudan grass roots from the pot 
cultures became infected. 


VAM spores were found within the filtrate of all 
soil samples, including controls, pointing toward pos- 
sible contamination. However, only one of the twelve 
control soils had infected sudan grass roots (Table 1.), 
suggesting that sterilization of the soils inactivated, 
but did not disrupt or destroy spores. The VAM spore 
occurring most frequently in all soil filtrates was iden- 
tified as the asexual chlamydospore of Glomus microcarpus 
Tul. & Tul. The spore is smooth, globose hyaline to 
light yellow, and approximately 50 um in diameter. It is 
attached singly to a hyphal extension. A large spore 
with a suspensor-like bulb attached to its base was 
found. This spore fit the description of the azygospores 
of the genus Gigaspora Gerdemann & Trappe. 


256 


Table l. 
VAM infection of sudan grass roots from pot cultures. 


Soil 1981 1983 1985 


Plots Unburned Burn Burn Burn 
la + + + - 
lb * * + — 
2a + + + + 
2b * + + + 
3a + + + - 
3b + + * - 

Control 1 x + - - 

Control 2 a ce fs - 

Control 3 r: * - o 


* = no plant 


Also investigated was the incidence of VAM infection 
within recently burned A. tridentata var. wyomingensis 
roots from two of the 1985 sites. One site experienced 
fire of low intensity, while the other a fire of higher 
intensity. 67% of roots from the low intensity burn were 
infected, while only 18% of roots from the higher inten- 
sity burn were infected. 


The preceding data suggest that VAM fungi are 
affected more adversely by a high intensity burn and 
that they recover from a burn within at most two years. 
However, after two years have passed, these areas are 
often highly invaded by cheat grass which does not nor- 
mally form mycorrhizae. Therefore, areas experiencing a 
low intensity burn in which the mycorrhizae are not sig- 
nificantly reduced, should be revegetated that same year 
before cheat grass invasion. In high intensity burn 
areas in which mycorrhizae are significantly reduced, 
revegetation programs should begin one to two years after 
the fire, giving the mycorrhizal propagules time to 
increase in numbers. In this case, steps would have to 
be taken to eradicate the cheat grass before reseeding. 


2o/ 


LITERATURE CITED 


Daniels, B.A.; Skipper, H.D. 1982. Methods for the 


recovery and quantitative estimation of propagules 
from soil. In: Methods and Principles of Mycor- 
rhizal Research. Sclenck, N.C. (Ed.), pp. 29-45. 
Minnesota: The American Phytopathological Society, 
Minn. 


Phillips, J.M.; Hayman, D.S. 1970. Improved procedures 


USS. 


for clearing roots and staining parasitic and vesic- 
ular-arbuscular mycorrhizal fungi for rapid assess- 
ment of infection. Trans. Br. Mycol. Soc. 55(1): 
158-160. 

Dept. Interior. 1979. Snake River Birds of Prey 
special research report. Bur. Land Manage. Boise, 
Idaho. 142 pp. 


Winward, A.H. 1985. Fire in the sagebrush-grass ecosys- 


tem - the ecological setting. Pages 2-6 In: K. 
Sanders, J. Durham, eds. Rangeland Fire Effects: 
A Symposium. Idaho State Office. U.S. Bur. Land 
Manage., Boise, Idaho. 


i , We i. 7) eae tal x / ea 
a? iy , 


bod Se aM) pitty. 2 ae rane ; 

. . ‘. > 
hers yah 8 bs os gee 
Won yore ah ap ann ae ee 


4? 


rae 3 koatd 


MYCOTAXON 


Vol. XXXIV, No. 1, pp. 259-266 January 20, 1989 


COMPATIBILITY AND FRUITING STUDIES OF AN ALBINO FORM 
OF AURICULARIA CORNEA 


George J. Wong 
Department of Botany 
University of Hawaii 

Honolulu, Hawaii 96822 


SUMMARY 


Compatibility and fruiting studies were carried out with an 
Auricularia collection with white basidiocarps. Aside from 
the basidiocarp color, this collection was morphologically 
referable to Auricularia cornea. Mating studies determined 
this stock to be bifactorial and compatible with normally 
pigmented stocks of A. cornea. Fruiting studies 
demonstrated that the white basidiocarp characteristic is 
inherent rather than an environmentally induced feature. 
Interstock crosses between pigmented and white strains 
produced hybrid basidiocarps with normal pigmentation. 
Only pigmented basidiocarps were observed from in- 
trastock crosses of compatible strains derived from hybrid 
basidiocarps. Based on mating and fruiting studies, it is 
concluded that the white basidiocarp collection is an albino 
form of A. cornea. Since only pigmented basidiocarps 
were observed in crosses between pigmented and white 
strains, a single gene, composed of a pair of dominant and 
recessive alleles, is thought to control pigmentation and al- 
binism in basidiocarp color respectively. However, a 
Mendelian 3:1 ratio was not observed, and white 
basidiocarps were not recovered in self crosses of isolates 
derived from hybrid basidiocarps. The absence of white 
basidiocarps is thought to be due to the low number of 
compatible crosses utilized in this study. 


INTRODUCTION 
The first white species of Auricularia Bull.:Merat was described in 


1872, as A. albicans Berk. (in Lloyd, 1925). There have since been two 
more white species of Auricularia described, A. leucochroma by 


260 


Kobayasi (1942), and A. eburnea by Li and Liu (1985). Auricularia al- 
bicans and A. leucochroma are presently recognized by Kobayasi (1981) 
as albino forms of A. auricula-judae (Bull.:Fr.) Schroet. and A. 
polytricha (Mont.) Sacc., respectively. 

Incompatibility studies have only been carried out in four species of 
Auricularia. A. auricula-judae, A. polytricha, A. delicata (Fr.) P. Henn., 
and A. fuscosuccinea (Mont.) Farl. These studies have been reviewed by 
Wong and Wells (1987). 

Other than in cultivation of Auricularia polytricha, there have been 
few fruiting studies carried out in this genus (Cheng and Tu, 1978; 
Quimio, 1982). Cheng and Tu (1979) described, macroscopically, 
basidiocarp development of A. polytricha grown in a sawdust and rice 
bran medium. Quimio (1982) studies the effects of aeration, light, and 
substrate supplementation on basidiocarp development. Hung (1983) de- 
scribed, macroscopically, basidiocarp development of A. polytricha, A. 
fuscosuccinea, and A. mesenterica. Mating and fruiting studies by Wong 
and Wells (1987) determined that the morphological differences between 
A. cornea, A. polytricha, and A. tenuis sensu Lowy (1951, 1952) and 
Kobayasi (1981), were environmentally induced, rather than inherent 
characteristics. Thus, the three species, in the sense of these authors are 
synonymous, and will, hereafter, be referred to as A. cornea, since it is 
the earliest species name validly published (Lowy, 1952). 

A specimen of Auricularia (GJW 820) with white basidiocarps, mor- 
phologically referable to A. cornea [=A. polytricha f. leucochroma 
(Kobayasi) Kobayasi] (Fig.1), was collected in Hawaii, on the island of 
Oahu, along the Manoa Falls Trail. Based on the morphology, in cross 
section, A. leucochroma basidiocarps, Kobayasi (1981) reduced this 
species to a form of A. cornea under the name A. polytricha f. 
leucochroma. However, this has not been substantiated with genetic evi- 
dence. In order to determine if GJW 820 is referable to A. cornea, in- 
trastock and interstock mating and fruiting studies were carried out. 
The criterion of compatibility between interstock crosses has been 
demonstrated as a useful tool in defining species in a number of studies 
(Boidin, 1986; Boidin and Lanquetin, 1984a,b; Capellano, 1985). How- 
ever, partially sterile hybrids have been observed in interspecific crosses 
between Typhula idahoensis Rembsberg and T. ishikariensis Imai 
(Christen and Bruehl, 1979; Bruehl and Machtmes, 1980). Fruiting 
studies of this collection would also determine if the white basidiocarp 
condition is an inherent, or an environmentally induced condition. 
Auricularia cornea is defined here as having a medullated basidiocarp 
with abhymenial hairs approximately 200-600 Im in length. Lowy 
(1951, 1952) and Kobayasi (1981) also used the width and morphology 
of the medulla in defining species, but these characteristics have been 
demonstrated to vary according to environmental conditions and to in- 
tergrade in presently recognized species of Auricularia (Wong and Wells, 
1987; Wong, unpubl.). 


261 


2 


Figs. 1-2. Auricularia basidiocarps. 1. White basidiocarps of GJW 
BZUweiy2s. 2. Basidiocarps of. GJW 820) (1. x»2))on right and 
basidiocarps of GJW 669 (9 x 11) on left. 1/3X. 


MATERIALS AND METHODS 


The method utilized for single spore isolation has previously been 
discussed (Wong and Wells, 1987). Mating strains derived from normally 
pigmented basidiocarps of A. cornea, GJW 802-18 (A,,B,,), GJW 802-16 
(A,B,3), GJW 669-9 (A,B) and GJW 669-11 (A,B,), were isolated and 
identified by Wong and Wells (1987). Isolates were maintained on malt- 
yeast-soytone agar (MYP; Bandoni et al., 1975). Inocula and matings 
were made on 3% malt agar (MA; Wong and Wells, 1987). Crosses were 


SEE ERRATA 
262 


made by cutting out 7 mm diameter plugs with a cork borer from the 
margins of inocula, and placing the inocula approximately 8 mm apart. 
After about 21-30 days, a small piece of agar was taken where the in- 
ocula have grown together, mounted in 3% KOH, and examined under 
phase optics. 

The method used for fruiting has previously been described by 
Wong and Wells (1987). The substrate was made up of 6 parts sugar 
cane bagasse and 1 part Old Fashioned Quaker Oats. The sugar cane 
bagasse-rolled oats medium was thoroughly mixed with an equal volume 
of tap water, and excess water squeezed out, before placing the mixture 
in a 1000 ml beaker. The beaker was covered with a 23.5 cm? sheet of 
aluminum foil placed on top of a paper towel of equal area and held in 
place with a rubber band. The beaker was then autoclaved at 121°C for 
90 min, followed by a 12 h cooling period. Following inoculation of the 
medium, the beaker was placed in the dark at 25°C until the mycelium 
had grown throughout the substrate. The substrate was then removed 
from the beaker and placed in a greenhouse with diffused lighting and 
an automatic misting system to keep the mycelium moist. 

Specimens used in this study have been deposited at BISH, and rep- 
resentative mating strains from each stock have been stored at ATCC. 

Hymenium colors, in brackets, of GJW 669, GJW 802 and 
basidiocarps formed from fruiting studies were cited from Ridgway 
(1912). 


RESULTS 


Intrastock crosses determined that GJW 820 was bifactorial. The 
results are summarized, and arranged according to their mating strains 
in Table I. The reactions are the same as those previously observed for 
A. cornea (Wong and Wells, 1987). Three types of reactions were ob- 
served in the contact zones where the monokaryons grew together: 


AJB]: Clamp connections at most septa. 
A]|B=: False clamps at most septa. 
A=B], or A=B=: Simple septa throughout. 


Clamp connections were observed only in the contact zones. Nuclear 
migration apparently did not occur. 

Compatible crosses of GJW 820 produced only white basidiocarps, 
while GJW 669 (9 x 11) and GJW 802 (16 x 18), under the same condi- 
tions, produced only pigmented basidiocarps (Fig. 2). Crosses between 
pigmented strains of A. cornea and GJW 820, GJW 820-1 x GJW 669-11 
and GJW 820-1 x GJW 802-18, were compatible, and produced fertile, 
pigmented basidiocarps. Twelve single spore isolates were obtained 
from each hybrid basidiocarp, and crossed in all possible combinations 
in order to identify compatible crosses. Only pigmented basidiocarps 
were produced in the latter compatible crosses. 


ERRATA 


The guest co-editors of MYCOTAXON 34(1) wish to correct some editorial 
oversights. These oversights were the responsibility of the guest co-editors and NOT 
the fault of the respective authors. 


*On p. 262, lines 30-32 should read: 


A#B#: Clamp connections at most septa. 
A#B=: False clamps at most septa. 
A=B#, or A=B=: Simple septa throughout. 


nyc Soi 
a 


ee es 


gen - 
FER eae a 


Le 


— 


— 


— 


—=AMNWNCWORNOW IH 


Table I 


Pairing Reactions of GJW 820 


1 6 2 4 5 7 8 10 ui 

1 ~ - + + + + £ + + 

6 - i in F F Ia) FE F 

) Ee = is 3 " so es 

4 Bs os so a a = 

5 = rs re > e 

7 # # s 

8 2 is a 
10 - ~ 
1] - 
12 

3 

9 

Symbols: 


1. -: Contact zone with simple septa. 
2. F: Contact zone with false clamps at most septa. 
3. +: Contact zone with clamp connections at most septa. 


Table II 


Pairing reactions 
of of 
GJW 820-1 X GJW 669-11 


meeseseile a DELO 2 5) 6 1) Lae ee el OY Ld 
se Seale Ee See Sad ci Sl Send Ss Gk ec a a 
AM a RE RR) Roy BQ ar Ligc ti arsenite (i aiboa 
Sreim ACe t tAPOCR FoF) 4 - - - - = 
a al eee SP eae oe ee minha ee sy Lin 
eee Paks Ee bebe yack | O - - = 
eR A ee a eat {) Lg re 
Sa ah ET Z 

- - - - - 7 

She Wee NS 

- - - 8 

- - 12 

- 6 


* Denotes compatible crosses not producing basidiocarps. 


12 


Pairing reactions 


y! 


ww 


Jem oii 9 Bae 9 Ree Poo Bc eas ew 
Elli Bs 9 lg 9 borg Bley lr 9 oleae 


oo 


263 


GJW 820-1 X GJW 802-18 


12 6 


> tay! tee) “wolee no olelay ileg| 
mt tte tt tt 


264 


Three compatible crosses did not produce basidiocarps in GJW 820- 
1 x GJW 669-11. The dikaryons of these crosses were slow growing and 
unable to infiltrate the bagasse-rolled oats substrate. The results of the 
interstock mating and fruiting studies are summarized in Table II. 

The pigmented, field basidiocarps from Wong and Wells (1987) 
showed the most variability. GJW 669 was dark brown [Dark Livid 
Brown to Warm Blackish Brown], to brownish black [Dusky Brown to 
Blackish Brown (1)] when dry, and reddish brown [Mahogany Red to 
Chestnut] to brown [Auburn] when wet. GJW 802 was light brown 
[Cacao Brown to Walnut Brown] to brown [Hay’s Brown to Seal Brown] 
when dry, and orange brown [Sayal Brown to Snuff Brown] to reddish 
brown [Vinaceous-Brown, or Sorghum Brown to Dark Vinaceous-Brown] 
when wet. Basidiocarps produced under greenhouse conditions were 
uniform with respect to color. The following crosses: GJW 669 (9 x 
11), GJW 802 (16 x 18), GJW (820-1 x 669-11), GJW (820-1 x 802-18), 
GJW (820-1 x 669-11) self crosses, and GJW (820-1 x 802-18) self 
crosses, produced basidiocarps in the same color range. These 
basidiocarps were brown black [Dusky Brown to Blackish Brown (1)] to 
dark grayish black [Dusky Purplish Gray, or Dusky Neutral Gray] to 
black [Black] when dry, and dark reddish brown [Liver Brown, or 
Chocolate] to brown [Carob Brown, Burnt Umber, or Vandyke Brown] 
when wet. 


DISCUSSION 


The compatibility of interstock crosses between GJW 820 and 
pigmented stocks of A. cornea demonstrated that GJW 820 is referable to 
A. cornea. Hybrid basidiocarps, i.e., basidiocarps derived from GJW 
820-1 x GJW 669-11, produced morphologically normal basidiocarps 
with abundant basidiospores, and do not represent partially sterile 
hybrids as those observed in interspecific crosses between Typhula 
idahoensis and T. ishikariensis (Christen and Bruehl, 1979; Bruehl and 
Machtmes, 1980). 

The characteristic of the white basidiocarp is inherent as 
demonstrated by intrastock crosses of GJW 820 (Fig. 2). Under the 
same environmental conditions, intrastock crosses of GJW 669 and GJW 
802 produced only pigmented basidiocarps (Fig. 2). Thus, genetically, it 
has been verified that white basidiocarp variants that are 
morphologically referable to A. cornea are albino forms of this species as 
Kobayasi (1981) suggested. 

Hybrid basidiocarps produced in this study were pigmented and 
uniform in color. This suggests that pigmentation is controlled by a 
single gene in which there is a dominant-recessive relationship in the 
pigmented and albino phenotypes. However, only pigmented 
basidiocarps were observed in compatible crosses in Table II, but 
because of the small number of compatible crosses fruited, a 3:1 
Mendelian ratio of pigmented to albino basidiocarps should not be 
expected. 


265 


The occurrence of the A. cornea albino form in Hawaii is rare. In 
the five years that I have collected in Hawaii, GJW 820 has been the 
only albino collection of A. cornea observed. The albino form also 
appears to be rare elsewhere, since Lowy (1951, 1952) did not record 
the occurrence of albinism in his treatment of Auricularia. Kobayasi 
(1981) recorded albino forms of A. cornea, as well as several other 
Auricularia species, but did not comment on the frequency of their 
occurrence in nature. 

The verification of GJW 820, as an albino form of A. cornea places 
another synonym under this species name. However, this has been done 
in concept only. Because a type probably does not exist for A. cornea, a 
neotype must be designated before A. cornea, A. polytricha, A. tenuis, 
and the albino form for this species can be reduced to synonymy. 


ACKNOWLEDGEMENTS 


This study was supported by funds from the University of Hawaii 
Research and Training Revolving Funds. 


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MYCOTAXON 


Vol. XXXIV, No. 1, pp. 267-268 January 20, 1989 


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MYCOTAXON 


Vol. XXXIV, No. 1, pp. 269-276 January 20, 1989 


TYPE SPECIMENS OF AGARICS, BOLETES AND 
GASTEROMYCETES IN THE SAN FRANCISCO STATE 
UNIVERSITY HERBARIUM (SFSU) 


BARBARA M. THIERS and ROY E. HALLING 


Cryptogamic Herbarium 
New York Botanical Garden 
Bronx, NY 10458-5126 


Agaricus amicosus Kerrigan, Mycotaxon (this issue) 1989. COLORADO. Grand Co.: 
Robber’s Roost, 21 Aug 1983, Kerrigan 1258. 


A. arorae Kerrigan, Mycotaxon 22: 427. 1985. CALIFORNIA. Santa Cruz Co., in live 
oak duff; Cap. Ext. Cemetery, 25 Nov 1981, Kerrigan 1199. 


A. cuniculicola Kerrigan, Mycotaxon (this issue) 1989. COLORADO. Grand Co.: Wil 
low Creek Pass summit, 22 Aug 1983, Kerrigan 1271. 


A. fuscovelatus Kerrigan, Mycotaxon 22: 424. 1985. CALIFORNIA. San Francisco Co.: 
San Francisco, under Cupressus macrocarpus, SW corner of Sunset Ave. & Lawton 
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A. perobscurus Kerrigan, Mycotaxon 22: 432. 1985. CALIFORNIA. San Francisco Co.: 
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A. rubronanus Kerrigan, Mycotaxon 22: 426. 1985. CALIFORNIA. San Mateo Co: 
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A. sequoiae Kerrigan, Mycotaxon 22: 420. 1985. CALIFORNIA. Mendocino Co.: 
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A. smithii Kerrigan, Mycotaxon 22: 428. 1985. CALIFORNIA. Humboldt Co.: Patrick’s 
Pt. State Park, scattered in humus in mixed woods, 12 Nov 1967, Thiers 21494 


A. vinaceovirens Kerrigan, Mycotaxon 22: 422. 1985. CALIFORNIA. San Mateo Co.: 
Colma, in a 10’ ring in bare lawn under cypress and Acacia, 7 Oct 1977, Kerrigan 
730. 


Amanita breckonii Thiers & Ammirati, Mycotaxon 15: 156. 1982. CALIFORNIA. San 
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A. constricta Thiers & Ammirati, Mycotaxon 15: 163. 1982. CALIFORNIA. San Mateo 
Co.: San Francisco Watershed, scattered to solitary under madrone and oaks, 22 Jan 
1965, Brzckon 302. 


270 


A, magniverrucata Thiers & Ammirati, Mycotaxon 15: 161. 1982. CALIFORNIA. San 
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A. pachycolea Stuntz in Thiers & Ammirati, Mycotaxon 15: 158. 1982. CALIFORNIA. 
Mendocino Co.: Jackson State Forest near Mendocino, 14 Nov 1967, Thiers 21502. 


Arcangeliella desjardinii Thiers, Sydowia 37: 300. 1984 (1985). CALIFORNIA. 
Mendocino Co.: Jackson State Forest near Mendocino, gregarious along road cut, 
under Douglas fir, 22 Nov 1981, Thiers 44020. 


A. parva Thiers, Sydowia 37: 301. 1984 (1985). CALIFORNIA. Sierra Co.: approx. 6 mi 
N of Sattley on Hwy. 89, buried completely in soil under pines and firs, 14 Jun 
1983, Thiers 45906. 


A. saylorii Thiers, Sydowia 37: 302. 1984 (1985). CALIFORNIA. Eldorado Co.: Silver 
Fork Rd., off Hwy 50 at China Flat, hypogeous under Abies, 8 Jul 1983, Saylor 
1574. 


A. variegata Thiers, Beih. Sydowia 8: 383. 1979. CALIFORNIA. Mendocino Co.: Hwy 
20 between Ft. Bragg and Willits, gregarious in humus in mixed woods, 1 Dec 
1967, Thiers 21738 


Armillaria olida Thiers & Sundberg, Madrofio 23: 451. 1976. CALIFORNIA. Eldorado 
Co.: Crystal Basin Recreation Area, Hwy. 50, 6 May 1972, Thiers 28816. 


Boletus abieticola Thiers, Calif. Mushrooms: A field guide to the boletes, 39. 1975. 
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B. amyloideus Thiers, Calif. Mushrooms: A field guide to the boletes, 104. 1975. 
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14 Nov 1970, Thiers 26903. 


B. barrowsii Thiers & A. H. Smith in Thiers, Mycotaxon 3: 262. 1976. ARIZONA. 
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Lodge, 21 Aug 1971, Thiers 27816. 


B. citriniporus Halling, Mycologia 69: 206. 1977. CALIFORNIA. Amador Co.: Pine 
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B. haematinus Halling in Thiers & Halling, Mycologia 68: 980. 1976. CALIFORNIA. 
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271 


B. mendocinensis Thiers, Calif. Mushrooms: A field guide to the boletes, 84. 1975. 
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Boletus mottiae (as ‘mottii’?) Thiers, Calif. Mushrooms: A field guide to the boletes, 36. 
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B. orovillus Thiers & Kowalski in Thiers, Mycologia 58: 825. 1966. CALIFORNIA. 
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B, pulcherrimus Thiers & Halling, Mycologia 68: 977. 1976. CALIFORNIA. Mendocino 
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B. puniceus Thiers, Mycologia 57: 529. 1965, non Chiu, Mycologia 40: 217. 1948. 
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B. rubripes Thiers, Mycologia 57: 532. 1965. CALIFORNIA. Mendocino Co.: Jackson 
State Forest, scattered to gregarious in humus in dense mixed woods, 27 Oct 1962, 
Thiers 9299. 


B. smithii Thiers, Mycologia 57: 530. 1965. CALIFORNIA. Mendocino Co.: near 
Mendocino, Jackson State Forest, scattered to subcespitose in humus in dense 
mixed woods, 12 Nov 1960, Thiers 8208 (leg. H. D. & B. M. Thiers). 


B. spadiceus var. rufobrunneus Thiers, Calif. Mushrooms: A field guide to the boletes, 
92. 1975. CALIFORNIA. Del Norte Co.: Jedediah Smith State Park, gregarious in 
soil under conifers, 12 Nov 1966, Thiers 17766. 


B. roseibrunneus Thiers, Mycologia 49: 707. 1957. TEXAS. Grimes Co.: near Navasota, 
gregarious in sand, mixed woods, 29 Sep 1951, Thiers 1402. 


B. silvaticus Thiers, Mycologia 49: 708. 1957. TEXAS. Grimes Co.: near Navasota, gre- 
garious in sand, mixed woods, 29 Sep 1951, Thiers 1403. 


Cortinarius velatus Thiers & A. H. Smith, Mycologia 61: 532. 1969. CALIFORNIA. 
Fresno Co.: Huntington Lake, scattered in soil under conifers, 6 Oct 1965, Thiers 
13414, 


C. thiersii Ammirati & A. H. Smith, Mycotaxon 5: 389. 1977. CALIFORNIA. 
Tuolumne Co.: Pinecrest Campground, gregarious in humus under confiers, 15 Jun 
1965, Thiers 12633 (leg. Thiers, Breckon, & Ammirati). 


C. verrucisporus Thiers & A. H. Smith, Mycologia 61: 533. 1969. CALIFORNIA. 
Amador Co.: Silver Lake, solitary in humus under conifers, 9 Jun 1966, Thiers 
16919. 


Destuntzia saylorii Fogel & Trappe, Mycologia 77: 741. 1985. CALIFORNIA. Sierra 
Co.: Hwy 49 at Sierra City, Wild Plum Campground, buried in rocky soil in mixed 
forest, 9 Oct 1983, Saylor 1645. 


Dermocybe sierraensis Ammirati, Mycotaxon (this issue) 1989. CALIFORNIA. Alpine 
Co.: Carson Pass & Blue Lakes, gregarious under Pinus contorta, 6500 ft., 17 Aug 
1974, Thiers 32671. 


272 


Clitocybe foveolata Bigelow, Beih. Nova Hedwigia 81: 326. 1985. CALIFORNIA. San 
Mateo Co.: Huddart Park, gregarious to subcespitose in humus in mixed woods, 24 
April 1965, Thiers 12475. 


Clitocybe gibba var. occidentalis Bigelow, Beih. Nova Hedwigia 81: 293. 1985. CALI- 
FORNIA. San Mateo Co.: San Francisco Watershed land, gregarious in humus un- 
der hardwoods, 4 Dec 1964, Thiers 11823 Z 


C. stercoraria Bigelow, Beih. Nova Hedwigia 81: 364. 1985. CALIFORNIA. Marin 
Co.: Muir Woods National Monument, gregarious to subcespitose on dung (cow?), 
21 Dec 1966, Thiers 18135. 


C. thiersii Bigelow, Beih. Nova Hedwigia 81: 355. 1985. CALIFORNIA. San Mateo 
Co.: San Francisco Watershed land, gregarious on dead hardwood log, 23 Dec 1966, 
Thiers 18196. 


Elasmomyces Sstipitatus Peters, Mycologia 54: 112. 1962. CALIFORNIA. Mendocino 
Co.: Jackson State Forest, 5 Nov 1961, H. A. Peters 651. 


Gastroboletus amyloideus Thiers in Thiers & Trappe, Brittonia 21: 251. 1969. CALI- 
FORNIA. Sierra Co.: Yuba Pass, hypogeous under lodgepole pine and red fir, 29 
Sep 1967, Thiers 21117. 


G. citrinobrunneus Thiers, Beih. Sydowia 8: 384. 1979. CALIFORNIA. Tehama Co.: 
near Mineral, erumpent under white fir, 29 Sep 1975, Showers 2901. 


G. suilloides Thiers in Thiers & Trappe, Brittonia 21: 248. 1969. CALIFORNIA. Mono 
Co.: Mammoth Mountain Ski Lift Area, in humus under lodgepole pines, 11 Sep 
1967, Thiers 20991. 


G. xerocomoides Trappe & Thiers in Thiers & Trappe, Brittonia 21: 247. 1969. CALI- 
FORNIA. Sierra Co.: Yuba Pass, gregarious in humus in conifer woods, 14 Sep 
1965, Thiers 13163. 


Gomphus thiersii Petersen, Evol. Higher Basidiomycetes, 363. 1971. CALIFORNIA. 
Calaveras Big Trees State Park, Squaw Meadow Campground, gregarious in humus 
under pines, 15 Oct 1967, Thiers 21238. 


Gymnopilus luteocarneus Hesler, N. Amer. sp. Gymnopilus, 62. 1969. CALIFORNIA. 
Marin Co.: Alpine Lake, solitary to scattered on dead conifer logs, 30 Nov 1964, 
Thiers 11815. 


G. obscurus Hesler, N. Amer. sp. Gymnopilus, 42. 1969. CALIFORNIA. Mendocino 
Co.: near Mendocino, Jackson State Forest, scattered on dead log in mixed forest, 1 
Dec 1962, Thiers 9677. 


G. parvisquamulosus Hesler, N. Amer. sp. Gymnopilus, 35. 1969. CALIFORNIA. 
Tuolumne Co.: Stanislaus National Forest, Dodge Ridge Ski Area, gregarious on 
dead conifer log, 13 Jun 1964, Thiers 11238. 


G. subtropicus Hesler, N. Amer. sp. Gymnopilus, 41. 1969. MISSISSIPPI. Harrison Co.: 
Biloxi, Popp’s Ferry area, gregarious on dead log, 15 Jun 1959, Thiers 6345. 


G. thiersii Seidl, Mycotaxon (this issue) 1989. CALIFORNIA. San Mateo Co.: Wunder- 
lich Park, 13 Mar 1987, Seidl 2291. 


*On p. 273, to be inserted under Lactarius: 


L. manzanitae Methven, Mycologia 77: 475. 1985. CALIFORNIA. Mendocino Co.: 
Jackson State Forest, under Arctostaphylos, 21 Nov 1982, Desjardin 1775. 


L. riparius Methven, Mycologia 77: 476. 1985. CALIFORNIA. El Dorado Co.: Audrian 
Lake, under Abies, Alnus, and Salix, 3 Oct 1982, Methven 1715. 


L. scrobiculatus var. montanus Methven, Mycologia 77: 478. 1985. CALIFORNIA. El 
Dorado Co.: Audrian Lake, under Abies and Pinus, 3 Oct 1982, Methven 1716. 


*On p. 273, to be inserted under Leccinunn: 


L. arbuticola Thiers, Calif. Mushrooms: A field guide to the boletes, 126. 1975. CALI- 
FORNIA. Nevada Co.: near Nevada City, scattered to gregarious under madrone, 
11 Nov 1972, Thiers 30514. 


273 


Hydnellum pineticola K. A. Harrison, Canad. J. Bot. 42: 1226. 1964. MICHIGAN. 
Chippewa Co.: N of Sheephead Lake, 23 Aug 1961, A. H. Smith 63973 (ISO- 
TYPE). 


Hygrophorus albinellus Largent, Mycotaxon 23: 384. 1985. CALIFORNIA. Santa Cruz 
Co.: Boulder Creek, gregarious in soil under hardwoods, 30 Dec 1970, Thiers 
27077. 


H. perfumus Largent, Mycotaxon 23: 394. 1985. CALIFORNIA. Shasta Co.: Lassen 
Volcanic National Park, site 381, elev. 7000 ft., solitary in humus under conifers, 
25 Jun 1965, Thiers 12786. 


H. pyrophilus Largent, Mycotaxon 23: 397. 1985. CALIFORNIA. Shasta Co.: Mt. 
Shasta area, gregarious to cespitose in burned areas, 24 Dec 1964, Thiers 12011 
(leg. J. Ammirati). 


H. subellenae Largent, Mycotaxon 23: 401. 1985. CALIFORNIA. Mendocino Co.: Jack- 
son State Forest, near Mendocino, cespitose in humus in mixed woods, 6 Dec 1969, 
Thiers 24459. 


Lactarius californiensis Hesler & A. H. Smith, N. Amer. sp.. Lactarius, 331. 1979. 
CALIFORNIA. Mendocino Co.: Jackson State Forest near Mendocino, gregarious to 
subcaespitose in dense humus, 23 Nov 1962, Thiers 9538. 


L. cocosiolens Methven, Mycologia 77: 472. 1985. CALIFORNIA. Sonoma Co.: Reedy 
Ranch, Alexander Valley, Chalk Hill Rd., gregarious in duff under Quercus and 
Arctostaphylos, 18 Dec 1982, Methven 2367. 


L. argillaceifolius var. megacarpus Hesler & A. H. Smith, N. Amer. sp. Lactarius, 369. 
1979. CALIFORNIA. Santa Barbara Co.: Los Padres National Forest, Fremont 
Campground, more or less buried in soil under valley oak, 28 Jan 1967, Thiers 

18556. 


Leccinum aeneum Halling, Mycologia 69: 208. 1977. CALIFORNIA. Mariposa Co.: near 
Greeley Hill, in soil under Arctostaphylos, 15 Dec 1974, Thiers 33241. 


L. armeniacum Thiers, Mycologia 63: 268. 1971. CALIFORNIA. San Mateo Co.: San 
Francisco Watershed, solitary in humus under hardwoods, 23 Dec 1966, Thiers 
18193. 


L. brunneum Thiers, Mycologia 63: 269. 1971. CALIFORNIA. El Dorado Co.: Luther 
Pass, Grass Lake, gregarious in humus under aspens, 6 Sep 1967, Thiers 20753. 


L. californicum Thiers, Mycologia 63: 273. 1971. CALIFORNIA. El Dorado Co.: 
Luther Pass, Grass Lake, scattered in grassy meadow near aspens, 3 Sep 1969, 
Thiers 23624. 


L. constans Thiers, Calif. Mushrooms: A field guide to the boletes, 134. 1975. CALI- 
FORNIA. Santa Cruz Co.: Boulder Creek, solitary in soil under madrone, 6 Dec 
1970, Thiers 26971. 


L. insigne var. brunneum Thiers, Mycologia 63: 272. 1971. CALIFORNIA. Madera Co.: 
Agnew Meadows, scattered in soil under aspens, 18 Aug 1969, Thiers 23609. 


L. largentii Thiers, Calif. Mushrooms: A field guide to the boletes, 142. 1975. CALI- 
FORNIA. Humboldt Co.: Samoa peninsula, gregarious to cespitose under toyon in 
soil, 30 Oct 1971, Thiers 28531. 


274 


L. manzanitae Thiers, Mycologia 63: 266. 1971. CALIFORNIA. San Mateo Co.: San 
Francisco Watershed, gregarious in soil under madrone and manzanita, 1 Feb 1970, 
Thiers 24736 (leg. R. Keller). 


L. manzanitae var. angustisporum (as ‘angustisporae’) Thiers, Mycologia 63: 20/1941. 
CALIFORNIA. Mendocino Co.: Jackson State Forest, gregarious in humus under 
madrone, Thiers 8774. 2 


L. montanum Thiers, Mycologia 63: 274. 1971. CALIFORNIA. El Dorado Co.: Luther 
Pass, Grass Lake, gregarious in humus under apsens, 6 Sep 1967, Thiers 20757. 


L. subalpinum Thiers, Mycotaxon 3: 264. 1976. UTAH. Summit Co.: Wasatch National 
Forest, Hwy. 150, Bald Mtn. Pass and Beaver River, gregarious in soil under pines, 
7 Aug 1970, Thiers 26823. 


Lepiota luteophylla Sundberg, Mycologia 63: 79. 1971. CALIFORNIA. San Mateo Co.: 
San Francisco Water Dept. property, scattered to gregarious to caespitose in humus 
under Cupressus macrocarpa, 10 Feb 1970, Thiers 24760. 


L. thiersii Sundberg, Mycotaxon (this issue) 1989. CALIFORNIA. San Mateo Co.: San 
Francisco Water Dept. Property, 10 Feb 1970, Sundberg IIJ-19-87-A-2. (Isotype). 


Marasmiellus ramealis var. californicus Desjardin, Mycologia 79: 132. 1987. CALI- 
FORNIA. Los Angeles Co.: Marshall Canyon, San Dimas, 9 Mar 1983, G. Wright 
2941. 


Marasmius applanatipes Desjardin, Mycologia 77: 894. 1985. CALIFORNIA. Sierra 
Co.: Yuba Pass, densely gregarious to subcaespitose in deep duff under mixed con- 
ifers, 8 Oct 1983, Desjardin 2330. 


M. thiersii Desjardin, Mycologia 79: 123. 1987. CALIFORNIA. Amador Co.: off Hwy. 
88 near Pine Grove, elev. 2000 ft., scattered, insititious on leaves of Pinus sp., 25 
Nov 1983, Desjardin 2611. 


Melanoleuca reai var. texana Thiers, Mycologia 50: 521. 1958. TEXAS. Brazos Co.: A. 
& M. College campus, forming large fairy rings in soil, 4 Sep 1953, Thiers 19/3. 


Micromphale arbuticola Desjardin, Mycologia 77: 897. 1985. CALIFORNIA. Marin Co.: 
Samuel P. Taylor State Park, densely gregarious, insititious on living madrone 
bark, 27 Nov 1982, Desjardin 1839. 


M. sequoiae Desjardin, Mycologia 77: 894. 1985. CALIFORNIA. Mendocino Co.: junc- 
tion of Hwys. 408 & 409, gregarious to scattered in needles of redwood, 13 Nov 
1982, Desjardin 1740. 


Phylloporus boletinoides A. H. Smith & Thiers, Contr. monogr. N. Amer. sp. Suillus, 
105. 1964. FLORIDA. Alachua Co.: low hammock west side of Newnan’s Lake, E 
of Gainesville, solitary in deep sand humus under pines, 31 Jul 1958, Thiers 4960. 


Protogautieria substriata Thiers, Beih. Sydowia 8: 386. 1979. CALIFORNIA. Fresno 
Co.: Huntington Lake, solitary in soil under conifers, 5 Oct 1965, Thiers 13405. 


Psathyrella atrospora A. H. Smith, Mem. New York Bot. Gard. 24; 272. 1972. CALI- 
FORNIA. San Diego Co.: Kitchen Creek near Mesa Grande, cespitose in humus 
near rotten logs (oak), 11 Mar 1970, Thiers 25248. 


275 


P. ellenae var. yubaensis Thiers & A. H. Smith in Smith, Mem. New York Bot. Gard. 
24: 75. 1972. CALIFORNIA. Sierra Co.: Yuba Pass, humus, 26 Oct 1965, Thiers 
13870. 


P. griseopallida Thiers & A. H. Smith in Smith, Mem. New York Bot. Gard. 24: 417. 
1972. CALIFORNIA. San Diego Co.: San Luis Rey Campground, 10 Mar 1970, 
Thiers 25079 (ISOTYPE). 


P. lithocarpi A. H. Smith, Mem. New York Bot. Gard, 24: 305. 1972. CALIFORNIA. 
Santa Barbara Co.: Lake Cachuma, on dead logs, 27 Nov 1964, Thiers 11801 (ISO- 
LY PE). 


P. sequoiae Thiers & A. H. Smith in Smith, Mem. New York Bot. Gard. 24: 152. 1972. 
CALIFORNIA. Humboldt Co.: Prairie Creek State Park, 29 Dec 1966, Thiers 
18282. 


P. texensis A. H. Smith, Mem. New York Bot. Gard. 24: 404. 1972. TEXAS. Sam 
Houston National Forest, near Richards, 23 Feb 1952, Thiers 1486. 


Suillus acerbus A. H. Smith & Thiers, Contr. N. Amer. sp. Suillus, 103. 1964. CALI- 
FORNIA. San Francisco Co.: The Presidio, 2 Dec 1962, Thiers 9685 (leg. Largent). 


S. fuscotomentosus Thiers & A. H. Smith in Smith & Thiers, Contr. N. Amer. sp. 
* Suillus, 65. 1964. CALIFORNIA. Santa Cruz Co.: Felton, scattered to gregarious 
under ponderosa pine, 2 Nov 1963, Thiers 10759. 


S. glandulosipes Thiers & A. H. Smith in Smith & Thiers, Contr. N. Amer. sp. Suillus, 
86. 1964. CALIFORNIA. Mendocino Co.: near Mendocino, Jackson State Forest, 
gregarious to cespitose in pine forest, 3 Nov 1962, Thiers 9335. 


S. kaibabensis Thiers, Mycotaxon 3: 266. 1976. ARIZONA. Coconino Co.: Kaibab 
National Forest, Hwy 67 between Jacob Lake and Kaibab lodge, 21 Aug 1971, 
Thiers 27813. 


S. monticola (as ‘monticolus’) Thiers, Madrofio 19: 154. 1967. CALIFORNIA. Nevada 
Co.: Donner Summit, gregarious in soil under Abies and Pinus contorta, 22 Sep 
1965, Thiers 13248. 


S. occidentalis Thiers, Mycotaxon 3: 268. 1976. ARIZONA. Coconino Co.: Grand Can- 
yon National Park, North Rim, Point Sublime, gregarious under ponderosa pine, 19 
Aug 1977, Thiers 27775. 


S. pungens Thiers & A. H. Smith in Smith & Thiers, Contr. N. Amer. sp. Suillus, 92. 
1964. CALIFORNIA. San Francisco Co.: campus of San Francisco State Univer- 
sity, gregarious under Monterey pine, 2 Nov 1962, Thiers 9330. 


S. reticulatus Thiers, Calif. Mushrooms: a field guide to the boletes, 205. 1975. CALI- 
FORNIA. Madera Co.: Agnew Meadows, solitary in humus in pine woods, 18 Aug 
1969, Thiers 23616. 


S. riparius Thiers, Madrofio 19: 156. 1967. CALIFORNIA. Tuolumne Co.: Pinecrest, 
gregarious to cespitose in vicinity of dead logs and stumps and along edge of 
/  $tream, 28 Sep 1965, Thiers 13283. 


S. volcanalis Thiers, Madrofio 19: 158. 1967. CALIFORNIA. Lassen Co.: Lassen Vol- 
canic National Park, Butte Lake Campground, scattered to gregarious under Jef- 
frey pines, elev. 6000 ft., 28 Jun 1965, Thiers 12800. 


i 


* 


Rie 10 


S q i? % fi 
PES LORE’ % 


276 


S. wasatchicus Thiers, Mycotaxon 3: 270. 1976. UTAH. Summit Co.: Uintas Mts., Yel- 
low Pine Campground, scattered in soil near ponderosa pines and aspen, 21 Jun 
1970, Thiers 25618. 


Thaxterogaster thiersii Calhoun, Mycotaxon (this issue) 1989. CALIFORNIA. Marin 
Co.: Audubon Canyon Ranch, 20 Jan 1980, Calhoun 80-1475. 


Tulostoma thiersii J. E. Wright, Mycologia 79: 155. 1987. TEXAS. Kimble Co: Llano 
River bottom land near Junction, gregarious in humus, near decayed oak and 
pecan log, 13 Oct 1957, Thiers 4643. 


Tylopilus ammiratii Thiers, Calif. Mushrooms: a field guide to the boletes, 221. 1975. 
CALIFORNIA. Shasta Co.: Castle Crags, 21 Nov 1971, Ammirati 6177. 


T. humilis Thiers, Mycologia 58: 815. 1966. CALIFORNIA. Mendocino Co.: near 


Mendocino, Jackson State Forest, gregarious to cespitose in soil near bishop pine, 
11 Nov 1965, Thiers 14157. 


SPECIES NAMED FOR HARRY D. THIERS 


DENNIS E. DESJARDIN 


Department of Botany 
University of Tennessee 
Knoxville, TN 37996-1100 


Amanita thiersii Bas, Persoonia 5: 382. 1969. nom. nov. for A. alba Thiers, Mycologia 
49: 719. 1957, non Pers. per Vitt., Tent. mycol. s. Amanit. Ill. 18. 1826. 

Cladonia thiersii Hammer, Mycotaxon (this issue) 1989. 

Clitocybe thiersii Bigelow, Beih. Nova Hedwigia 81: 355. 1985. 

Cortinarius thiersii Ammirati & A. H. Smith, Mycotaxon 5: 389. 1977. 


Gomphus thiersii Petersen, Interfamilial relationships in the clavarioid and cantharelloid 
fungi. 363. 1971. in R. H. Petersen, Ed. Evol. Higher Basidiomycetes. 


Gymnopilus thiersii Seidl, Mycotaxon (this issue) 1989. 

Lactarius thiersii Hesler & A. H. Smith, N. Amer. sp. Lactarius. 510. 1979. 
Leptonia thiersii Largent, Bibliotheca Mycologica 55: 104. 1977. 
Marasmius thiersii Desjardin, Mycologia 79: 123. 1987. 

Psathyrella thiersii A. H. Smith, Mem. New York Bot. Gard. 24: 85. 1972. 
Ramaria thiersii Petersen & Scates, Mycotaxon 33: 138. 1988. 
Thaxterogaster thiersii Calhoun, Mycotaxon (this issue) 1989. 


Tulostoma thiersii Wright, Mycologia 79: 155. 1987. 


*On p. 276, to be inserted between lines 27 and 28: 


Lepiota thiersii Sundberg, Mycotaxon (this issue) 1989. 


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A FESTSCHRIFT IN HONOR OF 
HAIRY ID. THUUEIRS 


PRGEAC Ee MUN ea Tie cit at wey a tha let rey a aaa cana er 
Harry D. Thiers: Reminiscences about a teacher and friend ............... 
Remembering the morel grower: Ron Ower, 1939-1986 ................. 
Dermocybe, subgenus Dermocybe, section Sanguineae in northern 
Ge Fe Tae 7) MMA VE DRA i AGUA Ht De UNO UNION IPR a Joseph F. Ammirati 
Lichens of Mount Diablo State Park, Contra Costa County, California. 
Doris E. Baltzo 
The genus Rhodocybe: new combinations and a revised key to section Rhodo- 
phana in North America.. Timothy J. Baroni and David L. Largent 
Quality control factors for Alternaria allergens. 
Harriet A. Burge, Marion E. Hoyer, William R. Solomon, 
Emory G. Simmons, and Janet Gallup 
Thaxterogaster thiersii: a new secotioid species from California. 
Cornelia J. Calhoun 
Studies on Marasmius from eastern North America. II. New species. 
Dennis E. Desjardin and Ronald H. Petersen 
A: synopsis of Colomimnan boletes 0) oe) Cee ea ta eg Roy E. Halling 
Cladonia thiersii: a new lichen from California......... Samuel Hammer 
Studies in Agaricus IV: new species from Colorado.. Richard W. Kerrigan 
A new, lignicolous species of Entoloma (Entolomataceae, Agaricales) from 


Cf 29" 4 CRO ERE MO ME NRT A OLS aR DE BRR ACU Pe 2 David L. Largent 
Notes on the genus Pronibera: a) oo ee ee al eis David Malloch 
Notes on Clavariadelphus. II. New and noteworthy species from North 

Wid. t= Ota DN sa DUM PLAC om PERE HW OTe: Andrew S. Methven 
A key to the species of Inocybe in California....... Florence H. Nishida 
Folicolous fungi 8: Capnodium in California......... Don R. Reynolds 
A new species of Gymnopilus from northern California... Michelle T. Seidl 
The lichens of serpentine rocks and soils in California... .. Lorene L. Sigal 


Lepiota sensu lato in California. II. Species with a hymeniform pileipellis. 
Walter J. Sundberg 
Examination of Stilbothamnium togoense for Aspergillus flavus group 
TEV COLONIAS cuticle D. T. Wicklow, R. F. Vesonder, Cesaria E. 
McAlpin, R. J. Cole, and Marie-France Roquebert 
The occurrence of vesicular-arbuscular mycorrhizae in burned areas of the 
Snake River Birds of Prey Area, Idaho..... Marcia Wicklow-Howard 
Compatibility and fruiting studies of an albino form of Auricularia cornea. 
George J. Wong 
Master’s degree students of Harry D. Thiers and thesis titles.............. 
Type specimens of agarics, boletes and gasteromycetes in the San Francisco 
State University herbarium (SFSU). 
Barbara M. Thiers and Roy E. Halling 
Species named for Harry D. Thiers). ieee Dennis E. Desjardin 


Guest Co-Editors: ROY E. HALLING and BARBARA M. THIERS 


NWNre 


115 
119 


129 
133 


153 
181 
197 
217 
221 


239 


249 
253 
259 
267 


269 
276 


CCP 
W974 
yl 34 


O Ce 


/Vo. 2 


“MY COTAXON 


AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION 
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS 


Vol. XXXIV January-March 1989 No. 2 


CONTENTS 


The anamorphs of Diatrypella prominens and Eutypella sabalina. 
Dean A. Glawe and John P. Jones 277 
A preliminary account of Xylaria of Mexico. 
Felipe San Martin Gonzalez and Jack D. Rogers 283 
Varicosporium scoparium, a new staurosporous hyphomycete. 
A. Roldan and M. Honrubia 375 
A new addition to the genus Gorgomyces ...............44. A. Roldan 381 
The occurrence of Tuber texense in Georgia. 
Richard T. Hanlin, Mei-Lee Wu, and Timothy B. Brenneman 387 
A survey of Pilobolus from Yellowstone National Park. 
K. Michael Foos and Judith A. Royer 395 
Two new species of Parmelia (Parmeliaceae, Lichenes), further new 
combinations and notes, and additional new lichen records from southern 
I re Bec ys Ss rea a's NG an aS Re Franklin A. Brusse 399 
Pollution atmosphérique et lichens dans la ville de Santiago du Chile. 
Manuel Mahu 407 
Studies on xylophilous fungi from Argentina. IV. Anamorphs of Basidiomycetes 
on Eucalyptus viminalis (Myrtaceae). 
A. I. Romero, D. Cabral, and S. E. Lopez 429 
Synopsis of a revised classification for the Entomophthorales (Zygomycotina). 
Richard A. Humber 441 
Mnemonic three-letter acronyms for the names of fungal families. 
Joseph E. Laferriére 461 
Canalisporium, a new genus of lignicolous hyphomycetes from Malaysia. 
A. Nawawi and A. J. Kuthubutheen 475 
Quadricladium aquaticum gen. et sp. nov., an aquatic hyphomycete with 
Getravadiate conidia ......... A. Nawawi and A. J. Kuthubutheen 489 
A new taxon in Colispora (Hyphomycetes) from Malaysia. 
A. Nawawi and A. J. Kuthubutheen 497 


[CONTENTS continued overleaf] 


ISSN 0093-4666 MYXNAE 34 (2) 277-738 (1969) 


Published quarterly by MYCOTAXON, LTD., P. O. Box 264, Ithaca, NY 14851. 
For subscription details, availability in microfilm and microfiche, 
and availability of articles as tear sheets, see back cover. 


- 


GEE ERRA 


[CONTENTS continued from front cover] 


Ascomycetes of western India XII]........ Alaka Pande and V. G. Rao 503 
The genus Chaetomastia (Dacampiaceae) in North America. 

Margaret E. Barr 507 
The genus Dothidotthia (Botryosphaeriaceae) in North America. 

Margaret E. Barr 517 
Marine fungi from Seychelles. VIII. Rhizophila marina, a new ascomycete 


from mangrove prop roots........ K. D. Hyde and E. B. G. Jones 527 
Marine fungi from India. III. Acrocordiopsis patilii gen. et sp. nov. from 
Man Grove: WOOO FS e yale ca Ws earahale de B. D. Borse and K. D. Hyde 535 


New species in the lichen genus Xanthoparmelia (Ascomycotina: Parmeliaceae). 
Mason E. Hale 541 
Sugiyamaemyces, a new genus of Laboulbeniales (Ascomycetes) on 
Clidicus (Scydmaenidae) .... Isabelle I. Tavares and Jean Balazuc 565 
Index to J. B. Ellis’ types of pyrenomycetes with amyloid ascal rings. 
Katia F. Rodrigues 577 
Notes on tropical and warm temperate basidiomycetes. 
A. L. Welden and C. L. Ovrebo 601 
Amanita protecta — a new species from coastal southern California. 
Rodham E. Tulloss and Greg Wright 615 
Contribution to the study of the myxomycetes in Spain. I. 
Gabriel Moreno, Carlos Illana, and Michel Heykoop 623 
A new Ophiostoma on polypores .... O. Constantinescu and S. Ryman 637 
Lithographa, a lichen genus new to continental North America. . 
JoAnn W. Flock 643 
Notes on one lichenicolous and one fungicolous discomycete. 
Wen-ying Zhuang and Richard P. Korf 647 
Noteworthy corticolous lichens in Nothofagus forests, north-western Patagonia. 
Susana Cavelo and Laura Lorenzo 655 
Gigaspora ramisporophora: anew species with novel sporophores from Brazil. 
Joyce Lance Spain, Ewald Sieverding, and Norman C. Schenck 667 
A numerical taxonomic study of some phialidic genera of hyphomycetes: 


cluster analysis ..... Angélica M. Arambarri and Marta N. Cabello 679 
The genus Cladidium (lichenized Ascomycotina) ........ Bruce D. Ryan 697 
Author INDEX i )5.0 2 2 siaile Was bees) Be vk Lh eS 713 
INDEX to Fungous and Lichen Taxa 02 0) 6. 0 Js ed.s yk ee 716 
REVICWERS FF 8 ede leis Wha eae Rion ke Eola ad gO toe 
Publication dates; MYCOTAXON Volumes 33 and 34(1) .............. 732 
FETraea (iiss ee op alae bh Shae wae lah ee pS ad ator eek lect 738 


[MYCOTAXON for October-December, 1988 (33: 1-524) 
was issued November 28, 1988] 
[MYCOTAXON’ s special issue, the THIERS FESTSCHRIFT, (34(1): 1-276) 
was issued January 20, 1989] 


ERRATA, VOLUME THIRTY-FOUR 


Cover 2 line 29 for Cavelo read Calvelo 


Pages. iv 53 for Cavelo read Calvelo 
713 31 for Cavelo read Calvelo 
714 24 for Cavelo read Calvelo 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 277-281 January-March 1989 


THE ANAMORPHS OF DIATRYPELLA PROMINENS 
AND EUTYPELLA SABALINA 


Acquisiti 

~ Ons Divje; 

Dean A. Glawe RECE|Vep “StON 

Department of Plant Pathology ; 
University.ole Tilinois 


Urbana, IL 61801 MEY 04) 959 
and AIR MANN LIBRARY 


John P. Jones 
Department of Plant Pathology 
Louisiana State University 
Baton Rouge, LA 70803 


Previous studies have shown that species of 
Diatrypaceae often form anamorphs with both percurrently 
and sympodially proliferating conidiogenous cells (e.g., 
Glawe and Rogers, 1984; Abe, 1986; Glawe and Jacobs, 1987; 
Rappaz, 1987). Recently we obtained collections of 
Diatrypella prominens (Howe) Ell. & Everh. and Eutypella 
Sabarina) (Cooke) Ell: & Everh., and studied the anamorphs 
of these species. As detailed herein, anamorphs of both 
ereetypically diatrypaceous in their features of conidial 
and conidiomatal morphology, and conidiogenous cell 
proliferation. 

Cultures were started as described previously (Glawe 
and Rogers, 1984), and maintained on Difco potato dextrose 
agar at 20 C under fluorescent light with a daylength of 10 
h. Collections were deposited in ILLS. 


Diatrypella prominens, anamorph, Figs. 1-6. 


Anamorph on host not seen. Cultures covering 9-cm- 
diam culture plates in 14 days, producing conida in 30 
days. One-month-old colonies white, cottony; reverse off- 
white to yellowish. Conidiomata pycnidial, composed of 
loosely woven dark hyphae; subconic, black, producing gray 
conidial masses. Conidiogenous cells cylindrical, tapering 
toward apex, proliferating sympodially (Figs. 1, 2, 4), 
perc Urrentiy (Figs. S, 6). or) both ways, (10:4-)12-18.4 
Pere) ke CL 6—):2-2.4 ium. Conidiavhyaline, filiform, 
slightly to moderately curved (Fig. 3), with flat bases, 
moe o-34.4) X 1.6—-2.4 pm. 

Collections from which cultures were started: Jones, 
we Po,, 26.V.1986, Louisiana State University campus, Baton 


278 


Figs. l=6. Anamorph of Diatrypella prominens in 
culture. 1. Conidiogenous cells with lateral succession 
scars (arrows) indicative of sympodial proliferation. =2% 
Conidiogenous cell with new conidium (arrow) developing to 
Side of, maturesconidium- 98... Conidium. | 4. Api Galea. 
of conidiogenous cell with lateral conidial succession scar 
(arrow)* indicative of «sympodial® proliferation. Spee 
Conidiogenous cells with annellations (arrows) indicative 
of percurrent proliferation: All xX2500, by Nomarska@ 
differential interference contrast microscopy. 


24g 


Rouge, East Baton Rouge Parish, Louisiana, on Platanus sp. 
Glawe, D. A. I1.1987, West Washington Street, Urbana, 
Champaign Co., Illinois,.on Platanus’ occidentalis L. 


Bucypella Sabalina,. anamorph, Figs. 7-17. 


Anamorph on host: Conidiomata immersed in host, 
pycnidial, multiloculate (Pigf)7), 0.6-0.8 mm diam,’ lined 
with palisades of conidiogenous cells. Conidiogenous cells 
arising from interwoven hyphae or pseudoparenchyma, 
cylindrical, tapering toward apex, proliferating 
mma iaienyy (Figs. 8-10)" or percurrently,.8.8—-22.4 X 1..6- 
Peer. Conidia hyaline, filiform, moderately to strongly 
meee ee comee ToS 11s 12), swith flat. bases, 12-17.6 X° 0.'8-1%.2 
Weer Cudtures Covering 9-cm-diam Petri plates in about 10 
days, producing conidia in 19 days. One-month-old colonies 
white, felty, with gray to black areas of aerial hyphae. 
Conidiomata sporodochium-like, producing conidia in light- 
colored masses. Conidiogenous cells arising from interwoven 
hyphae or pseudoparenchyma, or borne on short, branched 
conidiophores arising from hyphae; cylindrical, tapering 
Powend apex, proliferating sympodially (Figs. 13, 14). or 
Gegourrent ly a(Pigs. 15-17), 12-24 X% 1.6-2.8 um., Conidia 
hyaline, filiform, moderately to strongly curved, with flat 
paeee 2 20.X%, 1.2-1..6. um. 

Collection examined, and from which cultures were 
Seeted-s Ones, J. P., L7/s11.1984, East Baton’ Rouge 
Parrshy, Louisiana, On Sabal minor (Jacq.) Pers. 


Illinois Agricultural Experiment Station Project 68- 
eemene thank Drs. L.(M.’ Carris and J. D. Rogers for 
reviewing the manuscript. 


ECE RATUREYGLTED 


Abe, Y. 1986. Notes on some comon xylariaceous and 
Blatrypaceous fungi. on hardwoods in Japan. “if. Bull. 
Romeo ater OLe Str lOd jeReS US be OOP mL aL . 

ewe moon) sand 1K. oA mJacobs./ 1987. Taxonomic) notes: on 
Eutypella vitis, Cryptosphaeria populina, and Diatrype 
StLgmaw Mvcologia 79 +838 5—139% . 

GlaweyeD. 9A., and: Js"D. Rogers. 1984. ‘Diatrypaceae in the 
Pacific Northwest. Mycotaxon 20: 401-460. 

Rappaz;, F' 1987. Taxonomie et nomenclature des 
Diatrypacees a asques octospores. Mycol. Helvet. 2: 
285-648. 


280 


Figs. 7-17. Anamorph of Eutypella sabalina. 7=12% 
From host material. 7. Longitudinal section through 
conidioma showing palisades of conidiogenous cells lining 
locules, ca. X250. 8. Conidiogenous cell withetageras 
conidial succession scar (arrow) indicative of sympodial 
proliferation. 9, 10. Apical regions of conidiogenous 
cells with conidia (arrows) developing to the sides of 
mature conidia. 11, 12.)/ Conidia. (13-17. From cuopeueeo 
material. 13. Apical region of conidiogenous cell with 
attached developing conidium. 14, 15. Conidiogenous meee 
with lateral conidial succession scars (arrows) indicative 
of sympodial proliferation. 16, 17. Conidiogenous celgs 
with annellations (arrows) indicative of percurrent 
proliferation. All by Nomarski differential interference 
contrast microscopy, X2500 unless otherwise noted. 


281 


Se & 
i, 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 283-373 January-March 1989 


A PRELIMINARY ACCOUNT OF XYLARIA OF MEXICO 
Felipe San Martin Gonzalez 


Instituto Tecnologico de Ciudad Victoria 
AD POS t.e NO .meL/ oS 
cd. Victoria, Tamaulipas, MEXICO 


and 
Jack D. Rogers 


Department of Plant Pathology 
Washington State University 
Pullman, WA 99164-6430 


ABSTRACT 


A preliminary account of Xylaria in Mexico is pre- 
sented, based primarily on collections from the states of 
Chiapas, Nuevo Leon, Quintana Roo, and Tamaulipas. The 
following new species are proposed: X. alata, X. 
amphithele, X. claviceps, X. guazumae, X. magniannulata, 
X. scabriclavula, X. squamulosa, and X. uniapiculata. A 
new variety, tropica, is proposed for X. longipes. In 
addition, nine taxa are discussed, but remain unnamed 
Owing to condition or size of the pertinent collections. 
Several of these undoubtedly represent new species. In 
tetal, 63 taxa are described and represented in keys. 
Most of them are depicted among 155 figures. 


Xylaria Hill ex Schrank is a cosmopolitan genus, 
reaching the zenith of its diversity in the various 
subtropical and tropical regions of the world. Until 
about 15 years ago it was widely considered that most of 
the species of Xylaria had been collected and were 
represented by specimens in various herbaria. Intensive 
collecting in Asian and American subtropics and tropics 
in recent years, however, has made clear that many taxa 
remain to be described. Indeed, Xylaria taxonomy has 
become vastly more complicated with the new additions 
from the tropics. For example, Rogers et al. (1988) 
published a treatment of Xylaria from Cerro de la 


284 


Neblina, Venezuela. Forty-one taxa were described from 
the rather small and circumscribed area, including 6 
formally named new taxa and 9 additional potentially new 
taxa, 1.e., about 37% of the taxa examined were unknown 
to the investigators. Dennis (1970) had presented keys” 
to 47 taxa from Venezuela and adjacent countries. 

Roughly half of the taxa described by Rogers et al. 
(1988) were not included by Dennis (1970). Similar 
patterns are emerging elsewhere in the subtropics and 
tropics, including Mexico. 

The present study was initiated in 1986 by FSMG in 
examining xylariaceous fungi in the herbarium of Instituto 
Tecnologico de Ciudad Victoria (ITCV). He, his col- 
leagues, and his students later collected extensively in 
the northeastern states of Nuevo Leon and Tamaulipas and 
the southeastern state of Quintana Roo and the south- 
southwestern state of Chiapas. These collections were 
brought to Washington State University in January, 1988 
by FSMG where he spent the next three months in collabor- 
ation with JDR, thanks to financial support from USAID. 
Tn addition, JDR had previously examined about 80 
specimens from herbarium XAL kindly provided by Gaston 
Guzman. A substantial number of those specimens were 
immature and could not be identified with certainty. 
Those that were of use to this study have been cited 
herein and are denoted with an asterisk. Unfortunately, 
none of the FSMG or XAL material was in culturable 
condition. Current collecting in México, however, will 
emphasize the collection and preparation of specimens for 
culture studies. 

We consider the present study to be preliminary. 
Nonetheless, 63 taxa are reported herein. Eight new 
Species, one new variety, and 9 unnamed taxa are described 
and illustrated, i.e., 28% of the collections were 
unknown to us, following the experience of JDR with 
Venezuela material (see earlier herein). The only other 
Study of Xylaria based exclusively on Mexican material, 
primarily herbarium specimens, is that of Pérez-Silva 
(1975) where 12 species were discussed. We have encounted 
every species discussed by Peérez-Silva except X. fockei 
(Miq.) Cooke. Comments on her concepts and data are 
included in the text. 

It is premature to draw far- reaching conclusions 
about the distribution of Xylaria species in Mexico. It 
is noteworthy, however, that X. Tongipes -- primarily a 
north temperate species -- was found in Tamaulipas state, 
a region where a mixing of northern and subtropical 


285 


species is not unexpected, especially at higher altitudes. 
An atypical specimen of X. longipes was collected in the 
southern state of Chiapas and 7s considered by us to be a 
new variety. Other temperate to subtropical species 
encountered in the northern states are X. magnoliae, X. 
oxyacanthae, X. persicaria, and X. polymorpha. The former 
3 species were collected Targely from expected hosts or 
host families. On the other hand, basically subtropical 
or tropical species such as X. anisopleura are widespread 
in Mexico as are representatives of the more cosmopolitan 
arbuscula complex. 

t 1s noteworthy that representatives of various taxa 
show ostioles whitened by chemical deposits. It is as- 
sumed that such a condition arises under conditions of 
low relative humidity and/or high concentrations of cer- 
tain minerals. It seems too widespread to be accorded 
taxonomic significance. 


SPECIES WITH LARGE STROMATA ON WOOD 
(Stromata usually 5 mm or greater in diameter) 


1.  Stromatal surface more or less smooth. ..... .2 
1. Stromatal surface more or less roughened 

with perithecial contours, coarse ostiolar 

papillae, wrinkles, or combinations of 

EnecOmred. GULCSs mayvoMe ince, Vite? fet Fella Fees Preise 11 


2. Fertile part subglobose to oblong, 
blackish with white polygonal patches, 
becoming hollow, on narrow tomentose 
stipe, 3.5-5 cm total length X 4.5-5 
mm diam. Ascospores (25-)29-33(-36) 

X 7-8(-9) um, with abrupt pinched 


ENS ie Me eit Sih. eee bee ilo tes Ste} ts X. aff. comosa 
2. Combination of features differing 
PGOMPODOVE dea ainsi dials LSE SES A a) BN ea 3 


au Fertile part subglobose to clavate, brown 
to dark-brown, becoming hollow and inrolled, 
on tomentose to smooth, attenuated stipe 
with discoid base, 2-6 cm total length X 
5-10 mm diam, almost smooth with smal] 
wrinkles and prominent hemispherical 
ostioles. Ascospores 22-32(-35) X 6-8 um, 
with ends abruptly pinched or rounded, 
and short, straight or oblique’ germ slit... ... A 
Gems rence he as we ee eae Ct et ne SCeN 


286 


Combination of features differing from above. . . .4 


4. Fertile part conical, on short to 
long, stout, pannose stipe, 1.8-6.8 
cm total length X 0.4-3 cm diam; 
whitish to yellowish, with umbili- 
cate or somewhat raised ostioles. 
Ascospores (19-)20-23(-25) X 8-9.5 
(-10) um, with broadly or narrowly 


rounded ends and spiralling germ slit. .... 

LE MA RMR Se em oO GOR clr gtad, X. aff. enterogena 
4. Fertile part cylindrical, clavate, 

or irregular sr! yee Ee ae 5 
Stromata at first copper-colored, often 
D LAGKGMING sine Seize on: oooh es, ope dneutie tacaintl Oy WSs UN oh ann 6 
Stromata at first .of.another.color.. <. 72 8 


6. Fertile part clavate on short stipe, 

1.7-5 cm total length X 7-10 mm diam, 

often becoming hollow and inrolled, 

with finely papillate ostioles. 

Ascospores (7-)8-10(-11) X 4-5(-6) um, 

the germ slit mostly not observed. ...... 

SW ieee ot (oe on oe a eee . . X.*cubensts 
NOTE: What appears to be another form of this 
species has stromata with polygonal surface scales, 
bigger perithecia and ascospores 9-12 um long. 
6. Ascospore germ slits usually con- 

Spicuous, straight. Combination of 

other features differing from above. ..... 7 


Fertile part short clavate, on short or 

long stipe with pannose base, 1.6-2.3 cm 

total length X 4-6 mm diam, becoming hol- 

low, with finely papillate ostioles. 

Ascospores (8:5-)9-11(-12) X 4-5 um’. 0°) see 


Fertile part clavate to more or less 

cylindrical on short stipe, 5-8 cm total 

length X 1-1.5 cm diam, with finely 

papillate ostioles. Ascospores (9-)10-12 

West teammiiib ag reas oe Go A Meese X. allantoidea 


* Ascospore germ slits may be assumed to be straight and 
oriented with the long axis of the spore and run 
essentially the length of the spore, unless noted 
otherwise in the key. 


287 


8. Fertile part clavate to irregular 
on short to obsolete stipe, 5.5-21 
cm total length X 2-4 cm diam, at 
first whitish and smooth but becom- 
ing blackish and rugose on drying, 
with hemispherical ostioles. Asco- 
spores 14-17(-18) X 5.5-6.5 um... X. poitei 
NOTE: Fungi with somewhat smaller stromata 
and larger ostiolar discs are referable to X. 
guyanensis. See description herein. a 
8. Fertile part generally smaller and 
less conspicuously wrinkled on drying. .... 9 


Fertile part clavate-fusiform, on short 

to long stipe, yellowish to dull orange, 

up to 6.5 cm total length X 4-9 mm diam, 

becoming hollow prior to perithecial 

maturation, with umbilicate ostioles. 

Ascospores (17-)18-21(-22) X (5.5-)6-7 

Havent pS NOGG. ROD 1a QUE GEYMiSAd Carsaas Bien fei po geen s,s 


Fertile part 4-13 cm high, not usually 
fusiform. Ostioles and surface colora- 
tion making stromata appear conspicuously 


or weakly longitudinally striped. ........ 10 
10. Ascospores (9.5-)10-12(-12.5) X 
RD =) Gar OP IM Be rene eps Bi che te Sats Boe X. grammica 


NOTE: Fungi with similar stromata and sterile 
apices have been called X. venustula. 
10. Ascospores (27-)28-33.5(-35) X 


67h Ol SHRM aWICOeSPI Vale Oermasl itvgitor.. oc. 

TT Re RE he Meine Rie ate fade o MMCtY o's X. kegeliana 
PSCOSDOLeS#eUSUd Vile eOne LONGCH araie, wk tease 12 
Ascospores usually not longer than 14 um..... 19 


12. Fertile part clavate, compressed 

with sterile acute apices, on 

distinct stipe, 4-9 cm total 

length X 2-5 mm diam, blackish 

with brown scales or peeling outer 

layer, with papillate ostioles. 

Ascospores (15.5-)16.5-18(-20) X 

6-472 5) UMS WitheSpLLa lmgermesidtacs: eve 

wks Mie Bisa sew cL icc tea at ces X. cf. corniculata 
12. Combination of features differing 

ECOMPADOVE seme. tere tae teen tee ae RSet Ns 


288 


3". 


ike) 


Rays 


or 


I 


Avy 


Fertile part broad clavate with rounded 
fertile apex, on long stipe, 10-11 cm 
total length X 1.5 cm diam, verrucose, 
black with white umbilicate to more or 
less hemispherical ostioles. Ascospores 
almost black, (12-)14-17(-18) X 6-7 um, 


with: acute to. pinched ends). 0.59.2 -) eswnenes cee 
Na vagal oihina' tial Salus Sve eibey bate wie «cee Pau s X. pallide-ostiolata 
Features of stromata and ascospores 
di FEES Epi y ise oeeltes vouireu vals etre o. of feb tate aie mena 14 
14. Fertile part cylindrical, clavate, 

or Spathutates. st hk ue ar) ee 15 


14. Fertile part subglobose to..e)lipsoid. <> ae be 


Fertile part cylindrical to clavate, on 

short to long stipe, 3.2-6.2 cm total 

length X 3-7 mm diam, blackish, becoming 

hollow, with conical or hemispherical 

ostioles often covered with whitish 

granules. Ascospores (13-)14-16 X 5-6 

um, with. spirallinggerm-slit®....° .©.i.% ....) eee 
sat eet ote emers X. longipes var. tropica var. nov. 
Fertile part similar to above, but with 

solid flesh and whitish to brown poly- 

gona}. surface.scalesirn. Vitis hee, ie.) tar 16 


16. Stromata 4-7 cm total lenath X 
5-9 mm diam. Ascospores 13-16 X 


4-5(-5.5) um, with spiral germ slits... .. . 

si CN ORS Wn te, ak ane rea eee ee eee X. longipes 
16. Combination of features differing — 

From: aADOVE ee NE A ae Nok eee 17 


Fertile part variable in shape, with 

Short or long stipe and sometimes with 

long rooting base, 3-15 cm total lenath 

X 0.8-3 cm diam, blackish, roughened 

with wrinkles and verrucae, with incon- 

Spicuous to discoid ostioles. Asco- 

spores (22-)23-26(-28) X 7.5-8(-9) um, 

with short straight to oblique germ slit. ..... 
a AMP aU HOG nA Ny Pe hd Men Och Gu Nes X. polymorpha 
NOTE: A form of this species with long rooting base 
has been called X. ophiopoda. 

Fertile part cylindrical, clavate, or 

irregular, on short or long tomentose 

Stipe, 1.5-6 cm total length X 2-6 mm 

diam, blackish, sometimes with brown 


bey 


th 


289 


scales, with inconspicuous to hemi- 
Spherical ostioles. Ascospores 18-71 

-22) X 6-7 wm, with oblique to spiral- 
HAT CEGELM ESL ON oti seheet hott et tose mn . X. scruposa 
NOTE: Some collections which might represent tthis 
Species have prominent whitish hemispherical 
ostioles and ascospores (18-)19-25 X 6-8 um with 
abruptly pinched ends and spiralling germ slits. 


18. Fertile part subglobose to oblong- 
ellipsoidal, on short stipe, 1.5- 
2.5 cm total length X 0.5-1.6 cm diam, 
blackish, roughened by wrinkles and 
hemispherical ostioles. Ascospores 
21-26 X (6-)6.5-8 um, with short 
ODA OUCHOe MINIS Tt ie rei? ne . X. schweinitzii 
18. Fertile part subqglobose to elong- 
ated, sessile or short stipitate, 
0.3-5 cm total length X 0.1-1.2 cm 
diam, blackish, wrinkled and verru- 
cose, with inconspicuous to hemi- 
spheric ostioles. Ascospores 22-30 
(-31) X 7.5-9 ym, with short oblique to 
SPIN G WGermes Milman aaa SOpleura 


Fertile part cylindric-clavate to 

irregular with acute sterile apices, 

unbranched or branched, on short stipe, 

solitary or cespitose, 1.3-4 cm total 

length X 3-6 mm diam, exterior with 

brown scales, interior cream-yel low, 

with obscure to hemispherical ostioles. 

Ascospores (12-)12.5-14.5(-16) X 5-5.5 

(-6) um, with straight germ slit less 

Pienmes COCs Leno cos, Mons NONOG Ot amet tue e awit ewes 
Eo ' hot Maen Dhol epics Xylaria sp. (no. 303) 
Combination of features differing 

EOP avOVE.. a CEONdatanOnealCOtyWOOG itm ke Ru we 2 


20. Fertile part cylindrical to 
irregular, usually branched, on 
distinct stipe, sometimes with 
discoid pannose base, solitary, 
2.5-5.5 cm total length X 2-6 mm 
diam, brown to dark brown, strongly 
roughened with papillate ostioles. 
Ascospores 9-10(-11) X (3.5-)4 um, 


290 


MA 


PA he 


23% 


oo. 


with hyaline cellular appendage at 


ONERENG. a apes ome X. uniapiculata sp. nov. 
20. Combination of features differing 
FrompaboVes, Si lips. Pi a iol ee ee fe! 


Fertile part cylindrical, the uppermost 
portions often flattened, unbranched or 
branched, solitary, on short to long 
Stipe, 7-10 cm total length X 3-12 mm 
diam, reddish brown to dull black, 
Ornamented by reticulating cracks, with 
small papillae often surrounded by 
sunken discs. Ascospores (9.5-)10.5-13 
(-14) X 4-5(-5.5) um, with straight germ 


slit slightly less-than spore-length 2 0 5 .seeeeee 
SUleGANAN es oicocise: Vebl-calie cotiied Veblie bran cum Ee X. adscendens 
Combination of characters differing 

From, ADOVE .wes ces footed vs eudiote whe sinks vocals «oc en Ra 


ee. Fertile part clavate, about 3 
total length, on long stipe 
Originating from swollen base, 
unbranched or branched, 7-8 cm 
total length X 5-7 mm diam, 
blackish, with gray peeling layer, 
with whitish deposits surrounding 
papillate ostioles. Ascospores 
(8-)8.5-9(-10) X 4-5 um, nearly 
black, with inconspicuous germ slith.) eee 
wih coi tie 6) Coe ctig ones ERC wae Cai X. claviceps sp. nov. 
22.. Stromata;with short stipess =>. .»)-Ciewineeeeee 
Fertile part clavate, spathulate, 
or irregular, branched or unbranched, 
from pannose base, 2.5-6.5 cm total 
length X 5-10 mm diam, black, rugulose, 
with finely papillate to discoid 
ostioles. Ascospores (8-)8.5-11(-12) 
X 4-4,.5(-5) um, with germ slit con- 
SPI CUOUSSONEUNCONS Dil CUOUSE enrms meus ie X. feejeensis 
NOTE: Fertile part sometimes hollow, with ostioles 
covered by a white material. 
Fertile part clavate to cylindrical, 
unbranched or branched, 1.5-3.5 cm 
total length X 2.5-5 mm diam, blackish 
with whitish to yellowish scales, 
rugulose, with obscure to finely papill- 
ate ostioles. Ascospores (9.5-)10-11 


291 


X 4-4.5 um, with straight germ slit 
sometimes slightly less than spore- 
LCUGLIGM Te Men ae eee em sae ar, SecA cue nCuntd 


SPECIES WITH SMALL STROMATA ON WOOD OR SOT: 
(Stromata usually less than 5 mm diameter) 


ihe Stromatal surface more or less smooth ....... 2 
1. Stromatal surface more or less rough- 
ened with perithecial contours, coarse 
ostiolar papillae, wrinkles, or combina- 
ELONSs*OTeUNCSeml CaLuUVeS mere hat ener ee Lee, 9 


7S Fertile part fusoid or short cylin- 
drical with acute sterile apices, 
with inconspicuous to conspicuous 
perithecial contours, on very long 
filiform tomentose stipe ca. 3/4 
total length, solitary or branched, 
7.5-7.8 cm total length X 1-2 mm 
diam, black with dark brown peeling 
outer layer, with inconspicuous to 
more or less papillate ostioles. 
ASCOSPOKeSal | =—4 =O kmo=OeuM omens: 6) 6 5 


ee Fertile part various. Stipe 
SNC CHRONO. URI ODN emeseoe rete a en , fee ot 3 


She Fertile part subglobose to elongated, 
up to 6 mm long X 4 mm diam, on narrow 
Stipe 1-1.5 cm long, solitary, blackish 
gray with whitish scales, with incon- 
spicuous ostioles. Ascospores (29-)31-37 
(-40) X (11-)12-13(-14) um, with hyaline 


SERA Str ee eet 2 rae yon X. squamulosa sp. nov. 
3. Fertile part clavate, cylindrical, 

COUMeCINANO rae] GREQU d Gircencertcte ste rte’ Cin smeeirctn quem ee 4. 4 

fee LCOMd sia Ot =COLOVCOROCEWNAGCM iS Peak. 4. O. 5 

Ae ULOMaABRDrOWN acon darksbrownnorablackma. . .'.. .. 6 


5. Fertile part cylindrical on short or 
long stipe 0.5-3.5 cm total length X 


* Ascospore germ slits may be assumed to be straight and 
oriented with the long axis of the spore and run essen- 

tially the length of the spore unless noted otherwise in 
the key. 


Fae 


1-3 mm diam, with white granular material 

over much of clava, with black umbilicate 

or slightly raised ostioles. Ascospores 

6-9(-10) X 3-3.5(-4) um, with inconspicu- 

OUSGenmas | TiGeiwh. i Se hee LL oie ze X. “muscula" 
Fertile part clavate, conical to sub- 
globose on long stipe, 1.5-3.5 cm total 
length X 3-5 mm diam, at first white 
becoming cinereus, very smooth and hard, 
with discoid ostioles. Ascospores 
(8.5-)9-10(-11) X 3.5-4.5 um, with in- 
CONSPTCUOUS* GERM «SCs. eee ee eect ae X. pallida 


6. Fertile part cylindrical on dis- 

tinct stipe, with acute sterile 

apices, slightly to strongly pube- 

scent overall, unbranched or 

branched, 3 cm total! length X 1-2 

mm diam, black with brown peeling 

outer layer, with umbilicate to 

slightly papillate ostioles. 

Ascospores 11-12(-12.5) X (4-)5- 

5.5 um, with germ slit less than 

Sspore-Tength sw. ae. tetas coh X. maitlandii 
Gh. eobertile: pant g)aDVOuS. tie Genes es 7 


Fertile part cylindrical with more or 

less flattened sterile apices, on short 

stipe, 1-2.5 cm total length X 2-4 mm 

diam, with reddish brown to dark brown 

peeling outer layer, with siightly 

raised ostioles in small disc. Asco- 

spores (9-)9.5-11(-12):X 4{-4.5) ums ° 2) eee 

RR i hh cee Fodih aa aid en Xylaria sp.(nos. 39,306) 
NOTE: This species also is found on soil, probably 

associated with insect nests. 

Combination of characters differing 

from‘aboves 0s std ee Os ee 8 


8. Fertile part cylindrical to long- 
conical on obsolete or short to 
long glabrous or pubescent stipe, 
0.6-3.5 cm total length X 1-4 mm 
diam, blackish with gray to brown 
peeling outer layer splitting in 
bands, with umbilicate to slightly 
papillate ostioles. Ascospores 
(12-)13-18(-19) X (4-)4.5-6 um, 


ike 


1a 


293 


with straight to undulate germ 
Sielua Le NCOs AeSPONe=lENGchm mae cake mee 
Be av as rs es X. arbuscula 
NOTE: Some collections probably referrable to this 
Species are more robust and have strigose stipes. 
. Fertile part cylindrical to clavate, 
with more or less acute sterile 
apices, on short to long pubescent 
stipe, unbranched or branched, 
solitary to cespitose, 1-5.5 cm 
total length X 1-2 mm diam, black- 
ish with brown peeling outer layer, 
undulate from perithecial contours, 
with more or less punctate ostioles. 
Ascospores 9-11(12-) X (3.5-)4-4.5 
CALSD WATT Mee RT SSG AIAG cts a ios > a X. multiplex 


Stromata growing on soil (or wood, 
nese smsCedSUeLG WODUGICG NN. SOMMpDine ee mers aide pe) 2) 10 
PaO VamOTOWINOUON AWOOU carats ue) deh depleuneucpoti ted beite it's L2 


10. Fertile part of more or less naked 
perithecia on short or long filiform 
tomentose rachis, with long or short 
sterile apex, 3-12 cm tota! length X 
1-3 mm diam, blackish, with papillate 
ostioles. Ascospores 10-12(-13) X 
4-5 um, with straight germ slit less 


than spore-length. .... : - X. gracillima 
10. Fertile part bearing more or less 
AMMEN SEU DEULENCO Ld cube ubem fabs aieh uch lediouvs. voter ie at 


Fertile part cylindrical with sterile 

apex, on tomentose stipe, unbranched, 

solitary, 3-5.5 cm total length X 1-2 

mm diam, blackish with gray peeling 

outer layer, with conical ostioles. 

Ascospores with flattened hyaline cell- 

ular appendage at each end, 8-10 X 3-4 

um (including appendages) . ... X. alata sp. nov. 
Fertile part cylindrical with sterile 
apices, branched or unbranched, on 
short to long tomentose stipe, 4 cm 
total length X 1.5-3 mm diam, black- 
ish with brownish to gray outer layer, 
with prominent conical ostioles. Ascus 
with massive apical ring. Ascospores 
34-45(-47) X (14-)15-20(-21) um, with 


294 


Sk, 


Tor 


15. 


Loe 


hyaline noncellular appendages and 


hyaline sheaths 2 2 2). ) Xv magniannulata speenove 


12. Stromata growing on monocot wood. . 
12. Stromata growing on dicot wood... 


Fertile part more or less clavate, 
on short stipe, 1 cm total length X 
1.5-3 mm diam, reddish brown, very 
rough, with hemispherical ostioles. 
Ascospores 10-12(-14) X 3.5-4.5(-5) 
um, with germ slit less than spore- 


length? Mveee ly eeiice : . ov Xeonscabriclavula Ssounove 


Fertile part cylindrical, with more 

or less crowded naked perithecia, 

with acute sterile apex, on more or 

less short stipe from swollen base, 

1.5-4 cm total length X 1-2 mm diam, 
blackish with brown peeling outer 

layer, with small papillate ostioles, 
sometimes within a depressed annulus. 
Ascospores (10.5-)11-12 X 3-4 um, with 
germ slit obscure or perhaps absent... 


Nig engl bates Sos Mite se Xylaria sp., aff. bambooensis 


14. Fertile part bearing more or 
less immersed to conspicuous 
perithecia, the prominence of 
perithecial contours often vary- 
ing on different parts of the 
SAMGuSUCOMG tebe eR ane ae sullen concurs 
14. Fertile part bearing more or 
less naked individual perithecia 


or clusters of naked perithecia. ... 


Fertile part cylindric-compressed bear- 
ing immersed to almost naked perithecia, 
with acute sterile apex, unbranched or 
branched, on more or less short stipe, 
up to 7 cm total length X 2-4 mm dian, 
dull blackish, tomentose over all, with 
finely papillate ostioles. Ascospores 
10-13(-14) X (2.5-)3-4 um, subhyaline, 


without evident germ slit ..... LET 


Fert espa dD GOUSsi aries err 


16. Fertile part cylindrical to spathu- 
late with acute sterile apex on 
long, pubescent stipe, solitary 


dichotoma 


Wes 


ee 


ioe 


Lor, 


295 


or cespitose, usually branched, 

2.5-3 cm total length X 1-6 mm 

diam, blackish with white peeling 

outer layer, with conical to hemi- 

spherical ostioles. Ascospores 

(10-)11-13(-14) X 4-5(-6) um, with 

SDL ROCriNGS tLe tier eee or emanate mere tle feT gi 

rib Pcs aed Xylaria sp., aff. arbuscula (no. 202) 
16. Fertile part without conspicuous 

peeling outer layer or, if present, 

dark DGownnOyep Tacks eeee eeu oma erar et 17 


Fertile part cylindrical, with sterile 

acute apex, on ill-defined stipe, 

solitary or cespitose, 2-6 cm total 

length X 2-4 mm diam, black although 

apex often with white remains of 

anamorph, with somewhat raised ostioles. 

Ascospores (7.3-)9-10(-11) X 4-5(-5.5) um... .. 
. ol 3 RS OA cee oer eta tee X. cf. longiana 
Combination of features differing from above. . . 


18. Fertile part cylindrical to 
clavate with blunt to acute 
Sterile apex, on more or less 
long stipe, unbranched or 
branched, 1-5 cm total length 
X 1-3 mm diam, biackish with 
lighter brown peeling outer layer, 
with conspicuous crowded hemi- 
Spherical ostioles. Ascospores 
8-9(-10) X 4.5-5 um, some with 
hyaline cellular appendage on 


Onesend’ ohm eee 9, oe Xylaria sp.(nos. 38, 64) 
18. Combination of features differ- 
INGehrOMLADOVG mee ek Meher bro sai my aes 19 


Fertile part cylindrical, on short 

Stipe, 7-10 mm total length X 1-3 mm 

diam, blackish with brown peeling outer 

layer, with circumferential ly-oriented 

wrinkles, with white fringed, strongly 

conical ostioles. Ascospores (11-) 

13-14(-15) X 4-5 um, with germ slit 

ODSCULCE SE cern cen here, Xylaria sp. (no. 359) 
Combination of features differing 

ErOMZADOVCran wear en a tan monly Senn oe Eee ais eo Ll 20 


296 


alts 
rash 


reste 


Cte 


20. Fertile part cylindrical to clavate, 
with sterile apex, on short stipe 
originating from bulbous base, ca. 

2 cm total length X 2-3 mm diam, 
blackish, with finely papillate 
ostioles. Ascospores 10-11 X 


Se OAS UII Te otas Ghanienite Era ees Cees X. Cf. brevipes 
20. Combination of features differing 

FLOM AD OVE: ieee iehedes: on Ree en ten ies to) ooh coke 21 
Fertile part short-cylindric to clavate ..... 22 
Fertile. part ‘cylindrical to irregular’ 4 7.sen eee 25 


22. Fertile part cylindrical with acute 

Sterile apex, originating from 

tomentose discoid base, solitary, 

1.5-3 cm total length X 1.5-3 

mm diam, black with inconspicuous 

brown to dark brown peeling outer 

layer, with conical to hemispherical 

ostioles. Ascospores (8.5-)9-11 X 

(3-)3.5-4(-4.5) um, with straight 

germ slit less than spore-length. . » Tee es 

A ACE NRA Some be (Pt a a Xylaria sp. (no. 729) 
22. Fertile part cylindrical to cyT- 

indric clavate, on more or less long 

Stipe, 1.5-2 cm total length X 1-2 

mm diam, blackish with brown to 

dark brown peeling outer layer, 

with papillate ostioles. Ascospores 

9-10(-11) X (4-)4.5-5 um, without 

obvious germ slit. . 2’... Xylariaisps (yoemeeen 


Fertile part unbranched or branched, 

on short to long stipe, 1.5-3.5 cm total 

length X 2.5-4 mm diam, blackish with 

peeling outer layer, with hemispherical 

ostioles. Ascospores 9-10(-11) X 4-4.5 

(-5) pm... . Xylaria sp. aff. feejeensis (no. 47) 
Fertile part unbranched or branched, 

roughened with perithecial undulations, 

with attenuated sterile apex, on short 

to long stipe, 1.5-3.5 cm total length 

X 2-3 mm diam, black and lacking outer 

peeling layer, with inconspicuous conical 

ostioles. Ascospores 9~-10.5(-11) X 4-5 um, 

with straight germ slit less than spore- 

TONG Taare Pe eee ma eben Xylaria sp.(no. 331) 


a 


Coe 


1h 


297 


24. Ascospores usually longer than 


"ipsdosee? G1] afb ds Ar MEAD eMC Aer rues | bu A! 0 a a5 
24. Ascospores usually no longer 
Elaine co A UMasere rte ahem ca a tie Tene Ts aS) aoe 26 


Fertile part cylindrical, terete or 
compressed or irregular, with attenuated 
sterile apex, on long stipe, 1.5-3 cm 
total length X 1 mm diam, blackish, with 
grey peeling outer layer, pubescent over 
all, with umbilicate to slightly raised 
ostioles. Ascospores 14.5-17(-18) X 5- 
5.5(-6.5) um, with straight germ slit 
Slightly less than spore-length . ...X. juruensis 
Fertile part cylindrical, with sterile 
apex, on long stipe, unbranched or 
branched, 3-5 cm total length X 2-3 

mm diam, with outer peeling brown layer, 


with umbilicate or finely papillate 


ostioles. Ascospores (12.5-)13-16(-18) 
Kaeo Caries eT ek. lsat ts X. cf. mellisii var. nuda 


26. Fertile part cylindrical with 

Sterile apex, on short to long 

Stipe, unbranched or branched, 1.5- 

6 cm total length X 1-3 mm diam, 

blackish covered with white, yellow- 

ish, or brownish peeling outer layer, 

with slightly raised ostioles. 

Ascospores 10-11(-13) X (3.5-)4-4.5 

um, with straight germ slit less than 

SDOVETILCNO UN ee sais ete) ents X. coccophora 
26. Fertile part cylindrical with long 

acute sterile apex, on short glabrous 

Stipe, unbranched, 1.5-3 cm total 

length X 2 mm diam, blackish with 

white peeling outer layer, with 

Slightly raised ostioles. Asco- 

spores (9-)10-11 X 4-4.5(-5) um, 

with straight germ slit less than 

RUNS bes DOGS SON Likes maen fon r. ye Claimicroceras 


SPEGCTEST ON UEAVES,. PEN COLES. OR FRUITS. 


os Fertile part globose to some- 


what conical. sc. 55.5. cnn ooo: cele recat ae 3 
ae Fertile part cylindrical, clavate, 
OPT NV EGU As 6 lis hcl. See ole Ves ct cee men 4 


Fertile part composed of a head of more 

or less naked perithecia on long rachis 
that becomes the stipe, 3.7-5 cm total 
length X 1-2 mm diam, black, with conical 
ostioles. Ascospores (14-)15-18(-19) xX 
5.5-6 um, with hyaline noncellular append- 
age at each end, with straight germ slit 
slightly less than spore-length. On 


Guazuma ulmifoltia’ fruits ia. 2% i.e 2. 2) eee 


She EO DAA cay ogike ne in ounes X. guazumae sp. nov. 
Fertile part composed of a globose 

to somewhat conical head with nearly 

immersed to evident perithecia on long 

finely pubescent stipe, 1-1.6 cm total 

length X 0.8-2 mm diam, at first white 

becoming grayish, with papillate ostioles. 
Ascospores 9-10(-11) X (3.5-)4-4.5 um, with 

straight germ slit 3/4 spore-length. On 


Guazuma ulmi folia fruits. 00) eerie X. aristata 
4, Ascospores usually longer than 

12th taly) ot SE. vc TOME Pheer 28 Use 5 
4. ~ Ascospores-no ‘longensthan? 12) um.) . 20 2 eee 7 


Fertile part cylindrical to irregular, 
with scattered to crowded, more or 
less naked perithecia, with acute 
sterile apices, on short to long 
tomentose stipe, 4-7 cm total 

length X 1.5-3.5 mm diam, dark 

brown to black, with conical ostioles. 
Ascospores 12-16(-17) X 3-5 um, sub- 
hyaline to yellowish, without evident 
germ slit. On Magnolia schiedeana 


ae hs oe a eRe eae ey X. magnoliae 
Fertile part and ascospores differing 
From above sage a hark a coe. Fob rls Lote) oy eee enn 6 


* Ascospore germ slits may be assumed to be straight and 
oriented with the long axis of the spore and run 
essentially the length of the spore, unless otherwise 
noted in the key. 


299 


6. Fertile part cylindrical with scat- 
tered to crowded, more or less evident 
perithecia, with sterile apices, on 
short to long tomentose stipe, 3.5-7 
cm total length X 0.5-2 mm diam, 
with conical ostioles. Ascospores 
13.5-16 X (4.5-)5-6(-6.5) um, with 
Spiral germ slit. On Liquidambar 
Su. baC Lud: CU 1 CS menrsmeeinss tonne X. persicaria 
6. Fertile part cylindrical to some- — 
what conical, with sterile apices, 
on short or long glabrous or pubescent 
Stipe, 0.3-2.2 cm total length X 1.5- 
2.5 mm, blackish with gray peeling outer 
layer, with slightly raised ostioles. 
Ascospores (12-)13-16 X 4.5-6 um, 
with straight to undulate germ slit 
1/2 to 3/4 spore-length. (On Swietenia 
macrophylla and Quercus spp. fruits ...... 
I BL Ak bt ee a a Aaa X. arbuscula 
NOTE: X. arbuscula is more commonly encount- 
ered on wood. It is also keyed out with the 
smal] wood-inhabiting species. 


Fertile part irregular in shape, com- 
pressed, branched, bearing more or less 
naked perithecia, on long finely pubescent 
stipe, 2 cm total length X 1.5 mm diam, 
blackish, with umbilicate to slightly 
raised ostioles. Ascospores 10-11(-11.5) 
X 4-4.5(-5) ym. On Swietenia macrophylla 


Haun Utee ie unde Ihe “aren ho ueop yi X. cf. inaequalis 
Combination of features differing 
SOIC DONC 2 sts ea ks Ig ti,. cae ve tL CN Ae ee. 8 


8. Fertile part cylindrical to irre- 
gular, bearing more or less naked 
perithecia, with acute sterile apex, 
unbranched or branched, usually on 
ill-defined stipe, 2-13 cm total 
length X 1.5-2.5 mm diam, reddish 
brown to blackish, tomentose overall, 
with finely papillate ostioles. 
Ascospores 10-12(-13) X 4-5 um. 

On fruits of Guazuma ulmifolia and 
various pods, including Pithecellobium 
LICK. Cau lCieeeswesnee) cleeerew X. Tanthino-velutina 


WO WO 


Lies 


LD, 


ae Fertile part cylindrical with 
short sterile apex on long tomentose 
stipe, 3-7 cm total length X 1-2.5 
mm diam, blackish with grayish brown 
peeling outer layer splitting in 
lines, with papillate ostioles. 
Ascospores (9.5-)10-11.5(-12) X 
5-5.5 um. On buried fruits and 


superficial “unidentified pods. \..9.) se ; 

os CTY ec eenbvelh sl ealbe Peel ct Wk auc aaa evans wats X. oxyacantha 
Fertile part igiobose ses 2 Ra oy oe 10 
Fertile. part. more or less cylindrical > .9saeee 11 


10. Fertile part a small aggregation 
of more or less naked perithecia 
on rachis that often extends into 
a sterile apex, on long, glabrous 
stipe, up to 2 cm total length X 
1-2.5 mm diam, black, with barely 
papillate ostioles. Ascospores 
12-14(-15) X (6-)7-8 um, with 
hyaline noncellular appendage at 


CaCh end Seah aati berets X. amphithele sp. 
10. Fertile part globose with im- 
mersed perithecia, with acute 
Sterile apex, on long stipe, up 
to 2.3 cm total length X 1.5 mm 
diam, with stipe and sterile 
apex tomentose, with papillate to 
almost digitate ostioles. Asco- 
spores 10-12(-14) X 5-6 um, with 
hyaline cellular appendage at one 
CNV Ges pee sonal ane eee, “ “Xvlaria Spetnoe 
Fertile part short cylindrical, com- 
posed of aggregations of more or less 
naked perithecia, with acute sterile 
apex, on filiform glabrous stipe, 
0.5-1.3 cm total length X 0.5-1.5 mm 
diam, black, with papillate ostioles. 
Ascospores (5.5-)6-7 X 3-3.5(-4) um, 
with straight germ slit slightly less 
than spore-length...... Xylaria sp. (no. 


Fertile part cylindrical, bearing 
immersed perithecia, with short, acute 
Sterile apex, on more or less long 
Stipe, 0.3-3.8 cm total length X 0.5- 


nov. 


337) 


301 


1.5 mm diam, reddish brown, with strong- 

ly papillate ostioles. Ascospores 11-14 

X 5-6(-7) um, with hyaline sheath, with 

straight germ slit slightly less than 

SUUGCHM CHOC works een eins iromber Terug” sauic ie X. phyllocharis 


Xylaria adscendens (Fr.) Fr., Nova Acta Regiae Soc. Sci. 
Hosauimisencis) supe io. 1351. BAGS sihwes 


Stromata unbranched or branched from the base or 
above, cylindrical, terete or flattened particularly in 
apical parts, with apices sterile, on short or long 
stipes, 7-10 cm total length X 3-12 mm diam; externally 
reddish brown to dull black; internally whitish, sometimes 
becoming hollow. Texture hard. Surface smooth, orna- 
mented with reticulating cracks. Perithecia 0.4-0.8 mm 
diam. Ostioles papillate, often located in sunken 
circular areas 0.25-0.35 mm diam. Asci eight-spored, 
cylindrical, long-stipitate, 230-255 wm total length X 
6-6.5 um broad, the spore-bearing part 72-84 um, with 
apical ring bluing in Melzer's iodine reagent, quadrate, 
2X 2 um. Ascospores brown, unicellular, ellipsoid- 
inequilateral with broadly or narrowly rounded ends, 
smooth, (9.5-)10.5-13(-14) X 4-5(-5.5) um, with straight 
germ slit slightly less than spore-length. 


SPECIMEN EXAMINED: Garcia 5249, 22.VIII.1986 and San 
Martin 203, 18.VII.1987, "El Cielo" ranch, Gomez Farias 
municipality, Tamaulipas state, on Quercus spp. wood in 
cloud forest. 


NOTES: Our material seems much like that described and 
illustrated by Dennis, 1961; Rogers, 1984A. Ascospores 
and the ascus are depicted in Rogers (1984A). 


Xylaria alata San Martin & Rogers, sp. nov. 
FIGS. p35 aula 2s. 


Stromata cylindracea, aggregatis peritheciis plus 
minusve immersis, apicibus acutis sterilibus, in longis 
abrupte attenuatis tomentosis stipitibus, 3-5.5 cm longa 
X 1-2 mm crassa; extus nigella cum externo strato cinereo 
delapso; intus albida vel flavida. Textura mollis vel 
plus minusve dura. Superficies asperata ab formis 
peritheciorum et papillis ostiolorum. Perithecia 0.2-0.3 
mm diam. OQOstiola conica. Asci saepe biseriate 
octospori, cylindrici, stipitati, 76-110 longitudine tota 


302 


X 4-7 um crassi, partibus sporiferis 44-60 um, annulo 
apicali in liquore Melzeri cyanescente, quadrato, 1.5-2 ym 
X 1.5-2 pm. Asci inter conspicue crassas paraphyses. 
Ascosporae brunneae, ellipsoideo-inequilaterales, omni 
extremo cum cellulari appendicula complanata hyalina ca. 

2 um X 2 um, leves, 8-10 X 3-4 um includentes appendi- 
culas, rima germinative recta per longitudinem partis 
coloris sporae. Anamorphosis ignotus. 


Stromata cylindrical, with crowded partially 
immersed perithecia, with acute sterile apices, on 
abrupt, long thin tomentose stipes, 3-5.5 cm long X 1-2 
mm diam; externally blackish with gray peeling outer 
layer; internally whitish to yellowish. Texture soft to 
fairly hard. Surface roughened with perithecial contours 
and ostiolar papillae. Perithecia 0.2-0.3 mm diam. 
Ostioles conical. Asci eight-spored, the spores arranged 
in a uniseriate or partly biseriate manner, cylindrical, 
Stipitate, 76-110 um total length X 4-7 um broad, the 
Spore-bearing part 44-60 um, with apical ring bluing in 
Melzer's iodine reagent, quadrate, 1.5-2 um X 1.5-2 um. 
Asci among conspicuously broad paraphyses. Ascospores 
brown, ellipsoid-inequilateral with a flattened hyaline 
cellular appendage ca. 2 um X 2 um at each end, smooth, 
8-10 X 3-4 ym including appendages, with straight germ 
Slit running the length of the colored part of the spore. 
Anamorph unknown. 


SPECIMEN EXAMINED: San Martin 389, V1.1987, Gomez 
Farias, Tamaulipas state, on wood or fruits buried in 
soil in median subdeciduous rain forest (HOLOTYPE: ITCV). 


NOTES: This fungus differs from all other Xylarias in 
the wing-like flattened ascospore appendages. It is 
named on the basis of these ascospore "wings." Cellular 
appendages on ascospores of most Xylarias are essentially 
conical, i.e€., appearing approximately the same shape as 
the spore is rotated. The appendages of X. alata appear 
conical in some orientations and plate-like in others, 
7.e€., aS if one were rotating the bit of a spade or 
Spatula. They sometimes cur] upward. The centrum of 


Figs. 1-5." 152. Xylaria adscendens.,, 1. . Stromataran 
0.85. 2. Stromatal surface, X 16. 3,4. Xylaria alata. 
3. Stromatadicsurface. X 16. 94.0) Stromata.,aricneoe 
Xylaria allantoidea. Stromata, X l. 


303 


304 


this fungus is noteworthy in having very broad paraphyses 
for the small size of the perithecium. They are much 
larger than mature asci. 


Xylaria allantoidea (Berk.) Fr., Nova Acta Regiae Soc. 


Sci. Upsalio(ser..3) 1, p..1272 1851) ek igseee ae 


Stromata unbranched or branched at base, clavate to 
clavate-cylindric, with rounded fertile apices, on short 
Stipes, 5-8 cm total length X 1-1.5 cm diam, externally 
copper- to bronze-colored, becoming black; internally 
white to yellowish, becoming hollow. Texture hard. Sur- 
face smooth except for ostiolar papillae. Perithecia 
0.5-1 mm diam. Ostioles finely papillate. Asci eight- 
spored, the spores arranged in a uniseriate or partially 
biseriate manner, cylindrical, 150-180 um total length X 
4-5 um broad, the spore-bearing part 72-88 um long, with 
apical ring bluing in Melzer's iodine reagent, quadrate, 
2 X 2 um. Ascospores light brown to brown, unicellular, 
ellipsoid-inequilateral, smooth, (9-)10-12 X 3.5-4 um, 
with germ slit straight, somewhat less than spore-length. 


SPECIMENS EXAMINED: *Brown, D. 576, 19.VIII.1982, Parque 
Francisco Javier Clavijero, INIREB, near Xalapa-Coatepec, 
Veracruz state, on wood (JDR;XAL); Guevara 730, 31.1.1986, 
between Plan de la Noria and San Isidro, Jose Maria 
Morelos municipality, Quintana Roo state, on wood (ITCV). 


NOTES: Our material seems typical for the species (see 
Dennis, 1956; Rogers, 1984B). 


Xylaria amphithele San Martin & Rogers, sp. nov. 
Figs. 6,113 90065 


Stromata globosa vel conica, constantia ex aggregatis 
3-4 vel 20 peritheciorum nudorum 1-2.5 mm diam, interdum 
Sterilibus appendicibus ornata, in longis attenuatis 
glabris stipitibus usque ad 2 cm longitudine; extus ni- 
gella; intus alba. Textura mollis. Superficies capituli 


Figs. 6-12. 6. Xylaria amphithele. Stroma, X 10. 7,8. 
Xylaria anisopleura. 7. Stromata, X 3. 8. Stromatal 
Surface, X 16. 9. Xylaria allantoidea. Stromatal 
surface, X 16. 10-12. XyTaria arbuscula. 10. 
Stromata,/X1.5. 11. Stromatalisurtacemiir.o 2 oles 
Stromata, X 1.25. 


305 


306 


asperata a formis peritheciorum. Perithecia 0.3-0.5 mm 
diam. OQstiola plus minusve papillata. Asci octospori, 
cylindrici, stipitati, 129-175 um longitudine tota X 
9.5-11 um crassi, partibus sporiferis 79-102 um, annulo 
apicali in liquore Melzeri cyanescente, inverse petasi- 
formis, 3.5-4.5 um alto X 2.5-3 um crasso. Ascosporae 
brunneae, unicellulares, ellipsoideo-inaequilaterales 

cum rotundatis extremis, omni extremo cum cylindracea vel 
globosa hyalina noncellulari appendicula 2-5 um longa X 5 
um crassa, leves, parte colorata 12-14(-15) X (6-)7-8 um, 
rima germinativa recta per longitudinem sporae. Status 
anamorphosis ignotus. 


Stromata globoid to conical, composed of aggregations 
of 3-4 to 20 naked perithecia 1-2.5 mm diam, sometimes 
with sterile apiculi, on long thin glabrous stipes up to 
2 cm long; externally dull black; internally white. 
Texture soft. Surface of head roughened with perithecial 
contours. Perithecia 0.3-0.5 mm diam. Ostioles barely 
papillate. Asci eight-spored, cylindrical, stipitate, 
129-175 wm total length X 9.5-11 um broad, the spore- 
bearing part 79-102 um, the apical ring bluing in 
Melzer's iodine reagent, inverted hat-shaped, 3.5-4.5 um 
high X 2.5-3 um broad. Ascospores dark brown, unicel lu- 
lar, ellipsoid-inequilateral with rounded ends each 
bearing a cylindrical to globular hyaline noncellular 
(secondary) appendage 2-5 wm long X 2-5 ym broad, smooth, 
the colored part 12-14(-15) X (6-)7-8 um, with straight 
germ slit full-length. Anamorph unknown. 


SPECIMEN EXAMINED: San Martin 207, VII.1987, Gomez 
Farias, Tamaulipas state, on dried fallen leaves in 
median subdeciduous rain forest (HOLOTYPE: ITCV; 
ISOTYPE: JDR). 


NOTES: This curious little Xylaria is unlike any 
species known to us. It is named on the basis of the 
ascospore appendages, i.e., the two nipples. 


Figs... 13-20. 13,14. -Xylaria sp. aff. arbusculage im 
Stromata, X 1.5. 14. Stromatal surface, X 16. 15,16. 
Xylaria sp. (SM 337). 15. Fertile head, X 14. 16. 
Stromata, X 2.25. 17,18. Xylaria sp. aff. bambooensis. 
17. Stromata, X 2.25. 18. Stromatal surface, X 16. 
19,20. Xylaria cf. brevipes. 19. Stromatal surface, X 
162" 3200 mes tromata Ame Ce 


307 


308 


Xylaria anisopleura (Mont.) Fr., Nova Acta Regiae Soc. 
Sci. Upsat. (ser. 3)0l\-p. 127. 185).. Si gseeeeees 
X. globosa (Spreng.: Fr.) Mont., Ann. Sci. Nat. Bot. 
Tsér. 4) 3:103. 1855, fide Laessde in litt. 


Stromata unbranched or branched, subglobose to 
elongated or irregular, often moriform, sessile or 
stipitate, 0.3-5 cm total length X 0.1-1.2 cm diam; 
externally dull black; internally white to yellowish. 
Texture cheesy to woody. Surface usually roughened with 
wrinkles, verrucae, ostiolar papillae and, occasionally, 
tomentum. Perithecia 0.3-1 mm diam. Ostioles obscure to 
hemispherical. Asci eight-spored, stipitate, 230-275 um 
total length X 8-10 um broad, the spore-bearing part 
158-178 um, with apical ring bluing in Melzer's iodine 
reagent, more or less urn-shaped, 7-8 um high X 4-5 um 
broad. Ascospores brown to dark brown, unicellular, 
ellipsoid- inequilateral to navicular, smooth, 22-30(-31) 
X 7.5-9 um, with spiralling or oblique germ slit less 
than spore-length. 


SPECIMENS EXAMINED: Chacon 363, X1.1987, Ejido Julilo, 
Gomez Farias municipality, Tamaulipas state, on wood in 
subtropical forest with abundant Quercus spp. (ITCV;JDR); 
San Martin 35, 14.X.1986, Gomez Farias Retain 
Tamaulipas state, on wood in cloud forest (ITCV); San 
Martin 98, 8.X1I.1986, Ejido La Union, Othon P. Blanco 
municipality, Quintana Roo, on wood in evergreen rain 
forest (ITCV3JDR); San Martin 3245: 336, 399, Vilbeioee 
Ejido Boca de Chajul, Ocosingo municipality, Chiapas 
state, on wood in evergreen forest (324, 336 in ITCV;JDR; 
399 in ITCV); Tinoco 27, ViI.1987 ) "El Cielo! traneae 

Gomez Farias municipality, on wood in cloud forest 
(ITCV;JDR). 


NOTES: Our material mostly fits the common and wide- 
spread concept of X.anisopleura (see Dennis, 1956; 
Péerez-Silva, 1975; Rogers et al., 1988). One collection 


Figs.. 21-28. 21,22. -Xylaria claviceps. 21. Stromate 
X 1. 22. Stromatal surface, X 16. 23-25. Xylaria 
coccophora. 23. Stromatal surface, X 16. ?4. 
Stromatay Xue... 29... sStromata, x 2.5. ,26.27...0c Vee 
aff. comosas 260 -Stromata, X 3ie2a7... Stromata 
surface, X 8. 28. Xylaria cf. corniculata. Stromatal 
surface, X 16. 


309 


310 


(San Martin 399) is a curious form with small stromata 
and sterile acute apices. Laessge (in litt.) has 
reinstated X. globosa that, in his opinion, is an earlier 
name for what is universally called X. anisopleura. 


Xylaria arbuscula Sacc., Michelia 1:249. 1878. 
Figs. 10-12. 


Stromata unbranched or branched from the base or 
above, cylindrical to conical, with acute sterile apices, 
nearly sessile or on short to long, nearly glabrous to 
pubescent stipes, 0.3-3.5 cm total length X 1-4 mm diam; 
externally blackish with gray to brown peeling outer 
layer; internally white to yellowish. Texture fairly 
hard. Surface smooth to slightly wrinkled, stipes often 
with pubescence. Perithecia 0.2-0.6 mm diam. Ostioles 
essentially umbilicate to slightly papillate. Asci 
eight-spored, cylindrical, stipitate, 130-206 um total 
length X 6.5-8 um broad, the spore-bearing part 60-116 
um, with apical ring bluing in Melzer's iodine reagent, 
inverted hat-shaped, 3.5-5.5 um high X 2.5-3 um broad. 
Ascospores brown to dark brown, unicellular, ellipsoid- 
inequilateral, smooth, (12-)13-18(-19) X (4-)4.5-6 um, 
with germ slit straight to undulate, 1/2 to 3/4 spore- 
length to only slightly less than spore-length. 


SPECIMENS EXAMINED: Garcia 5337, 9.X.1986, "El Cielo" 
ranch, Gomez Farias municipality Tamaulipas State, on 
Quercus sp. fruit in cloud forest (ITCV); Garza 1, 
VI.1981, "El Cerrito", Monterrey municipality, Nuevo Leon 
state, on wood in submontane scrub (ITCV); Moreno 3, 
14.X.1986, Gomez Farias, Tamualipas state, on wood in 
subdeciduous rain forest (JTCV); Moreno 19, El Cercado, 
Santiago municipality Nuevo Leon state, on wood in 
submontane scrub (ITCV); *Perez-Silva, E. 6053, X1I.1972, 
near Tepoztlan, Morelos state, on Quercus (JDR; SXAL) s San 
Martin 4, VIII.1987, Gomez Farias, Tamaulipas state, on 
wood in median subdeci duous rain forest (ITCV); San 
Martin 14, VIII.1986, Gomez Farias, Tamaulipas state, on 


Figs. 29-34. 29. Xylaria cf. corniculata. Stromatanen 
0.8. 30° Xylaria cf. curta. Stromatal surface, XiGumea en 
Xylaria cf. dichotoma. Stromatal surface, X 16. 32. 
Xylaria. cf. curta. *Stromata, X 2.. 33. Xylariasets 
dichotoma. Stromata, X 1.25. 34. Xylaria aff. 
enterogena. Stromatal surface. 2X, 16 sees 


Sa 


312 


wood in cloud forest (ITCV;UDR); San Martin 314, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 


on Swietenia macrophylla King fruit (ITCV); Silva 9, 
17.VIT.1987 (same date as Garcia 5337, above) (ITCV). 


NOTES: Xylaria arbuscula is a species complex and our 
specimens undoubtedly encompass more than one taxon. 
Culturing would undoubtedly be of great utility in 
elucidating this assemblage, but our material was not in 
culturable condition. Our concept of the species is 
generally that of Miller (1942), Rogers & Samuels (1986), 
and Rogers et al. (1988). Several of the specimens cited 
here differ from typical material, as follow: San Martin 
eel dirfersiin conspicuous perithecial contours; San 
Martin 50 differs in the robust stromata; Garcia 5250 
differs in the white outer peeling layer. 

Xylaria arbuscula usually occurs on wood. Two col- 
lections are from woody fruits and, thus, little patho- 
logical or ecological significance is attached to these 
occurrences, 7.e., the fruits are viewed as akin to wood. 


Xylaria sp., aff. arbuscula (Moreno 202) Figs. 13 3148 


Stromata unbranched or branched, cylindrical to 
spathulate with acute sterile apices, on long pubescent 
Stipes, 2.5-3 cm total length X 1-6 mm diam; externally 
blackish with white outer peeling layer; internally white 
with brownish medulla. Surface slightly rugose and other- 
wise roughened by ostiolar papillae. Perithecia 0.2-0.5 
mm diam. Ostioles conical to hemispherical, conspicuous 
or obscure. Asci eight-spored, the spores arranged in a 
uniseriate or biseriate manner, cylindrical, stipitate, 
110-131 um total length X 7.5-8.5 um broad, the spore- 
bearing part 62-77 um, with apical ring bluing in 
Melzer's iodine reagent, quadrate to rectangular, 2-3 um 
high X 2-3 ym broad. Ascospores dark brown, unicellular, 
ellipsoid-inequilateral with broadly to narrowly rounded 
ends, smooth, (10-)11-13(-14) X 4-5(-6) um, with germ 
slit undulate to spiralling, more or less spore-length. 


Figs. 35-40. 35. Xylaria aff. enterogena. Stromata, X 
0.8. 36. Xylaria feejeensis. Stromata, X 1. 37,38. 
Xylaria gracillima. 37. Stromata, X 1.5. 38. Stromatal 
surface, X 16. 39. Xylaria feejeensis. Stromatal sur- 
face, X 16. 40. Xylaria grammica. Stromatal surface, X 
16. 


313 


314 


SPECIMEN EXAMINED: Moreno, A. 202, X.1987, El Cercado, 
Santiago municipality, Nuevo Leon state, on wood in scrub 
forest with Quercus spp. (ITCV). 


NOTES: This fungus looks much like a small-spored variant 
of X. arbuscula. The undulate to spiralling germ slits, 
however, are unlike that species. A similar fungus from 
Venezuela (Rogers et al., 1988) described as perhaps an 
atypical X. arbuscula probably represents the same taxon. 
In any case, the X. arbuscula complex is, as yet, poorly 
understood and it seems premature to formally erect 
another taxon until culturing has been accomplished (see 
notes on X. arbuscula). 


Xylaria aristata Mont., Ann. Sci. Nat. Bot. (ser. 4) 
2106. 1855. 


Stromata globose to somewhat conical with acute 
apices, up to 3 mm high X 2 mm diam, on long, thin finely 
pubescent stipes up to 1.3 cm high; externally white, 
becoming gray, stipes dark brown; internally white. 
Texture becoming fairly hard. Surface roughened with 
perithecial contours and ostiolar papillae. Perithecia 
0.3-0.5 mm diam. Ostioles papillate. Asci eight-spored, 
cylindrical, 100-115 um total length X 6-7.5 um broad, 
the spore-bearing part 33-45 um, with apical ring bluing 
in Melzer's iodine reagent, quadrate, 2 X 2 um. Ascospores 
brown, unicellular, ellipsoid-inequilateral with broadly 
or narrowly rounded ends, smooth, 9-10(-11) X (3.5-)4-4.5 
um, with straight germ slit ca. 3/4 spore-length. 


SPECIMEN EXAMINED: San Martin 213, 2.VIII.1987, Ocampo, 
Tamaulipas state, on fallen fruits of Guazuma ulmifolia 
Lam., aquiche in median subdeciduous rain forest (ITCV). 


NOTES: Our material is much as illustrated by Dennis 
(1956) except that perithecial contours of our material 
are more evident and the color is lighter. This species 
usually occurs on leaves; ours is the first report known 


Figs. 41-47. 41. Xylaria grammica. Stromata, X 1. 
2,43. Xylaria guyanensis. 42. Stromata, X 1. 43. 
Stromatal surface, X 16. 44. Xylaria kegeliana. Stroma, 
X 1.5. 45,46. Xylaria juruensis. 945. ,Stromataseneemee 
46. Stromatal surface, X 16. 47. Xylaria kegeliana. 
Stromatal surface, X 16. 


315 


316 


to us of its occurrence on fruits. See also description 
and notes on Xylaria sp. (SM 337) herein. 


Xylaria sp., aff. bambooensis Lloyd, Mycol. Notes 72, 
Mycol. Writ. 7:1285. 1924. Figs. l/l gene 


Stromata unbranched or branched, upright or pros- 
trate, cylindrical with crowded conspicuous to more or 
less naked perithecia, with long acute sterile apices, on 
short stipes arising from swollen bases, 1.5-4 cm total 
length X 1-2 mm diam; externally blackish with evidence 
of brown outer peeling layer; internally white. Texture 
soft. Surface roughened from perithecia and tomentum, 
the stipes and lower fertile parts particularly tomentose. 
Perithecia 0.2-0.3 mm diam. Ostioles finely papillate, 
the papillae sometimes surrounded by a depressed annulus. 
Asci eight-spored, arranged in uniseriate or biseriate 
manner, short-stipitate, 76-94 ym total length X 5.5-7.5 
um broad, the spore-bearing part 56-63 um, with apical 
ring bluing in Melzer's iodine reagent, quadrate, 2 X 2 
um. Ascospores light brown, unicellular, ellipsoid- 
inequilateral to allantoid, smooth, (10.5-)11-12 X 3-4 
um, with germ slit obscure. 


SPECIMEN EXAMINED: Guevara 731, 1986, between Plan de la 
Noria and San Isidro, José Maria Morelos municipality, 
Quintana Roo state, on monocot wood in an acahual (ITCV; 
JOR). 


NOTES: This fungus greatly resembles Lloyd's photograph 
of X. bambooensis (Lloyd, 1924B). The ascospores of our 
material are much smaller than reported by Lloyd, i.e., 
20 X 6 um. They are morphologically much like the asco- 
Spores described for X. magnoliae herein anc another 
taxon described elsewhere as Xylaria sp. aff. magnoliae 
(Rogers et al., 1988). The stromata likewise resemble X. 
magnoliae and X. ianthino-velutina, but the known host _ 
ranges of these fungi do not include monocots. Thus, it 


Figs. 48-54, 48. Xylaria cf. laevis. Stromata, X 1.3. 
49-50. Xylaria longipes. 49. Stromatal surface, X 16. 
50. Stromatay X 10555 51. Xylaria longipes varcmtegeeees 
Stromata, X 1.25. 52. Xylaria sp. aff. feejeensis. 
Stroma, X 3.25. 53,54. Xylaria guazumae. 53. Stromata 
On Evil tee OOM so oe werent nempalrt samo. 


OLY 


318 


is inappropriate at this time to assign this fungus to a 
known species or to erect a new one. 


Xylaria brachiata Sacc., Ann. Mycol. 4:75. 1906. 


Stromata unbranched or branched, upright or 2 
prostrate, the fertile part short cylindrical to fusoid 
with perithecia immersed or with evident perithecial 
contours, with acute sterile apices, 1.5-1.8 cm long X 
1-2 mm diam, on long thin tomentose stipes 6 cm or 
longer; externally black with dark brown outer peeling 
layer; internally white. Texture fairly hard. Surface 
smooth to roughened with perithecial contours, ostiolar 
papillae, and peeling outer layer. Perithecia 0.3-0.5 mm 
diam. Ostioles inconspicuous to papillate. Asci 
eight-spored, cylindrical, 130-160 um total length X 7-8 
um broad, the spore-bearing part 85-92 um, with apical 
ring bluing in Melzer's iodine reagent, rectangular, 
3.5-4 um high X 1.5-2.5 wm high. Ascospores brown, 
unicellular, ellipsoid-inequilateral with broadly rounded 
ends, smooth, 11-14(-16) X 5-6 um, with straight germ 
slit spore-length. 


SPECIMEN EXAMINED: Chacon 102, X.1987, Ejido Alta Cima, 
Gomez Farias municipality, Tamaulipas state, on wood in 
cloud forest (ITCV). 


NOTES: Dennis (1958) placed X. brachiata into synonymy 
with X. mellisii (Berk.) Cooke [as Xylosphaera mellisii 
(Berk.) Dennis]. Theissen (1909) likewise considered it 
close to X. arbuscula Sacc. (which Dennis also placed 
into synonymy with X. mellisii). Lloyd (1919) accepted 
X. brachiata and his photograph (Fig. 1561) looks very 
much Tike our material. Later, Lloyd (1924A) wrote that 
X. brachiata appeared to be too close to X. apiculata to 
be maintained as a separate species. Rogers et al. 
(1988) tentatively identified a collection growing on 
leaves from Venezuela as X. brachiata. The present 


Figs. 55-61. 55,56. Xylaria magniannulata. | S5 eee cama 
X' 2. 56. Stromatal.surface, X 12, 57,58. Xylaia 
magnoliae. 57. Stromatal surface, X 16. 58. Stromata 
on Magnolia fruit, X 1. 59,60. Xylaria maitlandii. 59. 
Stromata, X 2.3. 60. Surface of fertile part and upper 
stipe (left) and lower stipe (right), X 16. 61. Xylaria 
cf. mellisii var. nuda. Stromata, X 1.66. 


319 


320 


material, like Saccardo's type, is from wood. Cultural 
and other studies might clarify the status of X. 
brachiata. 


Xylaria cf. brevipes Sacc., Jour. Mycol. 12:47. 1906. 
Figs. °i9eeee 


Stromata cylindrical to clavate with acute sterile 
apices, on short tomentose stipes arising from bulbous 
bases, ca. 2 cm total length X 2-3 mm diam; externally 
blackish; internally whitish. Texture woody. Surface 
rugose and roughened by cracking and verrucae. Peri- 
thecia 0.3-0.5 mm diam. Ostioles finely papillate. Asci 
eight-spored, long-stipitate, 130-155 um total length X 
4-5 um broad, the spore-bearing part 75-85 um, with 
apical ring bluing in Melzer's iodine reagent, quadrate, 
2X 2 yum. Ascospores brown, unicellular, ellipsoid- 
inequilateral with broadly or narrowly rounded ends, 
smooth, 10-11 X 3.5-4 um, with straight germ slit 
spore-length. 


SPECIMEN EXAMINED: Acosta 8, 8.VI.1984, Raices, 
Monterrey municipality, Nuevo Leon state, on wood in 
submontane scrub (ITCV;JDR). 


NOTES: This is another small Xylaria that seems close to 
X. feejeensis. It resembles Lloyd's description and 
photograph (1924A) of X. brevipes. We have not seen type 
material of X. brevipes. 


Xylaria claviceps San Martin & Rogers, sp. nov. 
Figs. 21,22 ,3t= ease 


Stromata non ramosa vel ramosa, regulatim clavata 
apicibus rotundatis fertilibus, in stipitibus longitudine 
3/4 stromatis aequantibus gestis, e basibus conspicuis 
tumidis pannosis exorientibus, 7-8 cm longa X 5-7 mm 


Figs. 62-71. 62,63. Xylaria cf. microceras: » 629 
Stromata, X 2.3. 63. Stromatal surface, X 16. 64,65. 
Xylaria multiplex. 64. Stromatal surface, X 16. 65. 
Stromata X 3. 66,67. Xylaria "muscula". 66. Stroma X 
2.8. 67. Stromatal. surface, X 16.) 68.69. Xylariawem 
nigrescens. 68. Stromata, X 1.3. 69. Stromatal sur- 
face, X 16. 70,71. Xylaria oxyacanthae. 70. Stromata, 
Xi1 535 aye “Stromatailm@surntacewaur i l6% 


321 


322 


crassa; extus atra cum strato exteriore cinereo delapso, 
ostiolis ab albida materia circumcintis; intus alba. 
Textura similis casio vel lignosa. Superficies asperata 
ab parvis rugis, strato delapso, et papillis ostiolorum. 
Perithecia 0.3-0.5 mm diam. Ostiola papillata. Asci 
ostospori, cylindrici, longe stipitati, 92-122 um longi- 
tudine tota X 5.5-6 um crassi, partibus sporiferis 46-62 
um, annulo apicali in liquore Melzeri cyanescente, 
quadrato, 2 X 2 um. Ascosporae fuliginosae, unicellu- 
lares, late ellipsoideo-inequilaterales, leves, (8-)8.5-9 
(-10) X 4-5 um, rima germinativa inconspicua recta plus 
minusve longitudine sporae. Status anamorphosus ignotis. 


Stromata unbranched or branched, regularly clavate 
with rounded fertile apices, on long stipes representing 
ca. 3/4 of entire length, originating from conspicuous 
swollen pannose bases, 7-8 cm long X 5-7 mm diam; exter- 
nally dull black with gray outer shredding or flaking 
layer, the ostiolar areas surrounded by whitish material; 
internally white. Texture cheesy to woody. Surface 
roughened by small wrinkles, peeling layer, and ostiolar 
papillae. Perithecia 0.3-0.5 mm diam. Ostioles papill- 
ate. Asci eight-spored, cylindrical, long-stipitate, 
96-122 um total length X 5.5-6 um broad, the spore-bearing 
part 46-62 um, with apical ring bluing in Melzer's iodine 
reagent, quadrate, 2 X 2 wm. Ascospores nearly black, 
unicellular, broadly ellipsoid-inequilateral, smooth, 
(8-)8.5-9(-10) X 4-5 um, with inconspicuous straight germ 
Slit more or less spore-length. 


SPECIMEN EXAMINED: San Martin 356, VIII.1987, "E] 
Muneco", Ejido Loma Bonita, Ocosingo municipality, Chiapas 
state, on wood in evergreen rain forest (HOLOTYPE: ITCV). 


NOTES: This fungus has a great resemblance to X. 
hyperythra (Mont.) Fr. as illustrated by Dennis (1956). 
The surface of our fungus is grayish rather than 
cinnamon-colored and the ascospores are much smaller than 
those reported for X. hyperythra. 


Figs. 72-78. 72. Xylaria pallida. Stromata, X02 eee 

74. Xylaria pal.lide-ostiolata’ 73. Stroma, X’ Og6ceueaes 
Ostioles, X 16. 75,76. XylTaria persicaria. 75. Strom- 
atal ‘surface, X 16. 76. Stromata on fruit, xX) 2.9, eee 
Xylaria phyllocharis.° 77. Stromata;, X.3. 78 0e)Fenmtae 

Dances 


346) 


324 


Xylaria coccophora Mont., Ann. Sci. Nat. Bot. (ser. 3) 
7109. 1855. FIGS Siécneoe 


Stromata unbranched or with 2 or more branches, 
cylindrical bearing more or less conspicuous perithecia, 
with acute sterile apices, on short to long smooth 
stipes, 1.5-6 cm total length X 1-3 mm diam; externally 
black with remains of whitish, yellowish or brownish 
peeling outer layer; internally white. Texture fairly 
soft. Surface nodulose from perithecial contours. 
Perithecia 0.4-0.8 mm diam. Ostioles slightly raised. 
Asci eight-spored, cylindrical, stipitate, 150-163 um 
total length X 5-5.5 um broad, the spore-bearing part 
80-88 um, with apical ring bluing in Melzer's iodine 
reagent, quadrate, 2.5-3 X 2.5-3 um. Ascospores brown, 
unicellular, ellipsoid-inequilateral with rounded ends, 
smooth, 10-11(-13) X (3.5-)4-4.5 um, with straight germ 
slit less than spore-length. 


SPEICMENS EXAMINED: Chacon-Jimenez 192, 1.X1.1987, Ejido 
Alta Cima, Gomez Farias municipality, Tamaulipas state, 
on wood in cloud forest (ITCV;UDR); Garcia 5423, 
10.XIJT.1986, Felipe Carrillo Puerto, Quintana Roo state, 
On wood in median subdeciduous rain forest (ITCV;UNR); 

San Martin 101, 8.XII.1986, Ejido La Union, Othon P. 
Blanco municipality, Quintana Roo state, on wood in 
subdeciduous rain forest (ITCV;JDR). 


NOTES: This material fits our concept of X. coccophora 
(see Dennis, 1956; Rogers et al., 1988). Laessge (in 
litt.) indicates that some material from Mexico would be 
better placed in X. compressa Pat. et Gaill. (see also 
Joly, 1968 and Perez-Silva, 1975). 


Xylaria aff. comosa (Mont.).Fr., Summa Veg. Scand. Sect. 
BOoSt- preemie ocos Figs. cosas 


Figs. 79-87. 79. Xylaria polymorpha. "Qphiopoda" type 
Stroma, X 0.5. 80,81. Xylaria scabriclavula. 80. 
Stromata, X 5.75. 81. ‘StromataTl. surfaces xu16 aueoeeee 
Xylaria schweinitzii. 82. Stroma, X 2. 83. Stromatal 
surface, X 16. 84. Xylaria scruposa. Stromata, X 1. 
85,86. Xylaria sp. G 729. 85. Stromata, X 1.3. 86. 
Stromatal surface, X 16. 87. Xylaria scruposa. 
Stromatal surface, X 16. 


325 


326 


Stromata subglobose to pyriform to oblong, up to 1 
cm long X 5 mm diam, on long thin stipes up to 4 cm long 
X 1-1.5 mm diam; externally shiny black with white 
Sloughing patches; internally white and perhaps becoming 
hollow. Texture very hard. Surface smooth except for 
white sloughing patches, ostiolar papillae, remnants of 
conidial processes immediately below fertile part, and 
short hairs on stipe. Perithecia not seen intact. 
Ostioles strongly papilTlate. Asci not seen. Asco- 
spores brown, unicellular, ellipsoid-inequilateral with 
abrupt pinched ends, smooth, (25-)29-33(-36) X 7-8(-9) 
um, with germ slit straight, Slightly less than 
full-length. 


SPECIMEN EXAMINED: San Martin 350, "El Muneco, Fjido 
Loma Bonita, VIII. 1987, Ocosingo municipality, Chiapas 
state, on wood in evergreen rain forest (ITCV). 


NOTES: Our stomata are smaller than those reported by 
Dennis (1956) and Rogers et al. (1988). Material of the 
latter authors have ascospores with spiral germ slits, 
whereas our ascospores have straight germ slits. T. 
Laessde (in litt.) reports that the type of X. comosa is 
sterile. The lectotype selected by him has ascospores 
with straight germ slits, long secondary appendages, and 
according to our data, a cellular appendage on one end. 
Material having ascospores with a spiralling germ slit 
and devoid of appendages [such as described by Rogers et 
al. (1988)] is being described as X. comosoides Laessde. 
Our present material differs from X. comosa sensu Laessde 
in the generally smaller fertile parts and in having 
ascospores without appendages. We thus hesitate to call 
our material X. comosa until additional collections have 
been made and relevant cultures compared. 


Figs. 88-98. 88,89. Xylaria cf. longiana. 88. 
stromata, X 128. 89. “Stromatal surface eX sl6eeeoues 
Xylaria sp. SM 39. Stromatal surface, X 16. 91. 
XyTaria sp. SM 64. Stromatal surface, X 16. 92. 
Xylaria sp. SM 39, Stromata, X 2. 93. Xylaria sp. SM 

. otromata, X 2. 94. Xylaria sp. SM 211. Stroma on 
leaf, X-4.5. 95,96. Xylaria sp. SM 303. 95.  Stromatas 
X 1.75. 96.  Stromatalisurface, X 16. 97,98. Xylaria 
Sp. SM 359. 97. Stromata, X 7.. 98. Stromatal®suppaces 
X16. 


O27, 


328 


Xylaria cf. corniculata Sacc., Ann. Mycol. 4:75. 1906 
FIigS.ecGecee 


Stromata unbranched or branched, clavate to 
irregular, terete to compressed, with sterile acute 
apices, on short to long stipes, 4-9 cm total length xX 
2-5 mm diam; externally blackish with brown peeling outer 
layer; internally white. Texture woody. Surface rough- 
ened by wrinkles and ostiolar papillae; stipes sometimes 
tomentose. Perithecia 0.4-0.6 mm diam. Ostioles papill- 
ate. Asci eight-spored, cylindrical, stipitate, 144-186 
um total Tength X 7.5-9 um broad, the spore-bearing part 
104-126 um, with apical ring bluing in Melzer's iodine 
reagent, rectangular, 5-6 wm high X 4-5 um broad. 
Ascospores brown to dark brown, unicellular or with 
minute hyaline cellular appendage, ellipsoid-inequi lateral 
to navicular, with narrowed to acute ends, smooth, 
(15.5-)16.5-18(-20) X 6-7(-7.5) um, with spiralling germ 
slit spore-length. 


SPECIMENS EXAMINED: Chacon 110, XI.1987, Ejido La 
Libertad, Llera municipality, Tamaulipas state, on wood 
in riparian site (ITCV:JDR); Garcia 4913B, X.1985 and 
4949, 4.X.1985, "El Cielo" ranch, Gomez Farias 
municipality, Tamaulipas state, on wood in cloud forest 
(ITCV:JDR); San Martin 209, 16.VII.1987, data as Garcia 
(above) (ITCV:JDR). 


NOTES: These collections resemble X. scruposa in asco- 
spore size and gross morphology of stromata. Ascospore 
germ slits are spore-length in this material; they 
usually are less than spore-length in undoubted X. 
scruposa. Our material fits the Saccardo description of 
X. corniculata Sacc. and, although we have not seen type 
material, we tentatively consider it to be this species. 
Our material is reminiscent of Lloyd's photograph (Fig. 
2675)(1924A) of X. corniculata. 


Xylaria cubensis (Mont.) Fr., Nova Acta Regiae Soc. Sci. 


Uosalentserees sp yl 26m 185 le 


Stromata clavate with rounded fertile apices, on 
short stipes, 1.7-5 cm total length X 7-10 mm diam; 
externally copper-colored, blackening with age; internally 
white, becoming hollow and inrolled. Texture hard, but 
fragile. Surface smooth except for osiolar papillae and 
fine cracking. Perithecia 0.4-0.6 mm diam. 


329 


Ostioles finely papillate. Asci eight-spored, cylindri- 
cal, long-stipitate, 125-160 um total length X 4-6 um 
broad, the spore-bearing part 58-65 um long, with apical 
ring bluing in Melzer's iodine reagent, quadrate, 2 X 2 
um. Ascospores brown to almost black, unicellular, 
ellipsoid-inequilateral with rounded ends, smooth, (7-) 
8-10(-11) X 4-5(-6) wm, with germ slit unclear, but 
probably long. 


SPECIMENS EXAMINED: Guevara 723, no date, "San Miguel" 
ranch, Jose Maria Morelos municipality, Quintana Roo 
state, in acahual (ITCV,JDR); *Madrigal Xavier 2138-12, 
near Chocolhaito, Chakalah, Chiapas state (JDR,XAL); San 
Martin 1, 17.VIII.1986; 15, 1985; 24, 14.X.1986, Gomez 
Farias, Tamaulipas state, on wood in subdeciduous rain 
forest (ITCV,JDR); San Martin 387, VI.1987, Ocampo, 
Tamaulipas state, on wood in subdeciduous rain forest. 


NOTES: Most of the material fits our concept of X. 
cubensis (and see Rogers, 1984B; Rogers et al., 1988). 
Pérez-Silva (1975) has described and illustrated it from 
México. ne collection (San Martin 15) is atypical in 
having conspicuous surface scales, solid flesh at matur- 
ity, and somewhat longer ascospores (9-12 X 4-5 ym). 


Xylaria cf. curta Fr., Nova Acta Regiae Soc. Sci. Upsal. 
fers Men 26.) 1851. ElG Seman a2. 


Stromata unbranched or branched, cylindric-clavate 
with rounded fertile apices, on stipes often arising from 
pannose bases, 1.5-3.5 cm total length X 2.5-5 mm diam; 
externally blackish with whitish to tan sloughing scales; 
internally white. Texture woody. Surface roughened by 
wrinkles and scales. Perithecia 0.3-0.5 mm diam. 
Ostioles inconspicuously papillate. Asci eight-spored, 
cylindrical, long-stipitate, 117-158 um total length X 
5-6.5 um broad, the spore-bearing part 65-73 um, with 
apical ring bluing in Melzer's iodine reagent, quadrate, 
1.5-2 X 1.5-2 um. Ascospores brown, unicellular, 
ellipsoid-inequilateral with broad to narrow ends, 
smooth, (9.5-)10-11 X 4-4.5 um, with straight germ slit 
more or less spore-length. 


SPECIMEN EXAMINED: Garcia 5404, 2.VII.1986, Gomez 
Farias, Tamaulipas state, on wood in median subdeciduous 
rain forest (ITCV). 


330 


NOTES: This fungus is placed in X. curta primarily on 
ascospore size and the white scales on stromata. 
Stromata are less robust than usually encountered 
(Dennis, 1956; Rogers, 1983). 


Xylaria cf. dichotoma (Mont.) Fr., Nova Acta Regiae Soc. 
Gi UDSd SCs Rl Deel A ee Onl 
Figs. =sitooe 


Stromata unbranched or branched, cylindrical, terete 
or compressed, bearing perithecia with more or less 
evident contours, with acute sterile apices, on ill- 
defined stipes, up to 7 cm total length X 2-4 mm diam; 
externally dark brownish to blackish; internally white. 
Texture soft to somewhat woody. Surface roughened with 
perithecial contours, wrinkles, and tomentum, the stipes 
conspicuously tomentose. Perithecia 0.3-0.7 mm diam. 
Ostioles finely papillate. Asci eight-spored, the spores 
arranged in a uniseriate or partly biseriate manner, 
cylindrical, long-stipitate, 112-150 um total length X 
4-6.5 um broad, the spore-bearing part 59-75 yum long, 
with apical ring bluing in Melzer's iodine reagent, rec- 
tangular, 2-3 um high X 1.5-2 um broad. Ascospores 
yellowish, unicellular, ellipsoid-inequilateral to fusoid 
or crescentic, smooth, 10-13(-14) X (2.5-)3-4 um, with 
germ slit obscure. 


SPECIMEN EXAMINED: Garza-Ocanas 608, 18.VIII.1986, Ejido 
La Encantada Zaragoza municipality, Nuevo Leon state, on 
Quercus sp. wood in Quercus - Pinus forest (ITCV). 


NOTES: Our material seems to represent X. dichotoma in 

general stromatal morphology and ascospore size. However, 
the ascospores are paler and more crescentic than usually 
encountered and the germ slit obscure. There is the same 
problem in assigning this fungus without question to X. 

dichotoma as encountered in assigning certain collections 
to xe lanthino-velutina and X. magnoliae, as discussed by 


Figs; 99-107... 99,100." Xylaria sp. SM S3l9) 99% 

Stromata, X13.8.. 100. Stromatal surface, X) 16.0.s@ieee 
Xylaria sp. SM 372.. I0Ol1. Stromatal surface, X 16sel0e 
Stromata, X 2.5. 103, 104. Xylaria squamulosa.  s10a3 
Stromata, X 3.6. 104. Stromatal surface, X 16. 105-107. 
Xylaria uniapiculata. 105,106. Stromata, X 1.6.90 
Stromatal surface, X 16. 


331 


332 


Rogers et al. (1988). Until such species or species 
complexes are more widely collected and cultured and 
otherwise compared, their status will remain uncertain. 


Xylaria aff. enterogena (Mont.) Fr., Nova Acta Regiae 
Soc... Sci mUpsal wklser. ssiyaulenip ule) ee oe 
Figs. 34,35,146-148. 


Stromata conical to irregular with rounded fertile 
apices, on short to long stout stipes, 1.8-6.8 cm total 
length X 0.4-3 cm diam, externally dirty white to 
yellowish with black ostioles and fine cracks, the stipes 
reddish brown, pannose and spongy; internally whitish, 
sometimes with perithecia deep within the flesh. Texture 
very hard. Surface smooth except for major wrinkling and 
infolding. Perithecia 0.5-1 mm diam. Ostioles slightly 
raised. Asci eight-spored, cylindrical, stipitate, 
232-276 um total length X 8.5-12 um broad, the spore- 
bearing part 156-170 um long, with apical ring bluing in 
Melzer's iodine reagent, quadrate to slightly cuneate, 
3.5-4 wm high X 2.5-3.5 um broad. Ascospores dark brown, 
unicellular, ellipsoid-inequilateral with broadly or nar- 
rowly rounded ends, smooth, (19-)20-23(-25) X 8-9.5(-10) 
um, with spiralling germ slit. 


SPECIMENS EXAMINED: Jesus Garcfa 7, no date, "Rincon de 
la Sierra", Villa Juarez municipality, Nuevo Leon state, 
on wood in forest of Quercus spp. and Sargentia sp. 
(ITCV); Avalos 25, VIT.1987, Ejido Conrado Castillo, 
Hidalgo municipality, Tamaulipas state, on wood in cloud 
forest (ITCV, JDR); Martinez 26, 8.VII.1984, "El Cielo" 
ranch, GOmez Farias municipality, Tamaulipas state, on 
wood in cloud forest (ITCV, JDR). 


NOTES: Our fungus resembles X. enterogena in the color 
of the fertile part. It differs in being much more 
robust, in having spongy pannose stipes, and in having 
definitely raised ostiolar rims. It has apparently been 
encountered only in México and might represent a new 
taxon. 

Xylaria enterogena is a subject of confusion. 
Dennis (1956) considered it to be a form of X. telfairii. 
Rogers et al. (1988) considered it different from X. 
telfairii by reason of its smaller stromata, ascospore 
characteristics, stromatal color, and cultural 
characteristics. There is no question, however, that X. 
enterogena and X. telfairii are closely related. z 


333 


Martin (1970) indicated that X. enterogena is 
clavate, hollow, and has ascospores with spiralling germ 
slits. This latter feature agrees with our present 
material. However, the size of ascospores of our 
material averages larger than ascospores cited by Martin 
(1970) and Rogers et al. (1988). 

Saccardo (1882), citing Montagne, mentioned solid 
stromata and deeply embedded perithecia in addition to 
more common shallowly placed ones. Some of our material 
shows deeply embedded perithecia. 


Xylaria feejeensis (Berk.) Fr., Nova Acta Regiae Soc. 


Bimeupsaionuser.) 3) 1, p. 128. 71851. 
FIGSen Goce 


Stromata unbranched or branched, cylindric-clavate, 
spathulate, or irregular, with rounded fertile apices, on 
short to long stipes that sometimes originate from 
pannose bases, 2.5-6.5 cm total length X 5-10 mm diam; 
externally dull blackish, the ostiolar area sometimes 
surrounded by white deposits; internally white to 
yellowish. Surface roughened by wrinkles and ostiolar 
papillae and, frequently, by surface cracking and/or 
perithecial contours. Perithecia 0.4-0.8 mm diam. 
Ostioles finely papillate to discoid. Asci eight-spored, 
cylindrical, long-stipitate, 102-135 um total length X 
4.5-5.5 um broad, the spore-bearing part 62-70 um, with 
apical ring bluing in Melzer's iodine reagent, quadrate, 
2 um high X 1.5-2 wm broad. Ascospores brown to nearly 
black, unicellular, ellipsoid-inequilateral with rounded 
ends, smooth, (8-)8.5-11(-12) X 4-4.5(-5) um, with 
Straight germ slit spore-length or nearly so, sometimes 
inconspicuous. 


SPECIMENS EXAMINED: *Alonso, L. 38, San Rafael munici- 
pality, Mexico state, on wood (JDR;XAL); Garcia 3181, 
9.X.1983 and San Martin 391, IX.1987, El Cercado, 
Santiago municipality, Nuevo Leon state, On wood in 
submontane scrub (ITCV;JUDR); Garcia 51065 2.VI121986 and 
San Martin 723, 21.VII.1986, Gomez Farias, Tamaulipas 
state, on wood in median subdec iduous rain forest (5106 
in ITCV;JDR; 723 in ITCV); San Martin 241, 10.VIII.1987, 
Ocampo , Tamaulipas state, on wood in median subdeciduous 
rain forest (ITCV;JDR); San Martin 328, Ejido Boca de 
Chajul, Ocosingo municipality, Chiapas state, on wood in 
evergreen rain forest (ITCV3;JDR). 


334 


NOTES: Our concept of X. feejeensis is generally that of 
Dennis (1956). Most collections could probably be accom- 
modated as X. corniformis (Fr.) Fr. (see Laessde, 1988). 


Xylaria sp. aff. feejeensis Pigvapas 


Stromata unbranched or branched, cylindrical to ir- 
regular, terete to flattened, on abruptly narrowed short 
to long stipes, 1.5-3.5 cm total length X 2.5-4 mm diam; 
externally dull black with remains of a lighter colored 
outer layer; internally whitish. Texture hard. Surface 
rugose with cracks and verrucae. Perithecia 0.3-0.5 mm 
diam. Ostioles hemispherical. Asci eight- spored, cy- 
lindrical, long-stipitate, 112-150 um total length X 
5.5-7 um broad, the spore-bearing part 57-80 um long, 
with apical ring bluing in Melzer's iodine reagent, 
quadrate, 2 X 2 wm. Ascospores light brown to dark brown, 
unicellular, ellipsoid-inequilateral with broadly rounded 
or narrowly rounded ends, smooth, 9-10(-11) X 4-4.5(-5) 
um, with straight germ slit full-length. 


Figs. 108-122. 108-110. Xylaria magniannulata. 108. 
Ascospore with germ slit, X 940. 109. Ascospore ap- 
pendage, X 4125. 110. Ascus apical ring. Arrow points 
to the apex of ring, X 940. 111,112. Xylaria squamulosa. 
111. Ascospore with germ slit, X 940. 112.  Ascus 
apical ring, X 940. 113. Xylaria amphithele. Ascus 
apical ring, X..2055.9114 0 eX Vilaniams paeoMescas 

Ascospore with cellular appendage, X 750. 115,116. 
Xylaria amphithele. 115. Ascospore with noncellular 
appendages (arrows) X 3280. 116. Ascospore with germ 
slit, X 3280. 117-122... Xylaria alata. 117.2) AsSeocuane 
apex showing curled cellular appendage (arrow), X 10,000. 
118. Ascospore with cellular appendages oriented in the 
"broad" dimension (arrows), X 3300. 119. Ascospore with 
cellular appendages oriented in the "narrow" dimension 
(arrows), X 3300. 120. Section of closely packed broad 
paraphyses, X 3300. 121. Ascospores, one showing 
cellular appendages in the "narrow" dimension (arrows), X 
5000. 122. Ascospore with cellular appendages oriented 
in the "broad" dimension, X 7000. 

Figs. 114,115,118,119,120 by darkfield phase micro- 
scopy. Figs. 121,122 by scanning electron microscopy. 
Other figs. by brightfield microscopy. Figs. 110,112,113 
from mounts in Melzer's reagent. Other material examined 
by light microscopy from water mounts. 


335 


336 


SPECIMEN EXAMINED: San Martin 47, 8.X1I.1986, "Los 
Troncones", Ciudad Victoria municipality, Tamaulipas 
state, on wood along stream (ITCV). 


NOTES: This fungus probably represents a small-stromatal 
form of X. feejeensis., See notes on X. feejeensis 
herein. 


Xylaria gracillima (Fr.) Fr., Nova Acta Regiae Soc. Sci. 
Unsaeuttsering) utp eeke Seam boone 
Figs. 37,38, 146e0 eo" 


Stromata unbranched or branched, prostrate or up- 
_right, each stroma consisting of a rachis bearing scat- 
tered to crowded, naked perithecia, extended upward into 
short or long acute apex and downward into a short or long 
tomentose stipe, 3-12 cm total length X 1-3 mm diam. 
Externally black; internally white. Texture soft. Sur- 
face smooth except for naked perithecia. Perithecia 
0.4-0.8 mm diam. Ostioles papillate. Asci eight-spored, 
cylindrical, long-stipitate, 128-165 um total length X 


Figs. 123-144. 123. Xylaria longipes var. tropica. 
Ascospores showing portions of germ slit, X 2055. 124-126. 
Xylaria uniapiculata. 124. Ascospores, one with germ 

slit, X 2055. 125. Ascospore with cellular appendage 
(arrow), X 1800. 126. Ascus apical ring, X 2000. 

127-130. Xylaria guazumae. 127. Ascospore with germ 

slit, X 2055. I28. Ascospore, X 2055. 129. Ascus 

apical ring, X 2000. 130. Ascospore with noncellular 
appendages (arrows), X 1800. 131-133. Xylaria claviceps. 
131. Ascospores, X 2055. 132. Ascospore with germ slit, 

X 2055. 133. Ascus apical ring, X 2000. 134-136. Xylaria 
scabriclavula. 134. Ascus apical ring, X 2000. 135. 
Ascospores, X 2055. 136. Ascospore with germ slit, X 

2055. 137-139. Xylaria juruensis. 137. Ascospore with 
germ slit, X 2055. 138. Ascospore, X 2055. 139. Ascus 
apical ring, X 1560. 140-142. Xylaria cf. laevis. 140. 
Ascospore with slit, X 2055. 141. Ascospore, X 2055. 142. 
Ascus apical ring, X 2055. 143. Xylaria sp. aff. bambooensis. 
Ascospores, X 2055. 144. Xylaria gracillima. Ascospore 
with germ slit, X 2055. 

Figs. 125,130 by darkfield phase microscopy. Al] 
other figs. by brightfield microscopy. Figs. 126,129,133, 
134,139,142 from mounts in Melzer's reagent. Other 
material examined in water mounts. 


337 


338 


6-7.5 um broad, the spore-bearing part 74-84 um, with 
apical ring bluing in Melzer's iodine reagent, rectangular, 
4-4.5 um high X 2.5-3 um broad. Ascospores brown to dark 
brown, unicellular, ellipsoid-inequilateral with broadly 
or narrowly rounded ends, smooth, 10-12(-13) X 4-5 ym 

with straight germ slit less than spore-length. 


SPECIMENS EXAMINED: San Martin 111, 9.XII.1986, Felipe 
Carrillo Puerto, Quintana Roo state, on material buried in 
soil in median subdeciduous rain forest (ITCV); San Martin 
210, 16.VI1.1987, Gomez Farias, Tamaulipas state, on mate- 
rial buried in soil in median subdeciduous rain forest 
(ITCV;UDR). 


NOTES: Dennis (1970) considered this species a synonym of 
X. inaequalis Berk. & Curt. (as Xylosphaera). Our concept 
of X. inaequalis is of a somewhat more robust fungus with 
perithecia placed more or less on one or two sides of a 
rachis, much as illustrated by Dennis (1956). Miller's 
photos of X. gracillima (in possession of JDR) show fusoid 
ascospores. According to Lloyd (1918A) material of Jd. 
Rick has fusiform spores 24-28 X 6 wm; Lloyd accepted 
Rick's interpretation of the species. Clearly, the 
Xylarias with naked perithecia on a narrow rachis need to 
be studied as a group before their individual taxonomic 
and nomenclatural status becomes clear. 


Xylaria grammica (Mont.) Fr., Nova Acta Regiae Soc. Sci. 
Upsalmmiseyees) mite snl Oem one Figs. 40,41. 


Figs. 145-155. 145. Xylaria gracillima. Ascus apical 
ring, X 2055. 146-148. XylTaria aff. enterogena. 146. 
Ascospore with germ slit, X 2055. 147. Ascus apical ring, 
X 1560. 148. Ascospore, X 1560. 149,150. Xylaria kege- 
liana. 149. Ascospore with portion of spiralling germ 
Slats Xml500 wl bO ss BASCUS@apicalaining. e205 oR 151-153. 
Xylaria cf. nigrescens. 151. Ascospore with germ slit, X 
1560.9 9152. "Ascus apical ring, X 1560. 153. Ascospamemm 
1560. 154,155. Xylaria pallide-ostiolata. 154. Asco- 
spore with germ slit, X 2055. 155. Ascus apical ring, X 
1560. 

All figs. by brightfield microscopy. Figs. 145,147, 
150,152,155 from mounts in Melzer's reagent. Other 
material examined in water mounts. 


Gabe 


340 


Stromata unbranched or branched toward base, 
cylindric-clavate to cylindric-fusiform, with apices 
rounded and fertile or acute and sterile, on short to long 
stipes originating from pannose bases, 7-13 cm total 
length X 5-7 mm diam; externally at first whitish, dark- 
ening in such a manner that darker areas and remaining 
whitish areas appear as vertical stripes; internally 
whitish to yellowish, becoming hollow. Surface smooth 
except for stripes. Perithecia 0.4-1.2 mm diam. Ostioles 
punctate, in dark stripes. Asci eight-spored, cylindrical, 
long-stipitate, at least 150-162 um total length X 5-7 um 
broad, the spore-bearing part ca. 70 um long, with apical 
ring bluing in Melzer's iodine reagent, rectangular, 3 um 
high X 1.5-2 um broad. Ascospores brown to dark brown, 
unicellular, ellipsoid-inequilateral with narrowed to 
somewhat pinched ends, smooth, (9.5-)10-12 (-12.5) X 
(4.5-)5-5.5 wm, with straight germ slit spore-length. 


SPECIMENS EXAMINED: San Martin 325, 23.VIII.1987, Ejido 
Loma Bonita, Ocosingo municipality, Chiapas state, from 
wood in evergreen rain forest (ITCV,JDR); Garcia 4891, 
TX.1985, "El Cielo" ranch, Gomez Farias municipality, 
Tamaulipas state, on wood in cloud forest (ITCV); 
*Xelhuantzi, M. S. 100, 4.X1.1978, Cerro del Tepozteco; 
Tepoztlan, Morelos state, on wood (JDR;XAL). 


NOTES: Dennis (1958) believed that specimens differing 
from X. grammica in having thinner stromata and acute 
sterile apices should be referred to X. venustula Sacc. 
We have both kinds of stromata in our collections and 
believe that they represent one and the same species. We 
refer to all of these intergrading forms as X. grammica. 
Cultural and other data bearing on this problem would be 
desirable. Typical X. grammica from Mexico has been 
described and illustrated by Pérez-Silva (1975). 


Xylaria guazumae San Martin & Rogers, sp. nov. 
Figs. 53,54,127-130. 


Stromata discoidea vel subglobosa vel globosa, 
constantia ex aggregatis peritheciorum plus minusve 
nudorum 1-2 mm diam, stipitibus capillaribus usque ad 5 cm 
longis, plerumque solitaria sed interdum caespitosa. 

Extus nigra; intus alba. Textura mollis. Superficies 
asperata a formis peritheciorum. Perithecia 0.5-0.7 mm 
diam. Asci saepe biseriate octospori, cylindrici, 
stipitati, 148-212 um longitudine tota X 7-7.5 um crassi, 


341 


partibus sporiferis 100-132 um, annulo apicali in liquore 
Melzeri cyanescente, oblongo, 2.8-3.2(-3.8) wm alto X 2- 
2.5(-3) um crasso. Ascosporae brunneae, unicellulares, 
ellipsoideo-inaequilaterales cum late vel anguste 
rotundatis extremis, omni extremo cum rotundata hyalina 
noncellulari appendicula usque ad 5 um longa X 5 wm crassa, 
Teves, (14-)15-18(-19) X 5.5-6 um, rima germinativa 

leviter minus quam longitudo sporae. Status anamorphosis 
ignotus. 


Stromata discoid to subglobose to globose, composed 
of aggregations of more or less naked perithecia 1-2 mm 
diam, with hair-like stipes up to 5 cm long, usually 
solitary, but sometimes cespitose. Externally black; 
internally white. Texture soft. Surface roughened by 
perithecia! contours. Perithecia 0.5-0.7 mm diam. 
Ostioles conical. Asci eight-spored, the spores often 
arranged in a biseriate manner, cylindrical, stipitate, 
148-212 um total length X 7=7.5 um broad, the spore- 
bearing part 100-132 um, with apical ring bluing in 
Melzer's iodine reagent, rectangular, 2.8-3.2(-3.8) um 
high X 2-2.5(-3) um broad. Ascospores brown, unicellular, 
ellipsoid-inequilateral with broadly or narrowly rounded 
ends, each bearing a rounded hyaline noncellular appendage 
up to 5 um long X 5 um broad, smooth, (14-)15-18(-19) X 
5.5-6 um, with straight germ slit slightly less than 
full-length. Anamorph unknown. 


SPECIMENS EXAMINED: San Martin 208, 1987, Gomez Farias, 
Tamaulipas state, on fallen fruits of Guazuma ulmifolia 
Lam. ("aquiche" or "guazima") in median subdeciduous rain 
forest (HOLOTYPE: ITCV; ISOTYPE:JDR); San Martin 343, 
2.VIII.1987, Ocampo, Tamaulipas state, on fallen fruits of 
Guazuma ulmifolia Lam. in low deciduous rain forest 

(ITCV; UDR). 


NOTES: The present fungus resembles X. heloidea, but 
ascospores of that species do not bear conspicuous 
secondary appendages. 


Xylaria guyanensis (Mont.) Fr., Acta Regiae Soc. Sci. 
poouemsene oyelsnperle7s) Lop ls Figse14?.43. 


Stromata clavate with rounded fertile apices, on 
short stipes with discoid bases, 4-5 cm total length X 
0.8-1.2 cm diam; externally gray-yellowish, becoming 
blackened; internally white to yellowish, becoming hollow. 


342 


Texture hard to very hard but fragile. Surface smooth 
except for minute cracks and ostioles. Perithecia 0.8-2 
mm diam. Asci not intact, cylindrical, eight-spored, the 
spore bearing part 110-120 X 5.5-6 um, with apical ring 
bluing in Melzer's reagent, quadrate to almost rectangular, 
2.8-3 um high X 2-3 um broad. Ascospores brown, unicell- 
ular, ellipsoid-inequilateral to navicular, with rounded 
ends, smooth, (14-)15-17.5 X 4.5-6(-7) um, with straight 
germ slit spore-length. 


SPECIMEN EXAMINED: Garcia 2734, 10.XI.1982, from Catemaco 
to Montepio Balzapote, Veracruz state, on wood in ever- 
green rain forest (ITCV,JDR) 


NOTES: Xylaria guyanensis is much like X. poitei, but has 
generally smaller stromata and larger ostiolar discs. 
Xylaria poitei has an easily identifiable conidial state 
that is produced in culture. It is hoped that future 
collections of X. guyanensis can be cultured and compared 
with those of X. poitei (Rogers & Callan, 1986B). 
Perez-Silva (1975) described and illustrated X. guyanensis 
from Mexico. She gave ascospores as 18-20.4 X 6-7 um, 
considerably longer than reported here. Dennis (1956) 
reported ascospores of South American material as 14-21 X 
5-8 um. 


Xylaria ianthino-velutina (Mont.) Fr., Nova Acta Regiae 


SocumScisaUpsa lm Sergs)) aden Dem co eee oe 


Stromata unbranched or branched, cylindrical to 
irregular, bearing more or less naked perithecia, terete 
to flattened, with acute sterile apices, on ill-defined 
Stipes, 2-13 cm total length X 1:.5-2.5 mm diam; externally 
reddish brown to black, tomentose over all; internally 
white. Texture rather soft. Surface roughened with 
perithecial elevations and tomentum. Perithecia 0.3-0.6 
mm diam. Ostioles finely papillate. Asci eight-spored, 
cylindrical, stipitate, 85-120 um total Tength X 5-6 um 
broad, the spore-bearing part 60-90 um, with apical ring 
bluing in Melzer's iodine reagent, rectangular, 2-2.5 um 
high X 1-2 um broad. Ascospores light brown to dark 
brown, unicellular, ellipsoid-inequilateral with rounded 
to narrowly rounded ends, smooth, 10-12(-13) X 4-5 um, 
with straight germ slit spore-length. 


SPECIMENS EXAMINED: Garcia-Alanis 60, XI.1987, Aldama, 
Tamaulipas state, on fallen fruits of Pithecellobium 


343 


flexicaule (Benth.) Coult. ("maguacata") in mixed tall 
Scrub-low subdeciduous rain forest JAITCV): Guevara 724, 
1986, "San Miguel" ranch, José Maria Morelos municipality, 
Quintana Roo state, on pods in low subdeciduous rain 
forest (ITCV); San Martin 22, 14.X.1986, Gomez Farias, 
Tamaulipas state, on pod in median subdeciduous rain 
forest (ITCV). San Martin 377, IX.1987, (same data as San 
Martin 22, above), on fallen fruits of Guazuma ulmifolia 
Lam. (ITCV;JDR). 


NOTES: These collections generally fit our concept of X. 
janthino-velutina. Typical material has dark ascospores 
with evident germ slits (Dennis, 1956; Rogers, 1979). 
Some material has pale ascospores with less conspicuous 
germ slits, resembling in some respects X. magnoliae. 
That species, however, appears confined to Magnolia 
(Rogers, 1979). Discussions of X. ianthino-veTutina and 
allied forms or taxa with pale ascospores are in Rogers 
(1979) and Rogers et al. (1988). 

It is perhaps noteworthy that Guazuma ulmifolia is 
herein reported as a substrate for X. ianthino-velutina, a 
species usually reported from legume pods. Elsewhere 
herein, X. aristata and X. quazumae are also reported from 
fruits of G. ulmifolia. It is becoming evident that G. 
ulmifolia fruits represent a substrate for diverse and, 
perhaps ultimately numerous, species of Xylaria. 


Xylaria cf. inaequalis Berk. and Curt., J. Linn. Soc. Bot. 
30.0) 


Stroma branched, irregular, flattened, bearing a few 
more or less naked perithecia on one side of a branch, 
with acute sterile apices, on long pubescent stipe, 2 cm 
total length X 1.5 mm diam; externally brownish black, 
internally yellowish. Texture soft. Surface roughened by 
perithecial contours and pubescence of stipe. Perithecia 
0.2-0.4 mm diam. Ostioles more or less umbilicate. Asci 
eight-spored, cylindrical, long-stipitate, 116-127 um 
total length X 5.5-6 um broad, the spore-bearing part 
59-66 um, with apical ring bluing in Melzer's iodine 
reagent, quadrate, 1.5-2 X 1.5-2 wm. Ascospores brown to 
dark brown, unicellular, ellipsoid-inequilateral with 
mostly broadly rounded ends, smooth, 10-11(-11.5) X 
4-4.5(-5) um, with straight germ slit spore-length. 


SPECIMEN EXAMINED: San Martin 99, 8.VII.1986, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 


344 


on fallen fruit of Swietenia macrophylla King ("caoba"} in 


evergreen rain forest (ITCV). 


NOTES: This fungus is represented by only one stroma. It 
seems much like X. inaequalis as depicted by Dennis (1956) 
in the asymmetrical insertion of perithecia. See also 
"Notes" on X. gracillima herein. i 


Xylaria juruensis P. Henn., Hedwigia 43:262. 1904. 
Figs. 45,46,137-139. 


Stromata unbranched or branched, cylindrical to 
irregular, terete to compressed, bearing more or less 
naked perithecia, with short to long hair- like apiculi, 
on long thin stipe, 1.5-3 cm total length X 1 mm diam; 
externally blackish with gray peeling layer, pubescent 
overall; internally white. Texture fairly hard. Surface 
roughened by peeling layer, perithecial contours, and 
pubescence. Perithecia 0.2-0.6 mm diam. Ostioles 
umbilicate to slightly raised. Asci eight-spored, the 
spores usually arranged in a partly biseriate manner, 
cylindrical, stipitate, 127-152 um total length X 6-7(-10) 
um broad, the spore-bearing part 82-102 um, with apical 
ring bluing in Melzer's iodine reagent, rectangular, 5-7 
um high X 2-4 um broad. Ascospores brown, unicellular, 
ellipsoid-inequilateral with rounded to acute ends, 
smooth, 14.5-17(-18) X 5-5.5(-6.5) um, with straight germ 
slit slightly less than spore-length. 


SPECIMEN EXAMINED: San Martin 293, VIII.1987, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 
on wood in median subdeciduous rain forest (ITCV3;JDR). 


NOTES: The type was from monocot material from an Amazon 
jungle (Hennings, 1904). Rogers & Samuels (1987) 
described a collection from a palm midrib from Sulawesi. 
The present material is from dicot wood and fits Hennings' 
description well (Hennings, 1904). Carrol] (1964) illust- 
rated a spiralling germ slit for X. juruensis. 


Xylaria kegeliana (Lev.) Fr., Nova Acta Regiae Soc. Sci. 


Upsal. (ser. 3) 1, p. 125. 1851; sensu J. H. Miller. 
Figs. 44,47,149,150. 


Stromata unbranched or branched, cylindrical to 


eer et enn ante 


clavate with abrupt acute sterile apices, on short to long 
Stipes, 4-12 cm total length X 4-7 mm diam; externally 


345 


dirty white to light tan with black ostioles, often 
appearing vaguely longitudinally striped, with blackish 
stipes; internally white, becoming hollow. Texture fairly 
hard. Surface smooth except for ostiolar papillae and 
minute cracks. Perithecia 0.4-1.2 mm diam. Ostioles 
papillate, often in vertical rows. Asci eight-spored, 
cylindrical, long-stipitate, 317-337 wm total length X 
7.5-8 um broad, the spore-bearing part 177-192 um, with 
apical ring bluing in Melzer's iodine reagent, rectangular, 
5-6 um high X 3-3.5 um broad. Ascospores brown, unicell- 
ular, ellipsoid-inequilateral to navicular, with narrow to 
abruptly pinched ends, smooth, (27-)28-33.5(-35) X 
6-7.5(-8) um, with spirailing germ slit spore-length. 


SPECIMEN EXAMINED: San Martin 36, X.1986, Gomez Farias, 
Tamaulipas state, on wood in median subdeciduous rain 
forest (ITCV). 


NOTES: This material fits exactly J. H. Miller's concept 
of X. kegeliana. Unpublished photographs of Lloyd 
material by Miller (in possession of JDR) show stromata 
and ascospores like those described and depicted herein. 
Dennis (1956), discussing X. kegeliana, wrote, "In stature 
this recalls X. grammica, in color and spores it resembles 
X. dealbata." The present fungus greatly resembles X. 
grammica in gross morphology and somewhat resembles jt in 
the color and vague vertical striping of the stroma. It 
does not, however, resemble X. dealbata either in color or 
ascospore morphology. Rogers et al. (1988) described a 
fungus from Venezuela that was tentatively considered to 
be X. kegeliana. That fungus had coloration much more 
like X. dealbata than the fungus discussed herein. Its 
ascospores are much like the fungus discussed herein in 
general morphology, but are smaller and have a short, 
nonspiralling germ slit. The relationship of these 
distinct fungi to one another is unknown. 


Xylaria cf. laevis Lloyd, Xylaria Notes 1, Mycol. Writ. 
Pes NL OL oc Figs. 48 ,140-142. 


Stromata unbranched or branched at base, with rounded 
fertile apices, on short or long fluted stipes arising 
from pannose base, 1.6-2.3 cm total length X 4-6 mm diam, 
externally copper-colored, becoming blackish; internally 
white, becoming hollow. Texture hard. Surface smooth 
except for fine wrinkles and ostioles. Perithecia 0.3-0.5 
mm diam. OQstioles finely papillate. Asci eight-spored, 


346 


cylindrical, long-stipitate, 105-195 um total length X 5-6 
um broad, the spore-bearing part 50-69 um long, with apical 
ring bluing in Melzer's iodine reagent, quadrate to rec- 
tangular, 2-3 um high X 2 um broad. Ascospores brown, 
unicellular, ellipsoid-inequilateral with broadly or 
narrowly rounded ends, smooth, (8.5-)9-11(-12) X 4-5 um, 
with germ slit straight, spore-length. 


SPECIMEN EXAMINED: Chacon 111, X1I.1987, Ejido Loma 
Bonita, Llera municipality, Tamaulipas state, on wood in 
low subdeciduous forest (ITCV,JDR). 


NOTES: This fungus is close to X. cubensis, differing 
primarily in the slightly longer (average) ascospores with 
conspicuous germ slits. It is probably X. laevis, which 
was described as a small-spored form of X. nigrescens. 


Xylaria cf. longiana Rehm, Ann. Mycol. 2:175. 1904. 
Figs. 88,89. 


Stromata unbranched or branched from base or above, 
cylindrical, terete or flattened, bearing conspicuous 
perithecia, with acute sterile apices, with ill-defined 
stipes, 2-6 cm total length X 2-4 mm diam; externally 
black sometimes with remains of white peeling outer layer; 
internally white with black medulla. Texture fairly soft. 
Surface of stipe and usually fertile part conspicuously 
tomentose, roughened by perithecial elevations and peeling 
outer layer. Perithecia 0.2-0.5 mm diam. Ostioles 
umbilicate to somewhat papillate. Asci eight-spored, 
cylindrical, long-stipitate, 102-136 wm total length X 5-6 
um broad, the spore-bearing part 46-70 um long, with 
apical ring bluing in Melzer's iodine reagent, quadrate, 
1.5-2.5 X 1.5-2.5 um. Ascospores brown to dark brown, 
unicellular, ellipsoid-inequilateral with broadly or 
narrowly rounded ends, smooth (7.3-)9-10(-11) X 4-5(-5.5) 
um, with straight germ slit spore-length or nearly so. 


SPECIMENS EXAMINED: Chacon-Jiménez 360, XI.1987, Ejido 
Julilo, Gomez Farias municipality, Tamaulipas state, on 
wood in Pinus-Quercus forest (ITCV;JDR); Garcia Alanis 10, 
1987, Ejido Conrado Castillo, Hidalgo municipality, 
Tamaulipas state, on Quercus sp. wood in Pinus-Quercus 
forest (ITCV;JDR); *Vargas, J. 284, 12.VITI.1969, 20 km 
from Teotitlan, road to Huautla de Jiménez, Oaxaca state, 
on wood (JDR;XAL). 


347 


NOTES: These collections are much like X. longiana, but 
the heavy tomentum on the clavae and the stipes of the 
Chacon-Jimenez and Alanis specimens is unusual in our 
experience. Rehm remarked on the tomentose stipe in the 
Original description. Type material is from Quercus from 
Texas. We have not examined it. What we are calling X. 
longiana here is often considered to be a small-spored form 
of X. hypoxylon and has the same general morphology as that 
species. Pérez-Silva (1975) reported typical X. hypoxylon 
on the basis of immature specimens and cultures. We have 
not encountered typical (large-spored) X. hypoxylon, but 
would be surprised if it were not extant and, indeed, 
common in northern parts and/or high altitudes. 


Xylaria longipes Nits., Pyrenomycetes Germanici, p. 14. 
1867. Figs. 49,50. 


Stromata unbranched or branched, cylindrical to 
clavate to spathulate, with rounded fertile apices, on 
short to long stipes, 4-7 cm total length X 5-9 mm diam; 
externally dark brown with areas of sloughing whitish 
scales; internally white. Texture woody. Surface 
roughened with wrinkles, verrucae, and ostiolar papillae. 
Perithecia 0.3-0.8 mm diam. Ostioles papillate. Asci 
eight-spored, cylindrical, stipitate, 151-183 um total 
length X 6-7 wm broad, the spore-bearing part 106-116 um, 
with apical ring bluing in Melzer's iodine reagent, 
quadrate to rectangular, 2-2.5 um high X 2-2.7 um broad. 
Ascospores brown to dark brown, unicellular, ellipsoid- 
iInequilateral to navicular with narrowed ends, smooth, 
13-16 X 4-5(-5.5) um, with spiralling germ slit full- 
length. 


SPECIMEN EXAMINED: Chacon 384, XI.1987, Ejido Julilo, 
Gomez Farias municipality, Tamaulipas state, on wood in 
cloud forest (ITCV;JDR); San Martin 3, X.1985, "El Cielo" 
ranch, Gomez Farias municipality, Tamaulipas state, on 
wood in cloud forest (ITCV;JDR). 


NOTES: Xylaria longipes is widely considered to be an 
inhabitant of more northern parts of the temperate zone. 
It is widely associated with Acer spp. in North America 
and Europe. Dennis (1956) reported one collection from 
Brazil and, according to him, J. H. Miller reported a 
collection from Venezuela. Unfortunately, our material 
could not be cultured and compared with cultures from 
elsewhere (see Rogers, 1983). 


348 


Xylaria longipes Nits. var. tropica San Martin & Rogers, 


var. nov. 
Typical variety in Pyrenomycetes Germanici, p. 14. 
1867. Figs sofas 


A varietate typica differt in stromatibus cavis, in 
superficie rimosa, et in ascis brevioribus. 

Differs from typical variety in the hollow stromata, 
in the cracked surface, and in the shorter asci. 

Stromata cylindrical to clavate with rounded fertile 
apices, with short to long stipes, 3.2-6.2 cm total length 
X 3-7 mm broad; externally blackish with sloughing brown 
scales associated with cracking; internally white, 
becoming hollow. Texture woody. Surface roughened with 
wrinkles, scales, and ostiolar papillae. Perithecia 
0.3-0.7 mm diam. Ostioles conical to hemispherical, often 
coated with white granular material. Asci eight-spored, 
cylindrical, stipitate, 150-170 um total Tength X 6-7.5 ym 
broad, the spore-bearing part 90-95 um long, with apical 
ring bluing in Melzer's iodine reagent, quadrate to rec- 
tangular, 2-3 um high X 2-2.5 um broad. Ascospores brown, 
unicellular, ellipsoid-inequilateral to navicular with 
narrow ends, smooth, (13-)14-16 X 5-6 um, with spiral- 
ling germ slit spore-length. 


SPECIMEN EXAMINED: San Martin 327, VIII.1987, Ejido 
Bocade Chajul, Ocosingo municipality, Chiapas state, on 
wood in evergreen rain forest (ITCV:JDR). 


NOTES: This fungus appears to be a form of X. longipes, 
differing from typical material in becoming hollow, in the 
cracking of the surface, and in the shorter asci. The 
white material around ostioles is noteworthy, but is not a 
constant feature. It is accordingly described as a new 
variety. 


Xylaria magniannulata San Martin & Rogers, sp. nov. 
Figs. 55,56,108-110. 


Stromata non ramosa vel ramosa super stipites, cylin- 
dracea, peritheciis plus minusve immersis, apicibus acutis 
Sterilibus, in brevibus vel longis stipitibus, 4 cm longa 
X 1.5-3 mm crassa; extus atra, superficie cinereo- 
glaucescenti; intus alba. Textura plus minusve mollis. 
Superficies asperata formis peritheciorum et papillis 
ostiolorum magnarum. Perithecia 0.5-0.7 mm diam. Ostiola 
conica, valde conspicua. Asci saepe biseriate octospori, 


349 


cylindrici, breviter stipitati, 206-250 um longitudine 
tota X 17-24 um crassi, partibus sporiferis 182-230 um, 
annulo apicali in liquore Melzeri cyanescente, cylin- 
draceo, 21-40 um alto X 8.5-11 um crasso. Ascosporae 
brunneae, unicellulares, ellipsoideo-inequilaterales cum 
rotundatis extremis, totae ab acellularibus appendiculis 
hyalinis et vaginis hyalinis indutae, 34-45(-47) X (14-) 
15-20(-21) um, rima germinativa recta per longitudinem 
Sporae. Status anamorphosis ignotus. 


Stromata unbranched or branched above stipes, cylin- 
drical, bearing partly immersed perithecia, with acute 
Sterile apices, on short to long tomentose stipes, 4 cm 
high X 1.5-3 mm diam; externally black with grey bloom; 
internally white. Texture fairly soft. Surface rough 
with perithecial contours and large ostiolar papillae. 
Perithecia 0.5-0.7 mm diam. Ostioles conical, very 
conspicuous. Asci eight-spored, the spores arranged in 
partly biseriate or uniseriate manner, cylindrical, short- 
Stipitate, 206-250 um total length X 17-24 um broad, the 
spore-bearing part 182-230 um, with apical ring bluing in 
Melzer's iodine reagent, cylindrical, 21-40 um high X 
8.5-11 wm broad. Ascospores dark brown, unicellular, 
ellipsoid-inequilateral with rounded ends, bearing hyaline 
noncellular appendages and hyaline sheath, smooth, 34-45 
(-47) X (14-)15-20(-21) um, with straight germ slit 
spore-length. 


SPECIMEN EXAMINED: San Martin 292, VIII.1987, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 
on soil, possibly associated with insect nests, in median 
subdeciduous rain forest (HOLOTYPE:ITCV). 


NOTES: This fungus is notable in the large size of asco- 
spores and the massive apical ascus ring, among the 
largest known. It is named for this latter feature, i.e., 
the ascus is large-ringed. The large coarse ostiolar 
papillae and the ascospore sheath and noncellular append- 
ages are also highly characteristic. Everything 
considered, X. magniannulata is a highly distinctive 
fungus. oF 


Xylaria magnoliae Rogers, Canad. J. Bot. 57:941. 1979. 
AGS wo 7oos 


Stromata branched or unbranched, cylindrical to 
irregular, tomentose, bearing scattered to crowded more or 


350 


less naked perithecia, with acute sterile apices, on short 
to long fluted tomentose stipes, 4-7 cm total length X 
1.5-3.5 mm diam; externally dull blackish; internally 
yellowish. Texture woody. Surface roughened by peri- 
thecial contours and tomentum. Perithecia 0.3-0.4 mm 
diam. Ostioles papillate. Asci eight-spored, the spores 
arranged in uniseriate or partly biseriate manner, cylin- 
drical, stipitate, 110-150 um total length X 5-8 um broad, 
the spore-bearing part 75-85 um, with apical ring bluing 
in Melzer's iodine reagent, quadrate to rectangular, 1.5- 
2.2 um high X 1-1.5 um broad. Ascospores subhyaline to 
yellowish, uniceilular, ellipsoid-inequilateral with 
narrow to more or less acute ends, smooth, 12-16(-17) X 
3-5 um, lacking obvious germ slit. 


SPECIMEN EXAMINED: San Martin 254, I1X.1987, Ejido Conrado 
Castillo, Hidalgo municipality, Tamaulipas state, on 
fallen fruits of Magnolia schiedeana Schl. in cloud forest 
(ITCV;JDR). 


NOTES: This fungus seems typical of X. magnoliae, as 
described by Rogers (1979). Somewhat similar fungi have 
been found on non-Magnolia hosts in South America (see 
Rogers et al., 1988). 


Xylaria maitlandii (Dennis) D. Hawksw., Trans. Brit. 
Mycol. Soc. 61:199. 1973. Figs. 59960 


Stromata unbranched or branched, cylindrical bearing 
immersed perithecia, with acute sterile apices, on short 
to long pubescent stipes, up to 3 cm total length X 1-2 mm 
broad; externally black with brown peeling outer layer and 
Slight pubescence; internally at first white, becoming 
brown and then hollow. Texture hard. Surface smooth 
except for peeling layer and pubescence. Perithecia 
0.3-0.5 mm diam. Ostioles umbilicate to slightly raised. 
Asci eight-spored, poor condition in this material, the 
Spore-bearing part 78-83 X 5-6 um, with apical ring bluing 
in Melzer's iodine reagent, quadrate, 2 X 2 um. Ascospores 
brown, unicellular, ellipsoid-inequilateral with broadly 
or narrowly rounded ends, smooth, 11-12(-12.5) X (4-)5-5.5 
um, with straight germ slit less than spore-length. 


SPECIMEN EXAMINED: San Martin 320, VIII.1987, Ejido Boca 
de Chajul, Ocosingo municipality, Chiapas state, on wood 
in evergreen rain forest (ITCV;JDR). 


351 


NOTES: Our material looks much like X. maitlandii, 
described by Dennis (as Xylosphaera) from Africa (1958) 
except that the fertile part 1s not as hairy as type 
material. 


Xylaria cf. mellisii (Berk. ) Cooke var. nuda (Dennis) D. 
Hawksw., Trans. Brit. Mycol. Soc. 61:200. 1973. 
ElQe 01% 


Xylosphaera mellisii (Berk. ) Dennis var. nuda Dennis, 


Rev. Biol. 1:188. 1958. 


Stromata unbranched or branched, cylindrical bearing 
more or Tess naked perithecia, with acute sterile apices, 
on short to long stipes, 3-5 cm total length X 2-3 mm 
diam; externally blackish with whitish to brown outer 
peeling layer; internally white to cream. Texture hard. 
Surface roughened by perithecial contours and peeling 
layer. Perithecia 0.4-0.8 mm diam. Ostioles umbilicate 
to finely papillate. Asci eight-spored, cylindrical, 
stipitate, 157-200 um total length X 6-7 um broad, the 
spore-bearing part 97-105 um, with apical ring bluing in 
Melzer's iodine reagent, quadrate to rectangular, 2.5-3.5 
um high X 2-2.5 um broad. Ascospores brown, unicellular, 
ellipsoid-inequilateral with rounded ends, smooth, (12.5-) 
13-16(-18) X 5-6 um, with straight germ slit spore-length. 


SPECIMEN EXAMINED: San Martin 218, 16.VII.1987, Gomez 
Farias, Tamaulipas state, on wood in median subdeciduous 
rain forest (ITCV;JDR). 


NOTES: Our material is much as described by Dennis (1958) 
from Africa, except that the fertile parts of our fungus 
are longer (2 cm in some cases). Dennis considered X. 
arbuscula to be a synonym of X. mellisii. We have not 
evaluated this complicated situation and prefer to wait 
until the entire group of species can be studied and, 
preferably, cultured. We are, however, on shaky taxonomic 
and nomenclatural ground in using both X. mellisii and X. 
arbuscula herein. 7 , 


Xylaria cf. microceras (Mont.) Fr., Nova Acta Regiae Soc. 


UosaleiGSers oye, Pp. 128.7 1851. FT OS m0, 05. 


Stromata cylindrical bearing inconspicuous to con- 
Spicuous perithecial undulations, with long acute sterile 
apices, on short glabrous stipes, 1.5-3 cm total length X 


352 


2 mm diam; externally blackish with white peeling outer 
layer; internally white. Texture fairly hard. Surface 
smooth except for perithecial contours and peeling layer. 
Perithecia 0.3-0.6 mm diam. Ostioles slightly raised. 
Asci eight-spored, cylindrical, long-stipitate, 127-178 um 
total length X 5-6 um broad, the spore-bearing part 67-78 
um, with apical ring bluing in Melzer's iodine reagent, 
quadrate, 2 X 2 um. Ascospores brown to dark brown, 
unicellular, ellipsoid-inequilateral with rounded ends, 
smooth, (9-)10-11 X 4-4.5(-5) um, with straight germ slit 
less than full-length. 


SPECIMEN EXAMINED: San Martin 219, Gomez Farias, 
Tamaulipas state, on wood in median subdeciduous rain 
forest (ITCV). 


NOTES: Our material is much like X. coccophora reported 
herein but has slightly smaller ascospores. It might be 
more logically referred to X. coccophora because of the 
great resemblance of the stromata to those of that species 
(see Dennis, 1956; Rogers et al., 1988). 


Xylaria multiplex (Kunze) Fr., Nova Acta Regiae Soc. Sci. 
Upsal. (ser. 3) 1, p. 127. 1851; sensu Dennis, Kew 
Bul el956:: 4162 2956. Figs. 64,65. 


Stromata unbranched or branched, solitary to 
cespitose, cylindrical bearing perithecia partially 
immersed, with acute sterile apices, on short or long more 
or less pubescent stipes often arising from pannose bases, 
1-5.5 cm total length X 1-2 mm diam; externally blackish 
with dark brown peeling outer layer; internally white, 
often black at center. Texture hard. Surface smooth to 
undulate to nodulose from perithecial contours, sometimes 
with scattered hairs. Perithecia 0.3-0.5 mm diam. 
Ostioles more or less punctate. Asci eight-spored, 
cylindrical, stipitate, 109-142 um total length X 5-6 um 
broad, the spore-bearing part 65-74 um, with apical ring 
bluing in Melzer's iodine reagent, quadrate, 1.5-2 X 1.5-2 
um. Ascospores brown, unicellular, ellipsoid-inequilateral 
with broadly to narrowly rounded ends, smooth, 9-11(-12) xX 
(3.5-)4-4.5(-5) um, with straight germ slit spore-length. 


SPECIMENS EXAMINED: *Brown, D. 50, 28.1X.1981, park on 
old Xalapa-Coatepec road, Veracruz state, on wood 
(JDR;XAL); Garcia 3195, El Cercado, Santiago municipality, 
Nuevo Leon state, on wood in submontane scrub with Quercus 


353 


spp. (ITCV); *Martinez, D. 214, VII.1981, Estacion 
Biologica Los Tuxtlas, Catemaco, Veracruz State, on wood 
in evergreen rain forest (JDR; XAL) 3 San Martin 150, 
8.XII1.1986, Ejido La Union, Othon P. Blanco municipality, 
Ouintana Roo state, on wood in submedian subdeciduous rain 
forest (ITCV;JDR); San Martin 204, X.1987, Ocampo, 
Tamaulipas state, on wood in low Subdeciduous rain forest 
(ITCV;JDR). 


NOTES: These collections generally fit our concept of X. 
multiplex. Stromata are not as robust as those sometimes 
encountered. Xylaria sp. (SM 39, 306) described herein 
might be a variant of this species. Perez-Silva (1975) 
has described and illustrated this species from Mexico. 


Xylaria "muscula" Lloyd, Mycol. Notes 64, Mycol. Writ. 
6:994. 1920. Figs.-66,6/7. 


Stromata unbranched or several originating from 
common base, cylindrical with sterile or fertile apices, 
on short or long stipes, 0.5-3.5 cm total length X 1-3 mm 
diam; externally white with black ostioles; internally 
cream to light brown. Texture fairly hard. Surface 
slightly roughened by ostioles. Perithecia 0.2-0.4 mm 
diam. Ostioles umbilicate to slightly raised. Asci 
eight-spored, cylindrical, stipitate, 70-102 um total 
length X 5-6 um broad, the spore-bearing part 48-62 um, 
with apical ring bluing in Melzer's iodine reagent, quad- 
rate, 1-1.5 wm X 1-1.5 um. Ascospores light brown, 
unicellular, ellipsoid-inequilateral with broadly rounded 
ends, smooth, 6-9(-10) X 3-3.5(-4) um, with inconspicuous 
Straight germ slit spore-length. 


SPECIMENS EXAMINED: Garcia 4348, 14.X.1986, Gomez Farias, 
Tamaulipas state, on wood in median subdeciduous rain 
forest (ITCV;JDR); San Martin 94, 7.XII.1986, San Felipe 
Racalar, Othon P. Blanco municipality, Quintana Roo state, 
on wood in acahual (ITCV;JDR); San Martin 312, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 
on wood in median subdeciduous rain forest (ITCV;JDR). 


NOTES: Our material greatly resembles Lloyd's description 
and photographs. According to Dennis (1956) type material 
of X. muscula is sterile. If this is true, a neotype 
should be selected and a proper description published. 


354 


Xylaria cf. nigrescens (Sacc.) Lloyd, Xylaria Notes 1, 
Mycol J eWritemb Si nl918:. Figs. 68,695 U5 teas 


Stromata subglobose to clavate with rounded fertile 
apices, on short to long stipes, 2-6 cm total length X 
5-10 mm broad; externally dirty brown to dull black, 
internally at first white, becoming hollow and inrolled. 
Texture hard to very hard. Surface smooth except for minor 
wrinkles and ostiolar papillae; stipes smooth or somewhat 
tomentose, with discoid bases. Perithecia 0.5-1 mm diam. 
Ostioles hemispherical, prominent. Asci eight-spored, 
cylindrical, stipitate but stipes mostly deteriorated, the 
spore-bearing part 129-133 wm long X 8-9 um broad, with 
apical ring bluing in Melzer's iodine reagent, rectangular, 
6-9 um high X 4-5 um broad. Ascospores light brown to 
dark brown, unicellular, ellipsoid-inequilateral to 
navicular, with ends smoothly rounded or abruptly pinched, 
smooth, 22-32(-35) X 6-8 um, with germ slit straight to 
oblique, much less than spore-length. 


SPECIMENS EXAMINED: San Martin 348 and 370, VIII. 1987, 
Ejido Boca de Chajul, Ocosingo municipality, Chiapas 
state, on wood in evergreen rain forest (ITCV;JDR). San 
Martin 355, VIII. 1987, "El Muneco", Ejido Loma Bonita, 
Ocosingo municipality, Chiapas state, on wood in evergreen 
rain forest (ITCV,JDR). 


NOTES: Our material has a striking resemblance Lloyd's 
Fig. 1213 of Xylaria nigrescens (Lloyd, 1918B). His 
material, from Africa, was reported to have ascospores 20 
X 8 um. Ina later publication another African collection 
is described as having ascospores. 20-24 X 8 um and 
illustrated (Fig. 1564) (Lloyd, 1919). Photographs of 
ascospores of Lloyd material taken by J. H. Miller (in 
possession of JDR) are much like those of our material. 
Thus, although ascospores of our material average larger 
than given by Lloyd and the fact that his material was 
from Africa, we believe it likely that our fungus is X. 
nigrescens. = 


Xylaria oxyacanthae Tul., Selecta Carpologia Fungorum 2, 
Be We Nem Figs. ./037 


Stromata unbranched or branched toward base, 
cylindrical to irregular, terete to somewhat flattened, 
with short acute sterile apices, on long tomentose stipes 
originating from pannose bases, 3-7 cm total length X 


355 


1-2.5 mm diam; externally blackish with gray to brown 
peeling outer layer; internally white to yellowish. 
Texture soft to woody. Surface roughened with wrinkles 
and perithecial contours. Perithecia 0.3-0.6 mm diam. 
Ostioles finely papillate. Asci eight-spored, cylindrical, 
long-stipitate, 114-164 um total length X 6-7.5 um broad, 
the spore-bearing part 65-80 um, with apical ring bluing 
in Melzer's iodine reagent, quadrate, 1.5-2.5 X 1.5-2.5 
um. Ascospores brown to dark brown, unicellular, 
ellipsoid-inequilaterial with broadly or narrowly rounded 
ends, smooth, (9.5-)10-11.5(-12) X 5-5.5 um, with straight 
germ slit full-length or nearly so. 


SPECIMENS EXAMINED: San Martin 210A, 16.VII.1987, Gomez 
Farias, Tamaulipas state, on buried seeds of Vitaceae, in 
median subdeciduous rain forest (ITCV;JDR); San Martin 
339, VIII.1987, (data same as above), on buried leguminous 
pod (ITCV). 


NOTES: This material looks much like X. oxyacanthae from 
northern United States which occurs on Crataegus seeds 
(Stowell & Rogers, 1983). Material from northern USA has 
a paler peeling layer than the present material. The 
present material was found on seeds of a member of the 
Vitaceae and on a leguminous pod. According to Miller 
(1942) this fungus is common in Africa on unspecified 
seeds and fruits. Culturing of material would probably 
clarify the host range and geographical range of this 
species, but our material was not culturable. 


Xylaria pallida Berk. & Cooke, J. Linn. Soc. Bot. 15:395. 
1876. EV Gey 2cn 


Stromata clavate, conical, or subglobose bearing 
completely immersed perithecia, with rounded or acute 
sterile apices, on long narrow, smooth stipes, 1.5-3.5 cm 
total length X 3-5 mm diam; externally whitish becoming 
gray to dull black; internally white. Texture very hard. 
Surface smooth. Perithecia 0.5-1.0 mm diam. Ostioles 
finely discoid, slightly raised. Asci not seen. Asco- 
spores dark brown, unicellular, ellipsoid-inequilateral 
with broadly to narrowly rounded ends, smooth, (8.5-)9-10 
(-11) X 3.5-4.5 um, with inconspicuous straight germ slit. 


SPECIMEN EXAMINED: San Martin 340, VIII.1987, Ejido Boca 
de Chajul, Ocosingo municipality, Chiapas state, on wood 
in evergreen rain forest (ITCV). 


356 


NOTES: Our material looks much like that described by 
Dennis (1956), Perez-Silva (1975) from Mexico, and Rogers 
et al. (1988). 


Xylaria pallide-ostiolata P. Henn., Engler's Bot. Jahrb. 
38:128. 1905. Figs. 7357491545 1554 


Stromata broadly clavate with rounded fertile apices, 
with Tong stipes often extended into rooting bases, 10-11 
cm total length X 1.5-2 cm diam; externally black with 
white ostiolar regions, the stipes often with violet-brown 
tomentum; internally white. Texture cheesy to woody. Sur- 
face rugose and further roughened by perithecial contours 
and ostioles. Perithecia 0.3-0.7 mm diam. OQstioles in- 
conspicuous to hemispherical. Asci eight-spored, cylin- 
drical, poor condition in this material, the spore-bearing 
part 125-150 um long X 7-8.5 um broad, with apical ring 
bluing in Melzer's iodine reagent, rectangular, 2 wm high 
X 2.4-3.2 um broad. Ascospores almost black, unicellular, 
ellipsoid-inequilateral with abruptly pinched to acute 
ends, smooth, (12-)14-17(-18) X 6-7 wm, with straight germ 
Slit slightly less than full-length. 


SPECIMEN EXAMINED: Guevara 270, 11.1986, Jose Maria 
Morelos, Quintana Roo state, on wood (ITCV:JDR). 


NOTES: Our material is much like that described and 
depicted from African material by Lloyd (1918C), except 
that the ascospores are shorter. A photograph by J. H. 
Miller (in possession of JDR) shows ascospores as well as 
stromata. Dennis (1958) put this species into synonymy 
with X. scruposa (as Xylosphaera). Additional studies are 
required before the relationship of X. pallide-ostiolata 
and X. scruposa is clear. a 


Xylaria persicaria (Schw.: Fr.) Berk. & Curt., Grevillea 
sASu aL SbVA FigSia 45. on 


Stromata unbranched or branched, cylindrical, bearing 
crowded or scattered, more or less immersed perithecia, 
with sterile acute apices, on narrow short to long tomen- 
tose stipes, 3.5-7 cm total length X 0.5-2 mm diam; exter- 
nally blackish with remains of yellowish to brown outer 
peeling layer; internally white with brown medulla. Tex- 
ture fairly soft. Surface more or less roughened with 
perithecial contours. Perithecia 0.3-0.5 mm diam. 
Ostioles papillate. Asci eight-spored, the spores arrang- 


357 


ed uniseriately or biseriately, cylindrical, stipitate, 
127-162 um total length X 5.5-7 um broad, the spore-bearing 
part 87-100 um, with apical ring bluing in Melzer's iodine 
reagent, rectangular, 2-3 ym high X 1-2.5 um broad. Asco- 
spores brown, unicellular, ellipsoid-inequilateral to some- 
what navicular with broadly or narrowly rounded ends, 
smooth, 13.5-16 X (4.5-)5-6(-6.5) um, with long spiralling 
germ slit. 


SPECIMEN EXAMINED: Chacon-Jimenez 10, X.1987, Ejido Alta 
Cima, Gomez Farias municipality, Tamaulipas state, on 
fallen fruits of Liquidambar styraciflua L., in cloud 
forest (ITCV;UDR). 


NOTES: This seems like X. persicaria as described by 
Rogers (1979), except that the ascospores are larger than 
given in that publication. Ascospore size was erroneously 
cited therein and should have been given as 10-15 X 4-6 on 
the basis of collections examined. A more recent publica- 
tion (Rogers, 1986) gives ascospores of this species as 
10-15(-16) X 4-6 wm, based upon examination of additional 
collections. 


Xylaria phyllocharis Mont., Ann. Sci. Nat. Bot. (ser. 4) 
Salle 1055. FIGS sc A/ehGe 


Stromata cylindrical bearing fully immersed peri- 
thecia, with short acute sterile apices, on long smooth 
stipes, 0.3-3.8 cm total length X 0.5-1.5 mm diam; 
externally reddish brown with black ostiolar papillae; 
internally white. Texture soft. Surface smooth except 
for ostiolar papillae and peg-like structures that 
probably represent the remains of the anamorph located 
immediately below the fertile part. Perithecia 0.2-0.3 mm 
diam. Ostioles strongly papillate. Asci eight-spored, 
the spores arranged in a uniseriate or biseriate manner, 
cylindrical, stipitate, 103-128 um total length X 7-10 um 
broad, the spore-bearing part 70-83 um, with apical ring 
bluing in Melzer's iodine reagent, quadrate to rectangular, 
2-4 wm high X 2-3 um broad. Ascospores brown to dark 
brown, unicellular, ellipsoid-ineacuilateral with broadly 
or narrowly rounded ends, smooth, 11-14 X 5-6(-7) um, with 
hyaline sheath most evident on immature spores, with 
Straight germ slit slightly less than spore-length. 


SPECIMEN EXAMINED: San Martin 338, VIII.1987, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 


358 


on fallen leaves in median subdeciduous rain forest 
(ITCV;JDR). 


NOTES: This fungus equates well with X. phyllocharis as 
described and illustrated by Dennis (1956). Rogers et al. 
(1988) described a very similar fungus as Xylaria sp. 
(R1714) from Venezuela. That fungus differed from the 
present one only in its slightly larger ascospores, larger 
ascus tip ring, and geniculate surface. The Venezuela 
fungus is very close to the one described here and 
probably also represents X. phyllocharis. 


Xylaria poitei (Lev.) Fr., Nova Acta Regiae Soc. Sci. 
Upsaimalisence sia piel cose pico le 


Stromata unbranched or branched, clavate to irregular, 
with rounded fertile apices, on short bases or sessile, 
5.5-21.5 cm total length X 2-4 cm diam, externally whitish 
becoming brownish; internally white, sometimes becoming 
hollow. Texture hard. Surface smooth under field condi- 
tions, becoming rugose on drying; outer layer sloughing on 
old material. Perithecia 0.4-1 mm diam. Ostioles papil- 
late to hemispherical, particularly conspicuous on areas 
where outer layer has sloughed. Asci eight-spored, cylin- 
drical, long-stipitate, 198-229 um total length X 6-8 um 
broad, the spore-bearing part 98-121 um long, with apical 
ring bluing in Melzer's iodine reagent, rectangular, 3-3.5 
um high X 2.5-3 um broad. Ascospores dark brown, unicell- 
ular, ellipsoid-inequilateral with narrowed ends, smooth, 
14-17(-18) X 5.5-6.5 um, with straight germ slit less than 
spore-length. 


SPECIMENS EXAMINED: San Martin 10, 27.VII.1986, Guadalupe 
municipality, Nuevo Leon state, on Carya wood in oak 
forest (ITCV,JDR); Castillo 8020, "ET Naranjal", Ejido 
Felipe Angeles, Llera municipality, Tamaulipas state, on 
wood in a low subdeciduous rain forest (ITCV); *Ventura, 
F. 2901, 27.XI.1970, Paso de Ovejas, Veracruz state, on 
wood (JDR;XAL). 


NOTES: Our specimens seem typical for the species (see 
Rogers, 1984B; Rogers & Callan, 1986B). 


Xylaria polymorpha (Pers.: Fr.) Grev., Flora Edin., p. 
3554) 1824. Figvaaae 


359 


Stromata unbranched or branched, cylindrical to 
cylindric-clavate to spathulate or grotesque, usually with 
rounded fertile apices, but occasionally with somewhat 
acute sterile apices, with short to long stipes sometimes 
extended as rooting bases, 3-15 cm total length X 0.8-3 cm 
diam; externally blackish, sometimes with brownish peeling 
outer layer; internally white to yellowish. Texture hard. 
Surface rugulose to rugose, often with verrucae and 
ostiolar papillae. Perithecia 0.5-1 mm diam. Ostioles 
discoid to hemispherical, inconspicuous to pronounced. 
Asci eight-spored, cylindrical, long-stipitate, 197-282 um 
total length X 7-11 um broad, the spore-bearing part 133- 
182 um, with apical ring bluing in Melzer's iodine reagent, 
rectangular to urniform, 5-7 um high X 4-5 wm broad. 
Ascospores brown to dark brown, unicellular, ellipsoid- 
inequilateral to navicular with rounded to acute ends, 
sometimes pinched, smooth, (22-)23-26(-28) X 7.5-8(-9) um, 
with straight to oblique germ slit 1/2 to 2/3 spore- 
length. 


SPECIMENS EXAMINED: Garcia 1173, VII.1981, Pisaflores, 
Hidalgo state, on wood in cloud forest (ITCV) : Garcia 
2799 VAULT 1982, "Rincon de la Sierra", Juarez 
municipality, Nuevo Leon state, on wood in Quercus - 
Sargentia forest (ITCV); Garcia 4446, 1X.1984, ET Cercado, 
Santiago municipality, Nuevo Leon state, jon wood in low 
montane Quercus spp. forest ULC Vis Garcia 5347, 
14.X1.1986, Gomez Farias, Tamaulipas state, on wood in 
rain forest (ITCV); Guevara 500, II.1986, "Viejo" ranch, 
José Maria Morelos municipality, Quintana Roo state, on 
wood (ITCV ,JDR); Guevara 718, no date, Xan Santa Cruz, 
Jose Maria Morelos municipality, on wood (ITCV); San 
Martin 12, VIII.1986, Gomez Farias, Tamaulipas state, on 
wood in median subdeciduous rain forest (ITCV); San Martin 
394, 1987, Ciudad Victoria, Tamaulipas state, on wood in 
submontane scrub (ITCV). 


NOTES: Typical X. polymorpha seems to be most common in 
the northern parts of the temperate zones. In the tropics 
it is usually replaced by X. schweinitzii, a species from 
which it is often difficult to separate (Dennis, 1956; 
Rogers & Callan, 1986A) (see X. schweinitzii herein). One 
collection has subglobose fertile parts and long rooting 
stipe. Saccardo (1913) called this X. ophiopoda Sacc., 
but we consider it to be a form of X. polymorpha. Pérez- 
Silva (1975) described and illustrated X. polymorpha from 
México, but her concept of this species differs greatly 


360 


from ours. She gives ascospore dimensions as (15.3-)17 
(-18.7) X 6.8(-8.5) um, close to those of X. scruposa (see 
elsewhere herein). However, she describes and 7]Tustrates 
ascospores as having a long, straight germ slit, which is 
unlike either X. polymorpha or X. scruposa in our experi- 
ence. i 
Xylaria scabriclavula San Martin & Rogers, sp. nov. 

Figs. 80,81,134-136. 


Stromata non ramosa vel interdum aliquot exorientia 
ex eadem basi, clavata cum rotundatis vel leviter acutis 
extremis, peritheciis immersis praedita, stipitibus 
brevibus vel longis, 1 cm alta X 1.5-3 mm crassa; extus 
badia vel nigella cum atris papillis ostiolorum; intus 
eburnea. Textura mollis. Superficies scaberrima cum 
profundis fissuris et strato delapso et papillis 
ostiolorum, stipitibus saepe valde rimosis. Perithecia 
0.2-0.3 mm diam. Ostiola manifeste hemisphaerica. Asci 
saepe biseriate octospori, cylindrici, stipitati, 125-179 
um longitudine tota X 5-8 um crassi, partibus sporiferis 
67-104 um, annulo apicali in liquore Melzeri cyanescente, 
aliquantum cuneato, 0.75 um alto X 1.5 um crasso. 
Ascosporae brunneolae vel brunneae, unicellulares, 
ellipsoideo-inaequilaterales vel allantoideae, leves, 
10-12(-14) X 3.5-4.5(-5) um, rima germinativa inconspicua 
recta minus quam longituda sporae. Status anamorphosis 
ignotus. 


Stromata unbranched but several sometimes arising 
from common base, clavate with rounded or somewhat acute 
apices bearing immersed perithecia, with short or long 
Stipes, 1 cm high X 1.5-3 mm diam; externally reddish 
brown to blackish with black ostiolar papillae; internally 
cream-colored. Texture soft. Surface very rough with 
deep cracks and sloughing layer and ostiolar papillae, the 
stipes often especially cracked. Perithecia 0.2-0.3 mm 
diam. Ostioles hemispherical, prominent. Asci eight- 
Spored, the spores arranged in a uniseriate or partly 
biseriate manner, cylindrical, stipitate, 125-179 yum total 
length X 5-8 um broad, the spore-bearing part 67-104 um, 
with apical ring bluing in Melzer's iodine reagent, 
somewhat wedge-shaped, 0.75 um high X 1.5 um broad. 
Ascospores light brown to brown, unicellular, ellipsoid- 
inequilateral to allantoid, smooth, 10-12(-14) X 3.5-4.5 
(-5) um, with inconspicuous straight germ slit less than 
full-length. 


361 


SPECIMENS EXAMINED: San Martin 77, 8.XII.1986, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 
on Thrinax radiata Lodd. ex J. A. & J. H. Schult. wood in 
median subdeciduous rain forest (ITCV;JDR); San Martin 
341, VIII.1987, (same data as no. 77 above) (HOLOTYPE: 
ITCV; ISOTYPE:JDR). 


NOTES: This curious fungus has been collected twice at 
the same general location on monocot wood. It is 
noteworthy for its regular clavate stromata that are 
highly roughened when examined with a lens. It might be 
host-specific. 


Xylaria schweinitzii Berk. & Curt., J. Acad. Nat. Sci. 


Phimemcser sia) 12:284) yit853° ba GSwmocisass 


X. obovata (Berk.) Fr., Nova Acta Regiae Soc. Sci. 
Upsal. (ser. 3) 1, p. 127. 1851; fide Laessge in 
Tibt. 

Stromata subglobose to cylindrical-irregular, with 
rounded fertile apices, on abrupt short stipes or nearly 
sessile, 1.5-2.5 cm total length X 0.5-1.6 cm diam; 
externally dark brown to blackish; internally white to 
yellowish. Surface somewhat roughened by wrinkles, ver- 
rucae, and ostiolar papillae. Perithecia 0.3-0.9 mm diam. 
Ostioles discoid to hemispherical. Asci eight-spored, 
cylindrical, stipitate, 210-229 ym total length X 7-9 um 
broad, the spore-bearing part 140-145 um long, with apical 
ring bluing in Melzer's iodine reagent, rectangular, 
5.5-7(-7.5) um high X 3.5-5.5(-6) um broad. Ascospores 
brown to dark brown, unicellular, ellipsoid-inequilateral 
to navicular with rounded to acute ends, smooth, 21-26 X 
(6-)6.5-8 um, with oblique germ slits less than spore- 
length. 


SPECIMEN EXAMINED: Garcia 3059, Sieh eoe, eLOSIUXtlds 
Experiment Station, Balzapote, Veracruz state, on wood in 
evergreen rain forest (ITCV;JDR). 


NOTES: OQur concept of X. schweinitzii is the traditional 
one set forth by Dennis (1956), Carroll (1964), 
Perez-Silva (1975), Rogers & Callan (1986A), Lloyd (1919, 
1920), Miller (1934), and others. In our view X. 
schweinitzii is a tropical variant of X. polymorpha that 
usually exhibits a smoother surface and has ascospore germ 
slits that are less than spore-length and oriented oblique 
to the long axis of the spore. Our concept of X. obovata 


362 


(Berk.) Fr. is as a member of the X. polymorpha complex 
with a much smoother (often unwrinkled) surface (Rogers et 
al., 1988). 


Xylaria scruposa (Fr.) Fr., Nova Acta Regiae Soc. Sci. 
Upsal. (ser. 3) 1, p. 127.- 1851; sensu Dennis, Kew 
Bull. 919563436; 1956. Figs. 84,87. 


Stromata unbranched or branched, cylindrical, clavate, 
or irregular, with rounded fertile apices or somewhat 
acute sterile apices, on short to long stipes, 1.5-6 cm 
total length X 2-6 mm diam; externally dark brown to 
black, sometimes with brown scales; internally white to 
yellowish. Texture woody. Surface rugose, sometimes 
additionally roughened with tomentum or ostiolar papillae. 
Perithecia 0.5-1 mm diam. Ostioles inconspicuous to hemi- 
spherical. Asci eight-spored, cylindrical, stipitate, 
188-224 um total length X 7.5-8 um broad, the spore- 
bearing part 100-146 um, with apical ring bluing in 
Melzer's iodine reagent, rectangular, 5-6 um high X 3.5-5 
um broad. Ascospores brown to dark brown, unicellular, 
ellipsoid-inequilateral to navicular with rounded to acute 
ends, smooth, 18-21(-22) X 6-7 wm, with oblique to spiral- 
ling germ slit less than spore-length. 


SPECIMENS EXAMINED: *Brown, D. 14, 28.1X.1981, Parque 
Foo. Javier Clavijero near Xalapa-Coatepec, on wood . 
(JDR;XAL); *Magana 19, Michoacan, (JDR;XAL); San Martin 
251, VIII.1987; "El Muneco", Ejido Loma Bonita, @cosingo 
municipality, Chiapas state, on wood in evergreen rain 
forest (ITCV;JDR); San Martin 252, 365, 369, 373, 
VIII.1987, Ejido Boca de Chajul, Qcosingo municipality, 
Chiapas state, on wood in evergreen rain forest (ITCV; 252 
and 369 also in JDR). 7 


NOTES: Xylaria scruposa is a complex taxon, a small- 
spored member of the X. polymorpha complex. Its limits 
are unclear (see Rogers et al., 1988 and refs. therein for 
discussions of this taxon). 


Xylaria sp. (SM 38,64) Figs? 91937 


Stromata unbranched to branched, cylindrical to 
cylindric-clavate with acute to obtuse, sterile or fertile 
apices, on short to long tomentose stipes that arise from 
pannose bases, 1-5 cm total length X 1-3 mm diam; exter- 
nally blackish with brown peeling outer layer; internally 


363 


white. Texture soft. Surface rugose and roughened with 
ostiolar papillae. Perithecia 0.2-0.3 mm diam. Ostioles 
papillate, conspicuous and crowded. Asci eight-spored, 
cylindrical, long-stipitate, 90-125 yum total length X 5-6 
um broad, the spore-bearing part 60-65 um, with apical 
ring bluing in Melzer's iodine reagent, somewhat discoid, 
0.75 um high X 1.5 um broad. Ascospores brown to dark 
brown, ellipsoid-inequilateral with one end sometimes 
bearing a tiny cellular appendage, smooth, 8-9(-10) xX 
4.5-5 um, with straight germ slit spore-length. 


SUEGIMENSMEAAMINED:| *Basilio. M253. 20.X1.1979,° E) 
Teposteco, Morelos, on wood (JDR;XAL); San Martin 38, 
10.1X.1987, Gomez Farias, Tamaulipas state, on wood in 
median subdeciduous rain forest (ITCV;JDR); San Martin 64, 
7.X1I.1986, San Felipe Bacalar, Othon P. Blanco municipal- 
i.tye Quintana Roo state, on wood in acahual (ITCV;JDR). 


NOTES: This is another dark, small-spored Xylaria that 
does not fit into any species known by us. It possibly is 
X. aemulans Starback as understood by Lloyd (1921). We 
have not seen type material of that species. 


Xylaria sp. (SM 39, 306) Figs. 90,92. 


Stromata unbranched or branched, cylindrical, bearing 
completely immersed perithecia, with attenuated or acute 
sterile apices, on short stipes often arising from pannose 
bases, 1-2.5 cm total length X 2-4 mm diam; externally 
blackish with reddish brown to dark brown peeling outer 
layer; internally dirty white. Texture hard. Surface 
smooth. Perithecia 0.5-1 mm diam. Ostioles barely raised, 
located in flattened circular areas ca. 0.2 mm diam. Asci 
eight-spored, cylindrical, very long stipitate, 140-200 um 
total length X 5-6 um broad, the spore-bearing part /0-80 
um, with apical ring bluing in Melzer's iodine reagent, 
quadrate, 2 X 2 ym. Ascospores brown, unicellular, 
ellipsoid-inequilateral with broadly or narrowly rounded 
ends, smooth, (9-) 9.5-11(-12) X 4(-4.5) um, with straight 
germ slit spore-length. 


SPECIMENS EXAMINED: San Martin BOweS Nl) OO seit do" La 
Union, Othon P. Blanco municipality, Quintana Roo State, 
on wood in median subdeciduous rain forest (ITCV;JDR); San 
Martin 306, VIII.1987 (same data as no. 39) on soil 
(ITCV). 


364 


NOTES: This fungus might only be a variant of X. 
multiplex, differing from more typical material in having 
ostioles often surrounded by a disc and in lacking any 
perithecial contours. In surface features it resembles X. 
adscendens. 


Xylaria sp. (SM 211) Fig. 94. 


Stromata composed of cylindrical to conical aggrega- 
tions of more or less naked perithecia, with acute sterile 
apices, on filiform glabrous stipes, 5-13 mm total length 
X 0.5-1.5 mm diam; externally black; internally white. 
Texture fairly hard. Surface smooth except for perithecial 
elevations. Perithecia 0.2-0.3 mm diam. OQstioles papil- 
late. Asci eight-spored, cylindrical, stipitate, 73-83 
total length X 4-5 um broad, the spore-bearing part 47-50 
ym, with apical ring bluing in Melzer's iodine reagent, 
quadrate, 1-1.3 X 1-1.3 wm. Ascospores light brown to 
brown, unicellular, ellipsoid-inequilaterial with broadly 
or narrowly rounded ends, smooth, (5.5-)6-7 X 3-3.5(-4) 
um, with straight germ slit slightly less than spore- 
length. 


SPEGIMEN EXAMINED: San Martin 211, 16.VII.1987, G6mez 
Farfas, Tamaulipas state, on fallen leaves in median 
subdeciduous rain forest (ITCV). 


NOTES: This tiny Xylaria might well represent a new 
Species. Our collection is so small, however, that we 
decline to describe it as new. 


Xylaria sp. (SM 303) Figs. 95,96. 


Stromata solitary or many originating from common 
base, cylindric-clavate to irregular with subacute to 
narrowly rounded sterile apices, on short stipes, 1.3-4 cm 
total length X 3-6 mm diam; externally black, sometimes 
with brown tones; internally white to yellowish. Texture 
cheesy. Surface so deeply cracked as to appear verrucose, 
with tomentum on stipe and in patches on fertile part. 
Perithecia 0.3-0.5 mm diam. Ostioles hemispherical, often 
flattened, difficult to distinguish from verrucae. Asci 
cylindrical, stipitate, 148-166 um total length X 5.5-7 um 
broad, the spore-bearing part 80-90 um, with apical ring 
bluing in Melzer's iodine reagent, rectangular, 3-4.5 um 
high X 2-3 ym broad. Ascospores brown to dark brown, 
unicellular, ellipsoid-inequilateral with narrow ends, 


365 


smooth, (12-)12.5-14.5(-16) X 5-5.5(-6) um, with straight 
germ slit less than spore-length. 


SPECIMEN EXAMINED: San Martin 303, 31.VIII.1987, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 
on wood of Thrinax radiata Lodd. ex J. A. & J. H. Schult. 
in median subdeciduous rain forest (ITCV). 


NOTES: This fungus is probably another small-spored 
member of the X. polymorpha complex. It seems allied to 
X. feejeensis and X. Tuteostromata Lloyd. Most of our 
material 1S immature and even material showing fully 
colored ascospores is probably not entirely mature. We 
thus refrain from describing it as a new taxon, although 
it might well be such. 


Xylaria sp. (SM 331) Figs. 99,100. 


Stromata unbranched or branched, cylindrical to 
irregular with acute sterile apices, on abrupt thin short 
to long stipes, 1.5-3.5 cm total length X 2-3 mm diam; 
externally black without trace of peeling layer; 
internally yellowish. Texture soft to woody. Surface 
smooth except for perithecial undulations. Perithecia 
0.5-0.8 mm diam. Ostioles conical, inconspicuous. Asci 
not seen. Ascospores brown, unicellular, ellipsoid- 
jinequilateral with rounded ends, smooth, 9-10.5(-11) X 4-5 
um, with straight germ slit less than spore-length. 


SPECIMEN EXAMINED: San Martin 331, VIII.1987, Ejido Boca 
de Chajul, Ocosingo municipality, Chiapas state, on wood 
in evergreen rain forest (ITCV;JDR). 


NOTES: We do not know this fungus. Our material is 
overmature and asci are altogether lacking. 


Xylaria sp. (SM 337) PiGSem oe Loe l4s 


Stromata consisting of head of immersed perithecia 
1.5 mm diam on tomentose hair-like rachis that extends 
downward as a stipe and upward as a long apex, up to 2.3 
cm long; externally blackish with white flaking outer 
layer; internally white. Texture soft. Surface smooth 
except for ostiolar papillae. Perithecia 0.2-0.3 mm diam. 
Q@stioles papillate to almost digitate. Asci eight-spored, 
the spores arranged in a uniseriate to partly biseriate 
manner, cylindrical, long-stipitate, 117-154 um total 


366 


length X 6-10 um broad, the spore-bearing part 67-76 um, 
with apical ring bluing in Melzer's iodine reagent, 
quadrate to rectangular, 3-4 um high X 2-3 um broad. 
Ascospores brown to dark brown, ellipsoid-inequi lateral 
with one end bearing a cellular hyaline appendage ca. 2 um 
X 2 um, smooth, 10-12(-14) X 5-6 um, with straight germ 
slit spore-length. 


SPECIMEN EXAMINED: San Martin 337, VIII.1987, San Felipe 
Bacalar, Othon P. Blanco municipality, Quintana Roo state, 
on fallen leaves (ITCV). 


NOTES: The present fungus greatly resembles X. sicula 
Pass. and Beltr. f. major Ciccarone in the hair-Tike 
rachis with the intercalary fertile portion and size of 
ascospores (Ciccarone, 1946). It differs in having 
immersed perithecia, a tomentose rachis, and ascospores 
with a cellular appendage. This latter feature is easily 
missed unless critical microscopy is employed. It greatly 
resembles X. aristata sensu Dennis (1956) in the morphology 
of the fertile part and in the hairy stipe, but differs 
from his description in having a portion of the rachis 
extending beyond the fertile head and in having appendaged 
ascospores. It resembles X. aristata described herein, 
but differs in larger appendaged ascospores, in larger 
asci and in having a portion of the rachis extending 
beyond the fertile part. Dargan (1976) illustrated X. 
aristata as having a short mucro on the fertile portion. 
He described perithecia as being completely immersed, but 
illustrated them as being fairly conspicuous, i.e., as 
surface contours. Miller (1942) described X. aristata as 
having glabrous stipes and protruding perithecia. Our 
fungus also greatly resembles Dargan's Xylaria sp.-III in 
ascospore size and in the long portion of rachis project- 
ing beyond the fertile head (Dargan, 1976). In any case, 
it is probably most prudent to allow our present small 
collection to remain unnamed until additional studies of 
Similar fungi are undertaken. 


Xylaria sp. (SM 359) Figs. 97,98. 


Stromata cylindrical with fertile apices, on short 
pubescent stipes, prostrate in our material, 7-10 mm total 
length X 1-3 mm diam; externally blackish with brown outer 
peeling layer, with ostioles often white-fringed; inter- 
nally white. Texture soft. Surface with 
circumferentially-oriented wrinkles and roughened with 


367 


ostiolar papillae. Perithecia 0.4-0.7 mm diam. Ostioles 
strongly conical. Asci eight-spored, the spores arranged 
in a uniseriate to biseriate manner, cylindrical, 
stipitate, 132-148 um total length X 5.5-7 um broad, the 
spore-bearing part 82-88 um, with apical ring bluing in 
Melzer's iodine reagent, quadrate, 2 X 2 um. Ascospores 
brown, unicellular, ellipsoid- inequilateral, with broadly 
to narrowly rounded ends, smooth, (11-)13-14 (-15) X 4-5 
um, with germ slit obscure. 


SPECIMEN EXAMINED: San Martin 359, VIII.1987, Ejido Boca 
de Chajul, Ocosingo municipality, Chiapas state, on wood 
in evergreen rain forest (ITCV;JDR). 


NOTES: Our fungus highly resembles Lloyd's photograph 
(1921) of X. biformis Lloyd, but his description is of a 
fungus with much Targer ascospores. Laessge (in litt. bes 
considers X. Ba eatin to be a synonym of X. globosa (= X 
anisopleura). Our fungus also resembles a diminutive X. 
berkeleyi. In any case, we do not assign it to a known 
taxon herein. 


Xylaria sp. (SM 372) Figs melO lei 022 


Stromata cylindrical to cylindric-clavate with 
rounded fertile apex, on abruptly narrowed long stipe, 
1.5-2 cm total length X 1-2 mm diam; externally blackish 
with persistent brown outer sloughing layer; internally 
white. Texture hard, but fragile. Surface roughened by 
sloughing layer and perithecial undulations. Perithecia 
0.2-0.6 mm diam. Ostioles papillate. Asci eight-spored, 
cylindrical, poor condition in this material, the spore- 
bearing part ca. 50-60 um long X 5-6 um broad, with apical 
ring bluing in Melzer's iodine reagent, quadrate, 2 X 2 
um. Ascospores brown to dark brown, unicellular, 
ellipsoid-inequilateral with rounded ends, smooth, 9-10 
(-11) X (4-)4.5-5 um, without obvious germ slit. 


SPECIMEN EXAMINED: San Martin 372%, WA 1 987 sy EJ Vdoe Boca 
de Chajul, O@cosingo municipality, Chiapas state, on wood 
in evergreen rain forest (ITCV). 


NOTES: This fungus is not known to us and probably 
represents a new species. Perithecia of our material, 
however, were not in good condition and asci could not be 
fully characterized. 


368 


Xylaria sp. (G729) Figs. 85,86. 


Stromata cylindrical with acute sterile apex, without 
differentiated stipe, the base tomentose, 1.5-3 cm total 
length X 1.5-3 mm diam; externally blackish with brownish 
outer peeling layer; internally whitish. Texture hard. 
Surface smooth except for ostiolar papillae and remains of 
tomentum. Perithecia 0.3-0.4 mm diam. Ostioles conical 
to hemispherical, conspicuous. Asci eight-spored, cylin- 
drical, long-stipitate, 123-144 um total length X 4-6 um 
broad, the spore-bearing part 63-70 um, with apical ring 
bluing in Melzer's iodine reagent, quadrate to rectangular, 
1.5-2.5 wm high X 2 um broad. Ascospores light brown, 
unicellular, ellipsoid-inequilateraT with rounded ends, 
smooth, (8.5-)9-11 X (3-)3.5-4(-4.5) um, with straight 
germ slit less than spore-length. 


SPECIMEN EXAMINED: Guevara 729, 1986, Xan Santa Cruz, 
Jose Maria Morelos municipality, Quintana Roo state, on 
wood in an acahual (ITCV;JDR). 


NOTES: This small Xylaria is unknown to us. In general 
morphology it resembles X. myosurus in lacking a real 
Stipe, but the tomentose base 1s unlike that species. 


Xylaria squamulosa San Martin & Rogers, sp. nov. 
Figs. 103,104,112 idee 


Stromata subglobosa vel elongata, 5 mm diam vel 6 mm 
longa X 4 mm crassa, peritheciis omnino immersis, in 
abrupte angustatis stipitibus 1-1.5 cm longis X 1-1.5 mm 
diam; extus fumea sed sub lente nigella cum elevatis 
albidis squamis; intus alba. Textura dura. Superficies 
levis praeter fissuras inter squamas. Perithecia ca. 1 mm 
diam. Ostiola plus minusve umbilicata, inconspicua. Asci 
saepe biseriate ostospori, cylindrici, longe stipitati, 
370-430 longitudine tota X 14-27 um crassi, partibus 
Sporiferis 182-252 um, annulo apicali in liquore Melzeri 
cyanescente, urceolato vel obtrullato, 11-13 alto X 6-7 um 
crasso. Ascosporae brunneae, unicellulares, ellipsoideo- 
inequilaterales vel plus minusve rectangulatae cum late 
rotundatis extremis vel abrupte acutis extremis, leves, 
(29-)31-37(-40) X (11-)12-13(-14) um, saepe ab hyalina 
vagina indutae, rima germinativa recta per longitudinem 
Sporae. Anamorphosis ignotus. 


369 


Stromata subglobose to elongated, 5 mm diam to 6 mm 
long X 4 mm diam, with completely immersed perithecia, on 
abruptly narrowed stipes 1-1.5 cm long X 1-1.5 mm diam; 
externally blackish gray, but with lens seen to be black- 
ish with raised whitish scales; internally white. Texture _ 
hard. Surface smooth except for cracks between scales. 
Perithecia ca. 1 mm diam. Ostioles more or less umbili- 
cate, inconspicuous. Asci eight-spored, the spores 
oriented in a uniseriate or partially biseriate manner, 
long-stipitate, 370-430 um total length X 14-27 um broad, 
the spore-bearing part 182-252 um, with apical ring bluing 
in Melzer's iodine reagent, urn-shaped or coffin-shaped, 
11-13 um high X 6-7 um broad. Ascospores dark brown, uni- 
cellular, ellipsoid-inequilateral to more or less rec- 
tangular with broadly rounded or abruptly acute ends, 
smooth, (29-)31-37(-40) X (11-)12-13(-14) um, often with 
hyaline sheath, with straight germ slit spore-length. 
Anamorph unknown 


SPECIMEN EXAMINED: San Martin 63, 8.XII.1986, Ejido La 
Union, Othon P. Blanco municipality, Quintana Roo state, 
on wood in median subdeciduous forest (HOLOTYPE: ITCV). 


NOTES: This fungus appears to be an undescribed species. 
It is probably the same as Xylaria sp. (S 1836) described 
by Rogers et al., 1988). The fertile part of that fungus 
lacked the pronounced scaliness of the one described here, 
but shows conspicuous papillate ostioles. In other 
characteristics these fungi seem very similar. 


Xylaria telfairii (Berk.) Fr., Nova Acta Regiae Socescas 
oraersen: 23iiel fob 2ne ibot. 


Stromata clavate-fusiform with rounded fertile apices, 
on short to Tong stipes, up to 6.5 cm total length X 4-9 
mm diam; externally tan or yellowish to dull orange, often. 
black from spores; internally white, but becoming hollow 
and inrolled, often before maturation of perithecia. 
Texture hard, but fragile. Surface smooth. Perithecia 
0.4-0.7 mm diam. Ostioles umbilicate. Asci not observed. 
Ascospores light brown to brown, unicellular, ellipsoid- 
~Inequilateral with narrowed ends, smooth, (17-)18-21(-22) 
X (5.5-)6-7 um, with short, oblique germ slit. 


SPECIMENS EXAMINED: San Martin 220, VIII.1987; 352, VIII. 
1987, Ejido Boca de Chajul, Ocosingo municipality, Chiapas 
state, on wood in evergreen rain forest (ITCV,JDR). 


370 


NOTES: Qur material seems typical for the species. It 

has been described and illustrated from Mexican material 
by Perez-Silva (1975). See remarks by Dennis (1956) and 
Rogers et al. (1988). 


Xylaria uniapiculata San Martin and Rogers, sp. nov. ~- 
Figs. 105-107,124-126. 


Stromata plerumque maxime ramosa e basi vel super 
basim, plus minusve cylindracea vel teretia vel com- 
planata, plerumque apicibus acutis sterilibus, in 
stipitibus brevibus e basibus pannosis, 2.5-5.5 cm longa X 
2-6 mm crassa; extus rufa vel brunnea vel nigella cum 
strato externo fusco delapso; intus alba vel flavida. 
Textura lignosa. Superficies valde asperata ab verrucis 
prominentibus, fissuris, papillis ostiolorum, et formis 
peritheciorum. Perithecia 0.4-0.8 mm diam. Ostiola 
papillata. Asci octospori, cylindrici, stipitati, 115-138 
longitudine tota X 5-6 um crassi, partibus sporiferis 
70-90 um, ariulo apicali in liquore Melzeri cyanescente, 
quadrato vel oblongo, 2.5-3 um alto X 1.5-2 um crasso. 
Ascosporae brunneae, plerumque in uno extremo cellula 
minuta hyalina ornati, ellipsoidea-inequilaterales, leves, 
9-10(-11) X (3.5-)4 um, rima germinativa recta per 
longitudinem partis coloris sporae. Anamorphosis ignotus. 


Stromata usually highly branched from base or above, 
more or less cylindrical, terete or flattened, usually 
with somewhat acute sterile apices, on short stipes from 
pannose bases, 2.5-5.5 cm high X 2-6 mm diam; externally 
reddish brown, dark brown to black with peeling outer 
brown layer; internally white to yellowish. Texture 
woody. Surface strongly roughened with prominent ver- 
rucae, cracks, ostiolar papillae, and perithecia] 
contours. Perithecia 0.4-0.8 mm diam. Ostioles 
papillate. Asci eight-spored, cylindrical, stipitate, 
115-138 um total length X 5-6 ym broad, the spore-bearing 
part 70-90 um, with apical ring bluing in Melzer's iodine 
reagent, quadrate to rectangular, 2.5-3 wm high X 1.5-2 um 
broad. Ascospores brown, usually with a minute hyaline 
cell on one end, ellipsoid-inequilateral, smooth, 
9-10(-11) X (3.5-)4 um, with straight germ slit running 
the full length of the pigmented cell of the spore. 


SPECIMENS EXAMINED: Chacon 191, X.1987, Gomez Farias, 
Tamaulipas state, on wood in median subdeciduous rain 


371 


forest (HOLOTYPE: ITCV; ISOTYPE:JDR); Rodriguez 31 and 
Santos Lopez 4, same data as above (ITCV;JDR). 


NOTES: This species is probably related to X. feejeensis 
and X. rhytidophloea Mont. According to Dennis (1956) the 
latter name is a synonym of the former. Lloyd (1924A) 
gives the ascospores as 18 X 8 um, much larger than those 
of X. feejeensis or the present fungus. The main feature 
leading us to consider this a new species is the 
persistent cellular ascospore appendage. 


ACKNOWLEDGMENTS 


PPNS No. 0022, Department of Plant Pathology, Project 
1767, Washington State University, Agricultural Research 
Center, Pullman, WA, 99164. This study was supported in 
part by National Science Foundation Grant BSR-8710005 to 
JDR and by grants from USAID and the National Council of 
the Technological Educational System of Mexico to FSMG. 

We thank Michael Jd. Adams, Ruby Latham, and Brenda E. 
Callan, Washington State University, for aid with photo- 
graphy, for typing the manuscript, and for reading the 
manuscript, respectively. We thank Ruben Duran, 
Washington State University, for his friendship, 
encouragement, and translations from Spanish to English. 
We thank Jose Castillo Tovar and Gaston Guzman for 
encouragement and the latter also for specimens. We thank 
Lucrecia Garcia, Jesus Garcia, Efren Cazares, Gonzalo 
Guevara, Arnulfo Moreno, and Santiago Chacon-Jimenez for 
their aid with collecting. We thank Maria Concepcion 
Herrera Monsivais for her aid in identifying plant 
material. We thank Dr. Alberto Rodriguez-Fernandez for 
providing collecting facilities in Quintana Roo and 
Francisco Quinto-Adrian for aid in Quintana Roo. We thank 
Jorge Ayala-Guajardo for his friendship and aid in the 
rain forests of Chiapas and Javier Chavelas-Polito for 
collecting facilities and aid near the Belize border. 


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ae 


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MY COTAXON 


Vol. XXXIV, No. 2, pp. 375-379 January-March 1989 


VARICOSPORIUM SCOPARIUM, A NEW STAUROSPOROUS HYPHOMYCETE 


by 


A. ROLDAN & M. HONRUBIA. Depto. Biologia Vegetal (Botanica). Facultad 
de Biologia. Universidad de Murcia (Spain). 


Summary: Varicosporium scoparium sp. nov. is described in pure cultu- 
re from foam samples in Spanish streams. 


Conidia of the new species described below were illustrated 
by ROLDAN et al. (1988) from foam samples in the River Mundo (Albacete 
Spain). They have also been detected in several of its tributaries 
and in the River Vinalopo (Alicante, Spain). Its substrate remains 
unknown. Conidia only appear ater rains, and thus we believe it 
should be considered waterborne rather than strictly aquatic. 


Varicosporium scoparium sp. nov. (Figs. 1-3) 

Hyphomycetes Moniliaceae. COLONIAE (1% MA) pallide cremosae, patentes 
ad 4.5 cm diam/5 septimanis, glabrae, mycelio aerio absente, hyphae 
1.5-2 ym latae. SPORULATIO copiosa post 5 diebus (14-18°), summa 
aqua vel submersa. CONIDIOPHORA mononematosa, apicalia vel lateralia, 
semimacronematosa, simplicia vel bracchiata, usque ad 300 x 2-3 
ym. CELLULAE CONIDIOGENAE apicales (vel intercalares vel laterales), 
8-16 x 2-3 ym, monoblasticae vel polyblasticae, proliferationes 
sympodiales; cicatrices denticulatae. CONIDIA solitaria vel  laxe 
fasciculata, acrogena, typice in planibus variis, elementi cylindrici; 
axis principalis leve curvatus vel sigmoideus, 80-150 (raro ad 
350) x 2-3.5 ym, usque ad 6-septatus, cicatrix truncata; bracchia 
in 1-3 dispositionibus, lateralia, singularia, principale unilateralia 
recta vel tenue curvata 0-3 septata, insertio typice sinuosa et 
inaequaliter constricta. Disjunctio conidiorum schizolytica. Germina— 
tio in aqua et in agaro cum antibioticis. 


COLONY (1% MA) pale cream-coloured, reaching 4-5 cm diam/S weeks 
at 14-18°, glabrous, aerial mycelium wanting, hyphae 1.5-2 ym 
wide. SPORULATION on slices of culture partly submerged in sterile, 
distilled standing water for 5 days at room temperature (14-18), at 

or below water level, producing numerous highly branched conidia. 
CONIDIOPHORES mononematous, apical or lateral, semtmacronematous, 
Simple or branched, up to 300 x 2-3 ym. CONIDIOGENOUS CELLS apical 
(or intercalary or lateral), 8-16 x 2-3 \m, mono- or polyblastic, 
proliferating sympodially; secession scars denticulate. CONIDIA 
solitary or loosely fasciculate, acrogenous, typically branched 
in more than one plane, elements cylindrical, apices rounded; 
main axis slightly curved (or somewhat sinuate), sometimes with 
loss of apical domtnance, 80-150 (rarely. up to 350) x 2-3.5 wm, 


376 


Fig. 1.— Varicosporium scoparium in pure culture. A-D: conidiophores. 
E-H: conidia. (from holotype). A: scale a = 50 ym. B-H: scale b = 40 


ym. 


378 


Fig. 3.- Varicosporium scoparium. E: conidiophore. All to same scale, 


Dance oU sun: 


O72 


up to 6-septate, scar truncate; branches in 1-3 orders, lateral, 
Single, diverging in various planes and mostly on one side of 
the parent element, straight or slightly curved, characteristically 
Sinuous and asymetrically constricted at the insertion, 0-3 septate. 
Conidia germinate readily in liquefied foam or on isolation media 
Supplemented with streptomycin and penicillin (Descals et al. 
1977). 


COLLECTIONS: single conidia isolate from foam in River Mundo, 
Albacete (UTM WH 9457) (Roldan, Nov. 1987) (MA-fungi 20841 holotype 
ex MUB AR9851 isotype); other isolates from locus typici MUB AR 
9852, MUB AR9853; stream foam from River Mundo, Albacete (Roldan, 
Nov. 1985, A-25); stream foam from River Vinalopd, Alicante (UTM 
WHeocse) (Roldan, Nov. 1985, A-77). 


There are various staurosporous genera of aquatic hyphomycetes 
with conidia branching in more than one order and plane, such 
as Dendrospora Ingold, Polycladium Ingold and Varicosporium Kegel. 
In Dendrospora conidia are apical and solitary with conidiogenous 
cells sometimes proliferating percurrently (DESCALS & WEBSTER, 
1980). Furthermore, D. erecta Ingold (type species) displays vertici- 
llate branching, not seen in our species, although this character 
is less evident in D. polymorpha Roldan & Descals (ROLDAN et al., 
1987). Conidial insertion in Polycladium equiseti Ingold (1959) 
(type and only species) is clearly lateral. Varicosorium scoparium 
appears to be closely related to V. elodeae Kegel CINGOLD, 1942) 
(type species) but here conidia bear broadly divergent branches 
and secede by abstriction at a narrowed insertion. Conidia of V. 
delicatum (Iqbal, 1971) are typically larger than in V. scoparium 
and are produced singly and terminally. Cladoconidium articulatum 
Bandoni & Tubaki (1985) has repeatedly branched pale fuscous conidia, 
with somewhat sinuous constricted branched insertion. However, 
this species has olivaceous colonies and the conidia are mostly 
spanning 30-42, rarely up to 60 ym. 


REFERENCES 
BANDONI, R.J. & TUBAKI, K. (1985). Cladoconidium: a new hyphomycete 


anamorph-genus. Trans. Mycol. Soc. Japan, 26: 425-431. 
DESCAES, E.* & WEBSTER, J. (1980). Taxonomic studies on aquatic 


hyphomycetes. II. The Dendrospora aggregate. Trans. Br. mycol. 
DOG«we 4: (135-158. 

dees, bbs, WEBSTER, J. “& DYKO;° B.J.- (1977). Taxonomic studies 
on aquatic hyphomycetes. I. Lemonniera de Wildeman. Trans. Br. 
mycol. Soc., 69: 89-109. . 
INGOLD, C.T. (1942). Aquatic hyphomycetes of decaying alder leaves. 
mans-eors mycol. Soc.,:25: 339-417. 

PGORD SE C5560 1959)- sPolycladium—equiseti- gen. nove. sp. cnov.. an 
aquatic hyphomycete on Equisetum fluviatile. Trans. Br. mycol. 
Soc., 42: 112-114. 

IQBAL, S.H. (1971). New aquatic hyphomycetes. Trans. Br. mycol. 
Soc., 56: 343-352. 

ROLDAN, A., DESCALS; E. & HONRUBIA, M. (1987). Dendrospora polymorpha 
a new aquatic hyphomycete from Spanish streams. Mycotaxon, 29:21-2/7. 
ROLDAN, A., DESCALS, E. & HONRUBIA, M. (1988). Hifomicetos acuaticos 
en las cuencas altas de los rios Segura y Guadalquivir. Anal. 
Biologia, 12 (Biologia Vegetal, 3): in press. 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 381-385 January-March 1989 


A NEW ADDITION TO THE GENUS GORGOMYCES 


by 


A. ROLDAN, Depto. Biologia Vegetal (Botanica). Facultad de Biologia. 
Universidad de Murcia (Spain). 


Summary: Gorgomyces honrubiae sp. nov. is described in pure culture 
from foam samples in Spanish streams. Characters related with 
conidiogenesis in this genus are discussed. 


Gorgomyces honrubiae sp. nov. (Figs. 1-3) 


Hyphomycetes Moniliaceae. Coloniae (1% MA), albidae, patentes 
ad 3 cm diam/5 septimani, glabrae, mycelio aerio absente; hyphae 
Saepe cincinnatae, 1-2 ym latae. Sporulatio submersa et = sparsa. 
Conidiophora semimacronematosa, mononematosa, typice’ Jlateralia, 
cum stipite et capite conidiogeno, stipes irregulariter clavatus 
et curvatus, 5-8 x 2-3.5 ym, typice O-septatus; caput conidiogenum 
est racemus condensus, eodem tempore fructificans. Cellulae conidio- 
genae inflatae, 3-6 x 1.5-2.5 ym, monoblasticae, determinatae, 
collabentes. Conidia solitaria, apicalia (vel lateralia), filiformia; 
cum cellula basali et appendice apicale; corpus conidialis parce 
Subulatus, paene rectus vel spiralis vel sigmoideus, 35-45 x ca. 
2 ym, 8-9 septatus; appendix apicalis ad basim constricta et septata, 
9-16 ym longa, unicelularis, collabens ante secessionem conidii; 
cellula basalis ca. 2 x 1.5 ym, cum mucrone laterale breve, postea 
collabens et mucosa. Conidia in aqua singillatim vel aggregatim 
dispersa, ad basim adherentia. Germinatio e pluribus cellulis. 


Etym: named in acknowledgement to Dr. M. Honrubia (University 
of Murcia, Spain). 


Hyphomycetes, Moniliaceae. Colonies (1% MA) whitish, spreading 
Slowly, reaching 3 cm diam/5 weeks, glabrous, aerial mycelium 
absent; hyphae strongly coiled, 1-2 ym wide. Sporulation underwater, 
Sparse. Conidiophores semimacronematous, mononematous, typically 
lateral, with a stipe and an irregularly branched conidiogenous 
head, total length 12-22 ym; stipe irregularly clavate and curved, 
5-8 x 2-3.5 ym, typically O-septate; Conidiogenous cells variously 
Swollen, 3-6 x 1.5-2.5 ym, monoblastic, determinated, evacuated 
after conidial secession. Conidia solitary, apical cor lateral), 
consisting (initially) of a filiform body, an apical walled appendage 
- and a stalk cell. Conidial body slightly subulate, almost straight 
to helical (1-1.5 coils) or sigmoid, 35-45 x ca. 2 ym, septa initia- 
lly distant, later 8-9 by secondary septation; apical appendage 
constricted at basal septum, 9-16 um long, unicellular, becoming 
evacuated and collapsing prior to secession; stalk cell ca. 2 


382 


Fig. 1.- Gorgomyces honrubiae in pure culture (from holotype). 
A-G: conidiophores and developping conidia. Notice collapsed conidio- 
genous cells after conidial secession in B, F and G. Bar: 20 ym. 


Fig. 2.- Gorgomyces honrubiae in pure culture (from holotype). 
4: conidial group adhering to each other at their mucous base. 
B-J: single detached conidia. Bar: 20 ym. 


383 


384 


Fig. 3.- Gorgomyces honrubiae in pure culture (lactofuchsin mounts). 
A-B: mature conidial groups adhering to each other at their apical 
appendages (A) and base (B). C: conidiophore, arrows shows a constric-— 
tion process on the conidium body and the basal cell. D-E: Detached 
conidia with appendages being evacuated. Bar: 20 um. 


385 


x 1.5 ym with a short lateral peg, contents of the cell gelatinizing 
at a later stage and thus releasing the conidia. Conidia dispersed 
in water or trapped in foam individually or more typically in 
groups, adhering to each other at their mucous base or apical 
appendages; germinating readily from any cell on the main body. 


COLLECTIONS EXAMINED 


Single groups of conidia isolated from foam in Campamento San 
Juan stream, Ridpar, Albacete, Spain. Roldan Nov. 1987 (MA-FUNGI 
20840 holotype ex MUB AR 9761 Isotype. Other collections from 
the same locality and date (MUB AR 9765, MUB AR 9766). 


Gorgomyces hungaricus Gonczol & Révay (1985), type species, was 
only described from nature, and hence some important characters 
related with conidiogenesis may not have been seen. Unfortunately, 
according to Gonczol (in litt.) the type is damaged and unservicea- 
ble. Nevertheless conidial morphology is so characteristic that 
the assignation of our species to Gorgomyces is_ unquestionable. 
G. honrubiae is clearly distinct from the type species in that, 
in the latter, the conidiogenous head is a terminal verticil composed 
of only conidiogenous cells, while here there is an incurving 
branching system; secondly, there is no evidence of a basal appendage 
of a cellular nature on the conidium; thirdly, the apical appendage, 
altnought it appears to have a wall, is described as mucous, and 
develops by apical growth, while in our species the apical appendage 
is initiated by a localized constriction process on the fully 
developed cylindrical body of the conidium, followed by a collapse 
Greene )distal portion; »fourthly, ‘conidia of .G. hungaricus are 
straight to sinuous while ours are sinuous or helical; Tastly, 
they differ significantly in dimensions, i.e.: in G. hungaricus 
the conidial body measures 90-120 x 2.5-3.5 ym, while in G. honrubiae 
it does not exceed 45 x 2 ym. It is interesting that conidia of 
the type species have been seen attached to nematodes onleaf litter. 
Our species has so far been seen in nature only as conidia in 
foam. 


ACKONOLEDGEMENTS 


I wish to thank Dr. E. Descals (Majorca, Spain) and Dr. L. Marvanova 
(CCM) for critical reviewing this manuscript. I also am indebted 
to Dr. J. Gonczol for providing additional information on G. hungari- 
cus. 


REFERENCES 


GONCZOL, J. & REVAY, A. (1985). Gorgomyces gen. nov. an unusual 
hyphomycete from terrestrial litter of Hungary. Nova Hedwigia, 
4): 453-461. 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 387-394 January-March 1989 


THE OCCURRENCE OF 
TUBER TEXENSE IN GEORGIA 


Richard T. Hanlin and Mei-Lee Wu 


Department of Plant Pathology 
University of Georgia 
Athens, Georgia 30602 


and 


Timothy B. Brenneman 


Department of Plant Pathology 
Coastal Plain Experiment Station 
Tifton, Georgia 31793 


ABSTRACT 


A hypogeous fungus found among roots in a pecan 
orchard represents the first report of a truffle 
from Georgia. It is identified as Tuber texense 
Heimsch. An illustrated description of the Georgia 
material is presented. 


INTRODUCTION 


Records of ascomycetes reported from Georgia have been 
maintained for over 70 years (Miller, 1941; Hanlin, 1963), 
and during that time Elaphomyces has been the only genus of 
hypogeous ascomycetes recorded from the state. In September, 
1987, during a visit to a pecan orchard near Albany, 
Georgia, numerous ascomata were observed among roots of 
several large pecan trees [Carya illinoiensis (Wang.) K. 
Koch] that had been exposed by soil erosion caused by a 
recent rain. Several ascomata were taken to the laboratory 
and examined, and were subsequently identified as Tuber 
texense. As this is apparently the first report of a 
tuberaceous species from Georgia and since it differs 
slightly from previous descriptions, an illustrated 
description of the Georgia material is presented here. 


388 


MATERIALS AND METHODS 


General observations and measurements of asci and 
internal tissues were made on fresh material mounted in 
water. Material to be sectioned was cut into 5 mm* blocks, 
fixed in formalin-propionic acid-alcohol, dehydrated through 
a tertiary butyl alcohol series, embedded in paraplast, 
sectioned at 6 or 10 wm and stained in iron hematoxylin. 
Sections to be examined under the scanning electron 
microscope were mounted on an 18 mm round cover glass, 
deparaffined in xylene, then the cover glass was mounted on 
a stub and sputter-coated with gold-palladium in a Hummer 
sputter coater (Gaudet and Kokko, 1984). Light micrographs 
were taken with a Nikon Optiphot on Kodak Technical Pan film 
2415; scanning micrographs were taken on a Philips 505 SEM 
with Polaroid Type 55 P/N film. These procedures have been 
previously described in greater detail (Hanlin & Tortolero, 
1988). 


OBSERVATIONS AND DISCUSSION 


Tuber texense Heimsch 

Ascomata hypogeous, up to 5.5 cm across, oval to nearly 
globose, light brown to dark reddish-brown, surface smooth, 
often lobed, especially on one side (Fig. 1). Interior 
consisting of a gleba with light and dark veins surrounded 
by a cortex (= medullary excipulum) and an exterior layer (= 
ectal excipulum) of pigmented cells (Fig. 2). Cortex 240-280 
pm thick, composed of three regions (Fig. 4, 8). Outermost 
2-3 rows of cells angular (textura angularis), with somewhat 


“swe er we ewww nen ene ewe ew eee wee ww ewxe eee wenn inwninaenenaniuawectie eee ewee’ we we ewe eX’ ew ew ew fee ewww Neg | = 


Fig. 1-7. Tuber texense. Fig. 2-7, paraffin sections of 


ascomata. Fig. 1. Five ascomata. X0./1. Fig. 2. Séetion 
through ascoma. X1.2. Fig. 3. Close-up of gleba with 
convoluted veins. X3.2. Fig. 4. Section through cortex 


showing outer region of pseudoparenchyma cells and inner 
region of parallel hyphae, and fertile vein with asci in 
interior. X262. Fig. 5. Section through outer portion of 
cortex showing outer rows of enlarged, pigmented cells. 
X514. Fig. 6. Section through sterile vein bordered by 
asci. X250. Fig. 7. Close-up of asci in fertile area. X266. 


389 


390 


thickened, pigmented walls (Fig. 5). These surface cells 
intergrade into a region ca. 125 pm thick of small, 
compactly arranged pseudoparenchymatous cells with thin 
walls that vary in shape from globose to angular or short 
hyphal; the latter are oriented perpendicularly to the 
margin of the ascoma. Inner half of cortex (subcortex) 
composed of small, thin-walled, globose cells among tightly 
interwoven hyphae oriented parallel to periphery of ascoma; 
this region extends into gleba as sterile veins. Gleba 
composed of white, convoluted sterile veins (venae 
externae) bordered by brown fertile veins (venae internae) 
(Fig. 3). Sterile veins variable in width, branched, 
composed of hyaline, interwoven hyphae (textura 
intricata) (Fig. 10), becoming compressed by developing asci 
(Fig. 6, 9). Fertile veins brown, cells crowded, often 
indistinct and appearing as surrounded by mucus at maturity, 
enveloping numerous asci (Fig. 7, 11). Ascigerous areas in 
immature ascomata often separated by small veins of hyaline 
hyphae that form a loose, open network that is crushed as 
the asci develop. Asci unitunicate, thick- walled (Fig. 16), 
persistent, variable in size, (/0)-88-(122) X (34)-48-(58) 
pm (including stipe), subglobose to ovoid, short- or 
long-stipitate [stipe (8)-22-(58) wpm], containing 1-6 
(usually 4) spores (Fig. 14-15). Ascospores unicellular, 
oval to occasionally subglobose, golden-brown at maturity, 
(22)-28-(36) X (16)-19-(24) pm, containing oil droplets, 
densely covered with spines that vary from minute to long 
and distinct (Fig. 17-18), up to 2 pm in length. The bases 
of the spines interconnect to form a reticulate pattern 
(Eig elLone 


Collected among roots of Carya illinoiensis (Wang.) K. 
Koch, Dougherty County, Georgia, September 18, 1987, T. B. 
Brenneman and P. F. Bertrand. Specimens deposited in GAM 
(#12742) . 


The material collected contained asci in all stages of 
maturity, permitting observation of ascus development. Asci 
arise from croziers formed from ascogenous hyphae (Fig. 12). 
The ascus mother cell expands to form a clavate ascus (Fig. 
13), which then develops into a mature ascus with 
ascospores. Developing asci often have a eclamp-like 
structure at the base that results from the fusion of the 
tip and basal cells of the crozier; this apparently forms 
the basis for the erroneous reports in the literature that 
tuberaceous asci possess clamp connections (Alexopoulos and 
Mims, 1979). 


391 


oS es 


Figs. 8-11. SEM micrographs of Tuber texense. Fig. 8% 
Section through cortex and portion of gleba. Bar = 1.0 mn. 
Fig. 9. Close-up of sterile vein bordered by asci. Bar = Ome 
mm. Fig. 10. Close-up of interwoven hyphae comprising 
sterile vein. Bar = 10 wm. Fig. 11. Asci in fertile vein. 


Bars =.0.1 mn. 


392 


The structure of our material agrees well with that 
described for T. melanosporum Vitt. (Parguey-Leduc et al., 
1987), except for the large pyramidal scales that cover the 
surface of T. melanosporum. 


Gilkey (1939) recognized a single species of Tuber in 
North America with spinose ascospores, T. candidum Harkness, 
to which she later added T. harknessii Gilkey (1954). A 
third species, T. texense, was described from Texas by 
Heimsch (1958); this species differs from T. candidum and T. 
harknessii in the formation of a reticulum on the spore 
surface that is associated with the spines. On the basis of 
these characteristics, our material is considered to be T. 
texense. In addition to Texas and Georgia, this species has 
also been found near Gainesville, Florida (James Kimbrough, 
personal communication). 


Like our material, T. texense was found among roots at 
the base of a pecan tree. The occurrence of numerous 
ascomata in close association with roots of pecan suggests 
the possibility of a mycorrhizal association, but no direct 
evidence was found to support this. Tuber melanosporum has 
been demonstrated to form mycorrhizae with Corylus avellana 
L., Quercus spp., and other hardwood species (Delmas, 1976). 


ACKNOWLEDGMENTS 


The assistance of J. O. Owens and P. F. Bertrand is 
gratefully acknowledged. The manuscript was reviewed by J. 
W. Kimbrough and R. G. Roberts. 


393 


. - : 2 18 


ferameeee LOontisht (Fig. 12-15; 17) and SEM) (Fig.) 16, 18) 
micrographs of Tuber texense. Fig. 12. Young ascus mother 
cell formed from crozier. X1680. EL ee oe OUnm aS GUS 
X1000. Fig. 14. Two-spored ascus. X4/5. Bi gee Los 
Four-spored ascus. X375. Fig. 16. Close-up of ascus with 
ascospore. Note thickened ascus wall. Bar = 10 pm. Fig. 1/. 
Mature ascospore with spines. X950. Fig. 18. Close-up of 


ascospore showing spines with interconnecting bases. Bar = 
10 pm. 


394 


LITERATURE CITED 


Alexopoulos, C. J., and» C. WW. /Mims” 91979. “Intvodme tam. 
Mycology, 3rd ed. John Wiley & Sons, New York. 632 p. 

Delmas, J. 1976. La truffe et sa. culture. |InsteeeNagee 
Recher. Agron. Etude No. 60: 1-54. - 

Gaudet,.'D. “Ax, and. E: G. Kokko 2) 1984.) Appliicameame 
scanning electron microscopy to paraffin-embedded plant 
tissues to study invasive processes of plant-pathogenic 
fungi. Phytopathology 74: 1078-1080. 

Gilkey, H. M. 1939. Tuberales of North America. Oregon St. 
Monogr) Stud aFBOL, NO. eb loo. 

Gilkey, H. M. 1954. Tuberales. No. Amer. Flora, Ser. II(Pt. 
1S) sees OF 

Hanlin, R:* T. .1963. A revision of the Ascomycetesmmon 
Georgia. Univ. Georgia Agric. Expt. Sta. Mimeo Ser. N. 
Shue Wi foye t hhc sper 

Hanlin, R. T.,; .and O.. Tortolero.. 1988- ~MorphaiGe ae 
Sclerotium coffeicola, a tropical foliar pathogen. Can. 
erbOL ms OOmGI ne DLeSSiE 

Heimsch, C. 1958. The first recorded truffle from Texas. 
Mycologia 50: 657-660. 

Miller, J. H. 1941. The Ascomycetes of Georgia. PLE; Dice 
Reptyeescupp eis ago oF 

Parguey-Leduc, A.,' G. Montant, and M. Kulifajeeieage 
Morphologie et structure de l’ascocarpe adulte du Tuber 
melanosporum Vitt. (Truffe noire du  Périgord, 
Discomycétes). Cryptogamie, Mycol. 8: 173-202. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 395-397 January-March 1989 


A SURVEY OF PILOBOLUS FROM YELLOWSTONE NATIONAL PARK 


K. MICHAEL FOOS AND JUDITH A. ROYER 


Department of Biology 
Indiana University East, Richmond, IN 47374 


INTRODUCTION 


A survey of Pilobolus from Yellowstone National Park was undertaken to 
extend our current knowledge of the geographic distribution of this 
coprophilous fungus. While studies of Pilobolus indicate that the organism 
is widely found, those in the United States have been from eastern and 
midwestern states (1-3,7). None have been published describing the 
isolation of Pilobolus in the Rocky Mountains. This study was designed to 
determine whether Pilobolus is present in this region of North America. 


METHODS AND MATERIALS 


Isolates of Pilobolus collected in this study were all from fecal samples from 
herbivores within the boundaries of Yellowstone National Park, a site 
selected for this study because it contains many regions that are 
representative of the Northern Rocky Mountain area but are relatively 
undisturbed by domestic grazing and cultivation. The samples were 
collected during June and July of successive years from animals having 
access to native vegetation only. 


Fresh (less than one hour old) fecal samples were collected in 20 different 
areas to examine for Pilobolus. Fecal samples from twenty-four bison (Bison 
bison), twenty-six elk (Cervus canadensis), twenty-two moose (Alces alces), 
six mule deer (Odocoileus hemionus), ten antelope (Antilocapra americana), 
six bighorn sheep (Ovis canadensis), and one horse (Equus caballus) were 
collected. These collections were made aseptically in plastic baggies and 
were transferred within hours to aseptic plastic cups as described by Foos 
and Royer (4). During the field period light and temperature conditions 
varied considerably, and cultures were maintained under shaded outdoor 
conditions. Upon return to the laboratory all cultures were maintained at 
room temperature under cool white fluorescent lights (2000 Ix) with 
alternating 12 hour light and dark periods. 


Isolates were obtained by removing single sporangia from the sides or tops 
of preparation dishes with sterile inoculating needles and transferring these 
sporangia to petri dishes containing dung agar. After initial isolations, 
hyphal tip transfers were used to maintain stock cultures. 


396 


Sporangia to be studied were collected from the lids of petri dishes with 
sterile inoculating needles and mounted in lactophenol. One hundred 
spores from each sporangium were examined and measured at 1000X 
using brightfield microscopy. Columella were observed by removing 
sporangia with microforceps. 


Measurements of taxonomic structures were made from the original isolates 
on dung and from cultures growing on dung agar in the laboratory. 


RESULTS 


Seventy-two isolates of Pilobolus representing three species were 
recovered from 95 dung samples collected in Yellowstone National Park. 
These species were: P. crystallinus, P. kleinii, and P. roridus. 


Pilobolus crystallinus (Wiggers) Tode: Fries (5) 


Pilobolus crystallinus sporangiophores are 2 to 10 mm long, and are clear 
to pale yellow in color. Trophocysts develop submerged in the substratum 
and are usually 300 to 500 um long. Sporangia are covered with a dark wall 
and range from 80-300 um in diameter. Columella are papillate and extend 
deeply into the sporangia. Sporangiospores are pale yellow ellipses which 
measure 9.76 + 0.60 (6-14.5) um_ in length by 6.35 + 0.65 (5-11.5) um in 
width producing a length to width ratio of 1.53. 


Pilobolus crystallinus was isolated in 18 locations in Yellowstone National 
Park from the dung of antelope, bighorn sheep, bison, elk, and mule deer. 


Pilobolus kleinii van Tieghem (8) 


Pilobolus kleinii sporangiophores measure 1 to 10 mm in length and arise 
from dark yellow turnip-shaped trophocysts 250-500 um in diameter. The 
trophocysts are often partially submerged within the substratum. Sporangia 
have a dark, smooth wall measuring 80-450 um across. The columella are 
papillate and extend deeply into the sporangia. Sporangiospores are 
yellow and elliptical, measuring 10.95 + 0.21 (6.5-15) um in length by 7.52 + 
0.67 (5.5-12.5) um in width with a length to width ratio of 1.46. 


Pilobolus kleinii was isolated in 35 locations in Yellowstone National Park 
from the dung of antelope, bighorn sheep, bison, elk, horse, and moose. 


Pilobolus roridus (Bolt.) Pers. (6) 


Pilobolus roridus sporangiophores are 1 to 8 mm long. Sporangia are dark; 
smooth, and hemispherical. They average 80-450 um in diameter. 
Trophocysts are 250- 400 um in diameter, nearly spherical, and bright 
Orange in color. Sporangia are located on blunt rounded conical columella. 
Sporangiospores are pale yellow to colorless, oval in shape and measure 
5.47 + 0.23 (3-7.5) um in length and 3.66 + 0.39 (2.5-6.5) um in width. The 
length to width ratio is 1.49. 


Pilobolus roridus was isolated in 18 locations in Yellowstone National Park 
from the dung of antelope, bighorn sheep, elk, moose, and mule deer. 


byes 


DISCUSSION 


Pilobolus has been isolated in many places, but this is the first published 
report of Pilobolus from the Rocky Mountain Region of the United States. All 
fecal samples containing isolates of Pilobolus were recovered during 
summer months from seven different species of herbivores. These 
collections were of fresh material made several miles inside the Park 
boundaries. Of the ninety-five samples collected, seventy-one contained 
one or more isolates of Pilobolus. Twenty-two of twenty-four fecal samples 
from bison contained Pilobolus, as did five of twenty-two from moose, five of 
six from mule deer, twenty- five of twenty-six from elk, eight of ten from 
antelope, five of six from bighorn sheep, and the sample from the horse. 
Overall, more than seventy-five percent of the fecal samples contained 
isolates of Pilobolus indicating that this fungus is widely distributed among 
the different herbivores. 


It is not possible to make correlations between collecting sites in the Park 
and the particular species of Pilobolus found, each having been found in at 
least 18 different locations. Nor is it possible to generalize about a 
particular species of herbivore and a particular species of Pilobolus. All 
three species of Pilobolus isolated in this study were found in fecal samples 
of elk, antelope and bighorn sheep, while bison, moose and mule deer each 
contained two species. The species of Pilobolus found in Yellowstone are 
well distributed throughout the geographic area and the various species of 
herbivores in which they are found. 


It might be anticipated that in a study of this size a larger number of Pilobolus 
species would have been recovered. In other surveys four to six species 
have been isolated. The recovery of only three species may indicate that 
the environmental factors, particularly climatic conditions, restrict growth of 
other species of Pilobolus. However, with three-fourths of the fecal samples 
collected containing isolates of Pilobolus, it is clear that there are abundant 
representatives of the genus, even if only three species are present. 


LITERATURE CITED 


1. Bessey, E. A. 1946. Studies of Pilobolus: P. kleinii and P. longipes. 
Papers Michigan Acad. Science 32:15-26. 

2. Foos, K. Michael and James B. Rakestraw. 1985. A survey of Pilobolus 
from Lake County, Ohio. Ohio J. Science 83(3):137-138. 

3. Foos, K. Michael and Judith A. Royer. 1985. Isolation of the coprophilous 
fungus, Pilobolus, from Wayne County, Indiana. Proceed. Indiana Acad. 
Science 94:109-112. 

4. Foos, K. Michael and Judith A. Royer. 1986. Quick and easy methods for 
collecting coprophilous fungi. Proceed. Indiana Acad. Science 95:99-100. 
5. Fries, E. M. 1823. Systema Mycologicum. 2:308-309. 

6. Persoon, C. H. 1801. Synopsis methodica Fungorum. Part 1. p. 117-118. 
7. Sumstine, D. R. 1910. The North American Mucorales. Mycologia 2:125- 
154. 

8. Van Tieghem, P. 1876. Troisieme memoire sur les Mucorinees. Ann. Sci. 
Nat. 4:335-349. 


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MYCOTAXON 


Vol. XXXIV, No. 2, pp. 399-406 January-March 1989 


TWO NEW SPECIES OF PARMELIA (PARMELIACEAE, LICHENES), 
FURTHER NEW COMBINATIONS AND NOTES, AND ADDITIONAL 
NEW LICHEN RECORDS FROM SOUTHERN AFRICA 


FRANKLIN A. BRUSSE 


Botanical Research Institute, 
Private Bag X101, Pretoria, 
SOUTH AFRICA 


ABSTRACT 


Two new species of Parmelia (Parmeliaceae, lichenized Ascomycetes), P. agamalis 
Brusse and P. geckonalis Brusse, are described from southern Africa. Five new names are 
proposed: Parmelia mixta Brusse, P. perfunctata Brusse. P. phyllodactylaris Brusse, 
P. salax Brusse, and P. tantillum Brusse. Three new combinations are made: Parmelia 
protodysprosa (Hale) Brusse, P. saniensis (Hale) Brusse, and P. umtamvuna (Hale) Brusse. 
Notes on eight species in the Parmeliaceae pertinent to the flora of southern Africa are 
given. Five lichen species are newly recorded for southern Africa. 


NEW SPECIES 


PARMELIA AGAMALIS Brusse, sp. nov. Fig. 1 

Thallus foliosus, saxicola, adnatus, ad 6 cm diametro, 60—170 yum crassus. Lobi 
elongati, 1—3 mm lati. Thallus superne cinereus, nitidus, emaculatus, isidiatus, isidiis 
parvis, usque ad 0,1 mm crassis. Cortex superior 9-12 um crassus. Stratum gonidiale 
15—35 ym crassum, algis Trebouxiis, 5—16,5 um diametris. Medulla alba, 25—110 um 
crassa. Cortex. inferior 6—11 ym crassus. Thallus inferne piceus. Rhizinae simplices, 
sparsae. Apothecia non visa. Pycnidia hyalina, circa 150 um profunda et circa 90 um 
lata. Pycnidiosporae hyalinae, aciculares, 4—6 x 0,8 um. Thallus atranorinum, acidum 
hyposticticum, acidum hyposalazinicum, acidum sticticum, acidum hypoconsticticum et 
acidum consticticum continens. 

TYPUS—-SOUTH AFRICA, Cape Province, 3318 (Cape Town): Bothmaskloof Pass 
near Riebeeks Kasteel, Porseleinberg, on Table Mountain Sandstone outcrops on NW 
slope, alt. 300 m (—BD). F. Brusse 2782, 28.iv.1981 (PRE, holo-; COLO, LD, iso-). 
Figura 1. 

Thallus foliose, saxicolous, adnate, to 6 cm across, 60—170 um thick. Lobes elongate, 
1—3 mm wide. Upper surface grey, nitid, emaculate, isidiate, isidia small, up to 0,1 mm 


400 


thick. Upper cortex 9—12 ym thick. Algal layer 15—35 ym thick, algae Trebouxia, 5—16,5 
pm diam. Medulla white, 25—110 wm thick. Lower cortex 6—11 wm thick. Under 
surface black. Rhizines simple, sparse. Apothecia not seen. Pycnidia hyaline, about 150 
yum deep and about 90 um wide. Pycnidiospores hyaline, needle-like, 4—6 x 0,8 yum. 
Chemistry: atranorin in the cortex; hypostictic, hyposalazinic, stictic, pe setter ea 
and constictic acids in the medulla. - 
Etymology: agamalis (L) = relating to Agamas, a genus of lizards, often with some 
blue colouration on the head; specifically the rock agama, Agama atra. These and other 
saxicolous lizards, may play a role in the spreading of vegetatively propagating lichens 
such as Parmelia agamalis, as they scuttle over the rocks on which these lichens grow. 
There are no known species of the section Paraparmelia, that are isidiate, with a 
black lower surface and contain the hypostictic acid group with or without the stictic 
acid group of substances, in southern Africa. Parmelia ischnoides Kurok. (Hale 1976), 
may well be related, but this species is smaller and contains only the stictic acid group. 


On the other hand, there are several isidiate Paraparmelias in Australia with black 
lower surfaces, containing the hypostictic acid group with or without the stictic acid 
group. Of these, Parmelia numinbahensis Elix (1979) is undoubtedly the closest relative 
of this new species, but contains only the hypostictic acid group, without stictic and 
constictic acids. The lobes of P. numinbahensis also seem longer and narrower, than 
those of P. agamalis, but this may depend on the range of variation displayed by these 
two species, which at this time is unknown. 


Parmelia neoquintaria (Hale) Elix & Stevens, the first hypostictic acid containing 
Paraparmelia discovered (Hale 1976), is more distantly related, having a pale lower 
surface, and a looser thallus (again depending on the range of variation). Based on my 
experience with southern African lichens, these habit differences are probably not 
significant. 

The recently described, Paraparmelia subtropica Elix & Johnston (1986), is also 
isidiate, with a black lower surface, but is a small lobed, tightly adnate lichen (lobes 
0,4—1,0 mm wide), with only traces of hypostictic and hyposalazinic acids. The major 
component of P. subtropica is norstictic acid, and a trace of connorstictic acid is also 
present. Paraparmelia subtropica is therefore more distantly related to Parmelia agamalis, 
and is probably more closely related to Parmelia ischnoides. 

The type material (particularly the holotype in PRE) has white short barrel shaped 
insect eggs with umbonate lids, laid on it and the adjacent rock, and are probably those 
of a chinch bug (Lygaeidae). 

The holotype specimen is also growing with Parmelia (Neofuscelia) caliginosa Ess. 


At present this new species is only known from the type collection, near Riebeeks 
Kasteel in the south-western Cape Province. 


PARMELIA GECKONALIS Brusse, sp. nov. Fig. 2 

Thallus subcrustosus, saxicola, ad 3 cm diametro, 60—140 um crassus. Lobi elongati, 
0,3—1,3 mm lati. Thallus superne flavo-viridis, opacus vel grosse pruinosus, pustulatus. 
Pustulae capitatae, substipitatae, ad 1,5 mm diametris. Cortex superior 8—15 ym crassus. 
Stratum gonidiale 15—45 ym crassum, algis Trebouxiis, 5,55—18 ym diametris. Medulla 
albida, 20—95 um crassa. Cortex inferior 4,5—8 ym crassus. Thallus inferne pallide brun- 
neus. Rhizinae simplices, 30—60 ym crassae. Apothecia et pycnidia non visa. Thallus 


401 


FIGURE 1.—Parmelia agamalis Brusse, habit. F. Brusse 2782. Scale in mm and cm. 


FIGURE 2.—Parmelia geckonalis Brusse, habit. F. Brusse 5536. Scale in mm and cm. 


402 


acidum usnicum, acidum salazinicum, et acidum norsticticum (tr.) continens. 


TYPUS—SOUTH AFRICA, Cape Province, 3218 (Clanwilliam): 8 km W of Olyven- 
boskraal, Witelskloof. On large Table Mountain Sandstone boulder outcrop, on SW side. 
On outer edge of roof of rock overhang, high up on outcrop. Alt. 500 m (—BD). F. 
Brusse 5536, 25 iii.1988 (PRE, holo-; BM, iso-). Figura 2. 


Thallus subcrustose, saxicolous, to 3 cm across, 60—140 ym thick. Lobes elongate, 
0,3—1,3 mm broad. Upper surface yellow-green, matt to coarsely pruinose, pustulate. 
Pustules capitate, substipitate, to 1,5 mm across. Upper cortex 8—15 ym thick. Algal 
layer 15—45 pm thick, algae Trebouxia, 5,5—18 um diam. Medulla whitish, 20—95 um 
thick. Lower cortex 4,5—8 ym thick. Lower surface pale brown. Rhizines simple, 30—60 
yum thick. Apothecia and pycnidia not seen. Chemistry: usnic acid in the cortex; sala- 
zinic and norstictic (tr.) acids and the ‘chalybaeizans unknown’ in the medulla. 


Etymology: geckonalis (L) = relating to geckos. Since this lichen grows on the 
roofs of rock overhangs, only geckos could spread this lichen species, by the adhesion of 
pustular fragments on the foot pads and underbody. 


Parmelia geckonalis is a unique species, and has no close relatives. It does, however, 
resemble P. mougeotii Schaer. ex Dietr. P. mougeotii contains the stictic acid group only 
and has a black lower surface, whereas P. geckonalis contains salazinic acid and the 
‘chalybaeizans unknown’ and has a pale lower surface. P. geckonalis also has a coarse 
pruinose upper surface and grows on the roofs of rock overhangs, whereas P. mougeotii 
is glossy above and does not grow on the roofs of rock overhangs. Xanthoparmelia 
pustulifera Hale (1986) is not much different from P. mougeotii Schaer. ex Dietr. and is 
probably a synonym of it. The presence of P. mougeotii (or X. pustulifera Hale) in the 
Cape Province has yet to be confirmed. 

Parmelia phyllodactylaris Brusse (synonym: Xanthoparmelia eruptens Hale (1987)) 
is another small lichen from the roofs of rock overhangs, but this lichen has dactyls and 
a black lower surface, and contains evernic and lecanoric acids in the medulla. 


At present this species is known only from the type locality, Witelskloof near Clan- 
william. 


NEW COMBINATIONS 


Since the crucial proposal 291A (McNeill 1986), concerning lectotypification was 
not passed at the Berlin Congress (Greuter & McNeill 1987, McNeill 1987), it is still 
better to use Parmelia, typified by Parmelia conspersa Ach., as was first done by Clements 
& Shear in 1931 (Brusse 1988a). The following new combinations and new names are 
therefore proposed: 


Parmelia mixta Brusse, nom. nov. 

Basionym: Xanthoparmelia paradoxa Hale, Mycotaxon 29: 261, 1987. (non Parme- 
lia paradoxa Hale, Phytologia 27: 1, 1973). 
Parmelia perfunctata Brusse, nom. nov. 

Basionym: Xanthoparmelia natalensis Hale, Mycotaxon 29: 259, 1987. (non Parme- 
lia natalensis Steiner & Zahlbruckner, Bot. Jb. 60: 515, 1926). 
Parmelia phyllodactylaris Brusse, nom. nov. 

Basionym: Xanthoparmelia eruptens Hale, Mycotaxon 29: 254, 1987. (non Parme- 
lia eruptens Kurokawa in Hale & Kurokawa, Contrib. U.S. natn. Herb. 36: 153, 1964). 


403 


Etymology: phyllodactylaris (L) = relating to geckos of the genus Phyllodactylus, 
in particular the striped gecko, Phyllodactylus lineatus rupicolus. Since this lichen grows 
on the underside of rock overhangs, only geckos and perhaps bats could be important in 
dispersing this vegetatively reproducing (dactyls) lichen. 

Parmelia protodysprosa (Hale) Brusse, comb. nov. 

Basionym: Xanthoparmelia protodysprosa Hale, Mycotaxon 29: 262, 1987. 
Parmelia salax Brusse, nom. nov. 

Basionym: Xanthoparmelia applicata Hale, Mycotaxon 30: 320, 1987. (non Parme- 
lia applicata (Stiz.) Esslinger, Bryologist 76: 307, 1973). 

Synonym: Xanthoparmelia rugulosa Hale, Mycotaxon 30: 328, 1987. (non Parme- 
lia rugulosa (Nyl.) Bab. & Mitt. in Hooker, Flora Tasmaniae 2: 348, 1860; nec Parmelia 
rugulosa (Mill. Arg.) Vainio, Résultats de Voyage duS.Y. Belgica, Botanique: 14, 1903). 


Etymology: salax (L) = lustful, lecherous or sexy, in allusion to the dense covering 
of apothecia often displayed by this species. 


Parmelia saniensis (Hale) Brusse, comb. nov. 
Basionym: Xanthoparmelia saniensis Hale, Mycotaxon 27: 596, 1986. 


Parmelia tantillum Brusse, nom. nov. 
Basionym: Xanthoparmelia inconspicua Hale, Mycotaxon 29: 25 6, 1987. (non Par- 
melia inconspicua Sprengel, Mantissa Prima ad Floram Halens.: 57, 1807). 


Parmelia umtamvuna (Hale) Brusse, comb. nov. 
Basionym: Xanthoparmelia umtamvuna Hale, Mycotaxon 29: 266, 1987. 


NOTES ON SOME PARMELIACEAE 


Xanthoparmelia brevilobata Hale, Mycotaxon 29: 253, 1987. 

The chemistry of this species is identical to that of Parmelia saniensis (Hale) Brusse, 
containing the pigments usnic acid and skyrin, and hypostictic, hyposalazinic and hypo- 
constictic acids, in addition to a minor amount of lividic acid, and a pale UV3¢)+ sub- 
stance similar to the ‘chalybaeizans unknown’. However, this species does not have the 
bumps on it that P. saniensis has. If these bumps on the type of P. saniensis are abnormal- 
ities, then X. brevilobata is certain to be a synonym of P. saniensis. Xanthoparmelia 
naudesnekia Hale, may be a chemosyndromic variant of this species, containing steno- 
sporonic acid instead of lividic acid. 

Xanthoparmelia diffractaica Hale, Mycotaxon 29: 254, 1987. 

This lichen contains major amounts of the ‘chalybaeizans unknown’ and barbatic acid, 
and a minor amount of 4-O-demethylbarbatic acid, in addition to the salazinic, diffractaic 
and usnic acids originally reported. This lichen is identical to Parmelia lesothoensis (Hale) 
Brusse, except for a slightly larger amount of barbatic acid present. 


Parmelia lesothoensis (Hale) Brusse, Mycotaxon 31(2): 539, 1988. 
Synonyms: Xanthoparmelia lesothoensis Hale, Mycotaxon 27: 582, 1986. 
: Xanthoparmelia diffractaica Hale, Mycotaxon 29: 254, 1987. 

Although Brusse (1988b) detected the ‘chalybaeizans unknown’ in this lichen, in 
addition to usnic, salazinic and diffractaic acids as originally reported, he did not notice 
the minor amount of barbatic acid hidden by usnic acid on plates B and C, and the trace 
of 4-O-demethylbarbatic acid. Slightly larger amounts of barbatic acid occur in the type 
of Xanthoparmelia diffractaica Hale, but otherwise there are no differences between 


404 


these two lichens. 


Parmelia perfunctata Brusse 

Synonym: Xanthoparmelia natalensis Hale, Mycotaxon 29: 259, 1987. 

This lichen also contains barbatic acid in fairly large quantities and a trace of 4-O- 
demethylbarbatic acid in addition to the diffractaic, physodalic, succinoprotocetraric, 
fumarprotocetraric and usnic acids originally reported. This lichen is very similar to the 
pantropical P. adplanata Mill. Arg. (Syn: P. subramigera Gyeln.), but contains diffractaic 
and barbatic acids in addition to the usual constituents. Distributional patterns may clarify 
the status of this species. 

Parmelia saniensis (Hale) Brusse 

Synonym: Xanthoparmelia saniensis Hale, Mycotaxon 27: 596, 1986. 

This lichen contains a pale UV3,¢9 + substance similar to the ‘chalybaeizans unknown, 
as well as lividic acid in addition to skyrin, usnic, hypostictic, hyposalazinic and hypo- 
constictic acids as originally reported. The chemistry is identical to that of X. brevilobata, 
but P. saniensis is ‘isidiate’. These ‘isidia’ are too large and variable in size to be called 
dactyls even, but may actually be bullate regrowths from damaged lobes. For the time 
being the type specimen of P. saniensis is regarded as an abnormal, damage induced growth 
form, while the type specimen of X. brevilobata is considered the normal condition. 

As mentioned previously, X. naudesnekia Hale, reported to contain stenosporonic 
rather than lividic acid, may be a chemosyndromic variant of P. saniensis Hale. A final 
verdict, will depend on a TLC analysis of the type material. 

Parmelia spissa Brusse, Mycotaxon 31(1): 158, 1988. 

Additional isotypes of this species have been deposited at BM, COLO, UPS and US. 
Further specimens of F. Brusse 4592 (paratypes) have been deposited at UC, UPS and 
US, over and above the original BM, COLO, LD and PRE. 

Parmelia tantillum Brusse 

Synonym: Xanthoparmelia inconspicua Hale, Mycotaxon 29: 256, 1987. 

A thin-layer chromatographic study of this lichen revealed the presence of usnic, 
hypostictic, hyposalazinic and hypoconstictic acids, and the same unidentified pale 
UV360+ substance, similar to the ‘chalybaeizans unknown’ that occurs in P. saniensis 
(Hale) Brusse. Dull reddish pigments were not detected. 

Parmelia adhaerens Nyl, is closely related, but lacks hyposalazinic acid and contains 
stictic acid instead. 

Parmelia umtamvuna (Hale) Brusse 

Synonym: Xanthoparmelia umtamyvuna Hale, Mycotaxon 29: 266, 1987. 

An isotype specimen (PRE) contains stictic, constictic and usnic acids, but diffractaic 
acid is not present as originally reported. However, barbatic acid and a trace of 4-O- 
demethylbarbatic acid are present. 


ADDITIONAL NEW LICHEN RECORDS 


Recently Karnefelt (1987) reported a number of interesting new generic and species 
records for southern Africa, based on the high altitude collections of Ms. Elsie Esterhuy- 
sen. I recently took the opportunity to examine several high altitude localities personally, 
through the kind permission of the landowners, post-office authorities and forestry 
officials concerned, and found several further new generic and species records for the 


405 


southern African lichen flora, over and above those reported previously (Brusse 1988b). 


Acroscyphus sphaerophoroides Lé&v. 

SOUTH AFRICA, CAPE PROVINCE.—3319 (Worcester): Summit of Jonaskop in 
the Riviersonderend mountains, near Villiersdorp. On SE side, on top of Table Mountain 
sandstone outcrop in prominent position, and fully exposed. Sporadic. Alt. 1640 m 
(—DC). F. Brusse 5457, 21 iii.1988 (BM, LD, PRE, UPS). 


According to Tibell (1984) this is a rare but widespread lichen, occurring at high 
altitudes between the latitudes 43° N and 43° S. While the latitude of the present locality is 
within this range, the altitude is comparatively low (1 640 m), being much less than 
4000 m. It seems as though this lichen may occur at lower altitudes at higher latitudes. 
This is the first record of this lichen from the African continent, as it is known from 
Peru, Patagonia, Mexico, Bhutan, Yunnan, Sikang and Japan. Although sporadic in oc- 
currence at any particular locality, A. sphaerophoroides does not seem to be rare in the 
Riviersonderend Mountains, because it was collected again at Galgeberg, 16 kilometres 
to the east of Jonaskop, as the crow flies. That material was badly burned in a recent 
bush fire and is not cited here. 


Gomphillus calicioides (Delise) Ny]. 

SOUTH AFRICA, CAPE PROVINCE.—3318 (Cape Town): South side of Table 
Mountain, SSW slope overlooking Hely-Hutchinson Reservoir, on Hypnum cupressiforme 
Hedw. low down on rock. Growing with Megalospora tuberculosa (Fée) Sipman. Alt. 
950 m (—CD). F. Brusse 5422b, 19.iii.1988 (PRE). 

In an effort to extend the known range of Psathyrophlyctis serpentaria Brusse, a 
yellow granular lichen was collected on Hypnum cupressiforme Hedw. low down on a 
rock, on the south side of Table Mountain. The apotheciate material turned out to be 
Megalospora tuberculosa (Fée) Sipman, while the more olive-yellow material was Psa- 
thyrophlyctis serpentaria, as judged by the chemistry. In addition to these lichens, Gom- 
phillus calicioides, was also found. (Jacques van Rooy kindly named the moss). 
Hypogymnia subphysodes (Krempelh.) Filson 

SOUTH AFRICA, CAPE PROVINCE.—3318 (Cape Town): East side of Table 
Mountain, from Fernwood Buttress to Erica Buttress. On S sides of TMS outcrops and 
boulders, on steep E slope. Fairly common near crevices and in rock pockets. Alt. 1 000 
m (—CD). F. Brusse 5424, 19.iii.1988 (BM, LD, PRE). 3319 (Worcester): Jonaskop in 
the Riviersonderend Mountains near Villiersdorp. On S Table Mountain Sandstone cliffs. 
Tends to occur near crevices and in rock pockets. Alt. 1640 m (—DC). F. Brusse 5471, 
20.iii.1988 (BM, LD, O, PRE, UPS). Lit. Elix (1980). 

Hypogymnia subphysodes (Krempelh.) Filson var. austerodioides Elix 

SOUTH AFRICA, CAPE PROVINCE.—3318 (Cape Town): East side of Table 
Mountain, from Fernwood Buttress to Erica Buttress. On S sides of TMS outcrops and 
boulders, on steep E slope. Fairly common near crevices and in rock pockets. Alt. 1 000 
m (—CD). F. Brusse 5419. 19 .iii.1988 (PRE). Lit. Elix (1980). 

Parmelia (Xanthoparmelia) kurokawae Hale 

SOUTH WEST AFRICA/NAMIBIA.—2017 (Waterberg): Waterberg trail at camp- 
site, on sandstone rocks at S base of cliffs (—CA).F. Brusse 4219, 1984.03.22 (PRE, LD). 
Porpidia urbanskyana (Zahlbr.) Brusse 

SOUTH AFRICA, NATAL.—2828 (Bethlehem): 31 km S of Phuthaditjhaba (Wit- 
sieshoek), summit of Western Buttress (Mont-aux-Sources), on S basalt face, in heavy 


406 


seepage, on gentle S slope. Alt. 3080 m (—DB). F. Brusse 5548, 5.iv.1988 (BM, PRE). 
ACKNOWLEDGEMENTS 


The author is grateful to Dr J.A. Elix for reviewing this paper. Thanks are extended 
to the following herbaria for the loan of type and other material; BM, FH, G, GLAM, H, 
LD, TNS, TRH, TUR, VER, W and ZT. Mrs A.J. Romanowski prepared the photographs, 
Mrs M. van der Merwe typed and Mrs S.S. Brink typeset this manuscript, with my gratitude. 


LITERATURE CITED 


BRUSSE, F.A. 1988a. Three new species of Parmelia (Lichenes) from southern Africa. 
Mycotaxon 31(1): 155-162, p. 156-157. 

BRUSSE, F.A. 1988b. Five new species of Parmelia (Parmeliaceae, Lichenized Ascomy- 
cetes) from southern Africa, with new combinations and notes, and new lichen 
records. Mycotaxon 31(2): 533-555. 

CLEMENTS, F.E. & SHEAR, C.L. 1931. The Genera of Fungi (Edn 2). New York: 
Hafner. viii + 496 pp., 58 pl.; p. 322. 

ELIX, J.A. 1979. New species of Parmelia (Lichens) from Australia. Australian Journal 
of Botany 27(6): 873—883, p. 877-879. 

ELIX, J.A. 1980. A taxonomic revision of the lichen genus Hypogymnia in Australasia. 
Brunonia 2(2): 175—245. 

ELIX, J.A. & JOHNSTON, JEN. 1986. New species of Paraparmelia (Lichenised As- 
comycotina) from Australia and New Zealand. Brunonia 9: 139—153, p. 151—152. 

GREUTER, W. & McNEILL, J. 1987. Synopsis of Proposals on Botanical Nomenclature, 
Berlin 1987. Taxon 36(1): 174—281, p. 192. 

HALE, M.E. 1976. A monograph of the lichen genus Pseudoparmelia Lynge (Parmeli- 
aceae). Smithsonian Contributions to Botany 31: 1—62. 

HALE, M.E. 1986. New species of the lichen genus Xanthoparmelia from southern Africa 
(Ascomycotina: Parmeliaceae). Mycotaxon 27: 563—610. 

HALE, M.E. 1987. Additions to the Xanthoparmelia flora of southern Africa (Lichenized 
Ascomycotina: Parmeliaceae). Mycotaxon 29: 251—266. 

KAERNEFELT, I. 1987. Cetraria (Parmeliaceae) and some related genera on the African 
continent. Bothalia 17(1): 45—49. 

McNEILL, J. 1986. Proposals on lectotypification and the report of the Committee on 
Lectotypification. Taxon 35: 867—880, p. 873. 

McNEILL, J. 1987. XIV International Botanical Congress: mail vote and final Congress 
action on nomenclatural proposals. Taxon 36(4): 858—868, p. 859. 

TIBELL, L. 1984. A Reappraisal of the Taxonomy of Caliciales. Beihefte zur Nova 
Hedwigia 79: 597—713. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 407-428 January-March 1989 


POLLUTION ATMOSPHERIQUE ET LICHENS 


DANS LA VILLE. DE SANTIAGO DU CHILI 


MANUEL MAHU 


Departamento de Ciencias Ecoldégicas 
Facultad de Ciencias. Universidad de Chile 
Casilla 655 caroant1ago 


La Constituci6én asegura a todas las personas: 
El derecho a vivir en un medio ambiente libre 
de contaminacion. 
Es deber del Estado velar para que este dere- 
cho no sea afectado, y tutelar la preservacién 
de la naturaleza. 
La ley podra establecer restricciones especifi- 
cas al ejercicio de determinados derechos o li- 
bertades para proteger el medio ambiente. 

Art. 19. N2 8 
Constituci6én Politica de la 
Republica de Chile, 1980 


ABSTRACT 

Wevuemcnie Aim. Ol ,1indingy, biological” indicators of 
Pereeporiurion, an. inventory of lichens “collected, .in dif- 
ferent areas of Santiago de Chile was made. The distribu- 
mrOong,Ol) lichens is discussed im relation to “geographic 
fee eeone Or, the city, its climatic characteristics and 
Peeepoeruancs. ~Alvthougth the «distribution of,- lichens in 
the city varied according of species, lichens were present 
in all areas studied. 


408 


INTRODUCTION 

Pendant les derniéres années les habitants de Santiago 
sont preoccupés par la pollution atmosphérique accentuée 
que chaque année semble aller en augmentation. 

Les autorités du Ministerio de Salud a travers du La- 
boratorio de Contaminaciodn Atmosférica ont effectué des 
mesures des contaminants en différentes lieux de la ville. 
En plus, il contrédle periddiquement les sources fixes des 
contaminants. D'autre part, les autorités du Ministerio 
de Transporte y Telecomunicaciones ont adopté des disposi- 
tions qui restreignent la circulation véhiculaire dans 
l'aire centrale de la ville selon dernier numéro de la ma- 
triculation, une fois par semaine, de Lundi a Vendredi et 
a disposé que toute véhicule avec moteur a explosion doit 
6étre contrédlée, au moins, une fois par an dans ses aspects 
mechaniques et d'emission des contaminants (% CO). Mais, 
tous le deux organismes n'ont pas obtenue la disminution 
des indices de contaminants atmosphériques. 

Le probléme de la pollution atmosphérique qui affecte 
beaucoup des villes a originé différents publications. En 
Santiago, par example, on a analysé des aerosols de l'aire 
urbaine et suburbaine en trouvant 25 éléments chimiques 
(Préndez et al., 1984). 

L'action du CO de l'air dans l'accroissement des li- 
chens a été étudié en Anglaterre et Gales par Hawksworth 
& Rose (1970); les lichens comme bioindicateurs naturelles 
ont été étudié en relation aux éléments stables et radiac- 
tives de l'air par Looney & al. (1986). Les cryptogames 
ont été employées comme organismes moniteurs de pollution 
de l'air par metaux, selon Rasmussen et al. (1980) et la 
ville de Caracas fit l'objet d'une étude de la distribu- 
tion des lichens d'accord au degré des contaminants atmos- 
phériques par Vareschi & Moreno (1973). 

Etant donnée que la ville de Santiago présente une 
pollution ‘atmosphérique elevée, fait que je décidaiawa 
y a deja quelques années de récolter les lichens qui se 
trouvaient dans les différents lieux, avec l'objet de de- 
terminer jusqu'a quel point la pollution atmosphérique au- 
rait un effect negatif sur les populations lichéniques. 

Fn ce travail on montre la distribution des lichens 
dans l'aire urbaine de Santiago et les indices de contami- 
nants mesurés par les Stations distribuées dans la ville. 


SITUATION DE LA VILLE DE SANTIAGO 

La ville de Santiago est située dans une depression 
du Chili centrale comprise entre les 322 10' - 322 54' § 
et ‘les 702 29' - 702 OO W. Le dimension N-S demcenee 


409 


vallée sont 83 km et sa latitude E-W de 54 km environ. Au 
Nord la depression est limitée par un cordon de collines 
gui forment la Cuesta El Manzano, au Sud par 1'Angostura de 
Paine, au l'Est par la Cordillére des Andes. Les sommets 
plus importants de cette Cordillére sont le Cerro Colocalan 
540) m), “le Cerro La’ Provincia (2.731 '>m), le)Cerro’ San 
Bane 3-249 mM). le),Cerro..Llos. .Cristales:(.3.057,.m)). et. le 
Cerro Pabellones (2.497 m). Par l'Ouest cette depression 
est limitée par le cordon montagneux de la Cordillera de 
la Costa et ses sommets plus importants sont le Cerro Ro- 
DremAlcoOwt(c.0cy my), le Cerro Perro’ ‘Muerto. (1.391. m)) et. le 
Cerrcowcantillana (2.281 m). La ville de Santiago est situé 
Bema! OS et. (02: 38" Wh a-600 m) (Fig. 1): 


CLIMAT 

D'accord aux indices de Thorthwaite, Santiago a un 
climat modéré semi-aride, avec un hiver bénigne et un été 
sec. 


CARACTERISTIQUES DES CONSTITUANTS CLIMATIQUES 


a) Vents: La frequence des directions du vent en Juin 
et Janvier, dans la période 1831-1960, avec trois obser- 
Tartons: par jour (O8°%hrs.: 14 hrs. et 20 hrs..)) nous indique 
que au Juin les vents du SW occurrent dans le 12.1%, il'a 
un période de calme dans le 70.6% et le reste dans d'autres 
directions. En général, les vents qui dominent procedent 
du SW pendant le jour et de l'Est dans la nuit avec une 
faible vitesse. Les vents provoquent des inversions ther- 
miques frécuentes en hiver qui souvent subsistent toute 
la journée. Ces inversions causent la formation d'un 
"plafont" qui en été arrive aux 1.000 m et en hiver, entre 
les 300 et 400 m sur la ville, dessous duquel l'air conta- 
miné est gris ou obscur. On a observé en Aofit dans un jour 
de grande inversion thermique, la température au niveau 
Gumeeory est de 42°C et dans .la. ‘sommet |de’ 1'inversion: 
(1.000 m) est de 162 C. La masse supérieure d'air froid 
empéche que les contaminants montent aux couches supérieu- 
res de l'atmosphére restant retenus au-dessous d'elles 
Cet gemc. 3.) tet’ 5). 

La pollution atmosphérique est plus intense lorsque 
se présentent des inversions fortes de témperature avec 
des vents faibles dans les masses baisses de la atmos- 
phére. 


b) Precipitations: Les precipitations sont, en moyenne 
annuelle, de 329.8 mm et les mois de Mai a Aoait, les plus 
pluvieux, avec une moyenne de 64 mm. 


410 


® ia 


Fig. 1.- Région centrale du Chili. A: Ville de Santiago dans la 
vallée centrale et ses limits N-S-E-W. Image captée par le Satélite 
Landsat-2, le 22-III-1975, & 09 hrs. 57 min. 20 sec. a ume altitude 
de 900 km (Courtaisie d'Instituto Geografico Militar, Santiago). 


c) Témperature et Humidité Relative de 1'Air: La tém- 
perature moyenne annuelle est de 14.52 C et la humidité 
moyenne relative de l'air 83% (08 hrs), 48% (14 hrs) et 
647 aC2Dehrson 


PROTECTION DE L'ENVIRONNEMENT 

Par disposition du Ministerio de Salud on a communiqué 
les normes sanitaires destinées a prévoir et a contrdler 
la pollution au Chili. Dans cette resolution on défine la 
pollution atmosphérique comme "la presence dans l'air d'un 
ou plus contaminants ou quelconque combination d'eux en 


411 


C C2 de Renca 
903 m 


Fig. 2.- Vue W de la ville de Santiago couverte par le nuage de con- 
taminants, depuis le Cerro San Cristdébal, 880 m (7-V-1987). 


Santa Lucia 
628 m 


Fig. 3.- Vue S de la ville de Santiago couverte par le nuage de con- 
taminants, depuis le Cerro San Cristébal, 880 m (7-V-1987). 


412 


C2 San Ramon 


Fig. 4.- Vue E de la ville de Santiago couverte par le nuage de con- 
taminants, depuis le Cerro San Cristdébal, 800 m (7-V-1987). 


Fig. 5.- Vue de la ville de Santiago couverte par le nuage de con- 
taminants, depuis les Cerros de San Carlos de Apoquindo, a l'Est de la 
ville, 970 m (6-XI-1987). 


413 


concentration au niveau tels qui dérangent ou détruisent 
la vie, la santé et le bienétre humain, la flore et la 
faune ou dégradent la qualité de l'air, des biens, des re- 
cours nationales et particuliers". 


SOURCES DE POLLUTION 

Dans la ville de Santiago il y a environ 2.500 indus- 
tries» 300.000 automoviles, 1.500 taxis et 4.000 autobus 
gui dépensent mensuellement 5.000 m3 d'essence. D'autre 
sources de la pollution la constituent la brdlure du bois, 
du charbon et du mazut pour la chauffage et la incinération 
des déchets ont fait que la pollution atmosphérique dans 
la ville de Santiago soit alarmante. On voit clairement 
les effets de la pollution atmosphérique dans les feuilles 
d'arbres: la poussiére sédimentaire qui couvre la lame fo- 
liaire (Fig. 6) fait disminuer ses activitées vitales et 
la tombée des feuilles (Fig. 7). 


Fig. 6.- Feuilles de Magnolia Fig. 7.- Acacia retinodes avec 
grandiflora et de Eriobotrya ja- des branches inférieures defoliées 
ponica couvertes par la poussiére a cause de la pollution atmosphé- 

sédimentaire. La moitie droite rique. 

des lames sont propres. _ Ave. Grecia 2976 
Rue Diagonal Paraguay/Carmen 


414 


MESURAGE DES CONTAMINANTS 

Pour le mesurage des contaminants atmosphériques on 
a instalé quelques Stations “dans Ja ville, (Hig eee 
données obtenues sont representées dans les Fig. 9-13. 
L'enregistrement du Co a été effectué seulement dans la 
Station 1, les autres contaminants ont été enregistrés_dans 
les différents Stations établies. 


CONSTITUANTS DE LA POLLUTION ATMOSPHERIQUE 


OXIDE D'AZOTE (NO2) 

Les enregistrements ont été effectué avec un appareil 
semi-automatique de gazes RAC, Modéle PV. Pour l'analyse 
on emploie la méthode colorimétrique avec arsenite de 
sodium (NaAsO2), 24 heures du jour, deux fois par semaine, 
selon calendrier établi au commencement de l'année. 

La Norme de Qualité de 1'Air au Chili pour le NO2 est 
100 ug/m3 (moyenne annuelle) ou, 300 ug/m3/24 heures (1 
liGiso 1 Dal sau 

Dans la période 1977-1986, aucune Station a registrée, 
en moyenne, un excédent a la norme. La Station 2, situé 
dans l'aire centrale de Santiago, registrait l'indice le 
plus haut; 87% ug/m3’ et les Stations 5 et’ 6 *réeratranene 
76 et 69 ug/m3 respectivement. Les trois Stations se trou- 
vaient dans les aires de haut flux véhiculaire. Les Sta- 
tions 13 et 14 ont régistré les indices les plus bas avec 
28 et 26 .ug/m3,- tous. les’'deux sont loins du Centremyverre 
dans les aires de bas flux véhiculaire. 

La norme a été dépassait plusieurs fois. La Station 
2 en 1980 avec 112 ug/m3 et en 1981 avec 149 ug/m3, la 
Station 5 en 1981 avec 110 ug/m3, en 1985 avec 112 ug/m3 
et en 1986 avec 116 ug/m3, et la Station 6 avec 132 ug/m3, 
en 1983 avec 111 ug/m3 et en 1986 avec 117 ug/m3. Les in- 
dices les plus hauts apparaisent pendant les mois d'Aofit 
et d'Octobre. 

Source: Le NO2 provient principalement des industries 
et des véhicules 4 essence; Fig. 9. 


MONOXIDE DE CARBONE (CO) 

Les mesurages ont été effectué dans la Station 1, 
aire centrale de la ville, au moyen d'un Detector Ecolyzer 
(1977-1980) et un Detector Beckman, Modéle 866 (1981-1986). 
Le mesurage est faite pendant les 24 heures, tous le jours. 

La Norme de Qualité de 1'Air au Chili pour le CO est 
35 ppm/1 heure (1 fois par an) ou, 9 ppm/8 ‘heures (1# fois 
par an). 

Dans les années 1979 et 1986 ont a registré, en moyen- 


415 


STATION 


1 


2s 


15 
16 


Fig. 8.- Réseau de Vigilance de Qualité de 1'Air. 
Distribution des Stations 


COMUNE 


Santiago 
Santiago 
Santiago 
Santiago 


Santiago 
Providencia 


Santiago 
Maipu 
San Miguel 
Nufioa 
Conchali 
Pudahuel 


Maipt 
La Granja 


La Florida 
Las Condes 


DIRECTION 


Agustinas/Estado 
Santo Domingo 978 
Mac Iver 541 
Santo Domingo 2398 


Avda. Matta 576 
P. de Valdivia 963 


Independencia 939 
5 de Abril 4936 
Ismael Valdés 2020 
Marathon 1000 
Independencia 3450 
Teniente Cruz 1087 


Avda. Cerrillos 674 
La Pintana 


Vic. Mackenna 7196 
Las Condes/Las Clarisas 


GrEU 


Iglesia 
San Agustin 
Comandancia de 
Bomberos 
Ministerio de 
Salud 
Inst. de Salud 
Ocupacional 
72 Cia. de Bomberos 
I. Municipalidad 
de Providencia 
Esc. Salud Publica 
Inst. Comercial N2 7 
32 Cia. de Bomberos 
Inst. de Salud Publica 
12 Cia de Bomberos 
Consultorio 
Carlos Avendafno 
22 Cia. de Bomberos 
Cooperativa 
José Maza 
92 Cia. de Bomberos 
202 Cia. de Bomberos 


416 


ne 35.6 et 36.4 ppm/1l heure, respectivement. Les indices 
dans les autres années étaient inférieures a la norme. En 
1978 on a registré 14 ppm/1 heure, le plus bas, en 1983 et 
en 1984 moins que 20 ppm/1 heure. 

Les variations horaires du CO sont associées au flux 
véhiculaire et aux conditions climatiques. En Hiver,; ete 
CO se présente en plus grande quantité dai a sa moindre 
diffusion dans l'atmosphére. 

Ce contaminant constitue un des plus grands risques 
sanitaires par tout le monde. 

Source: Le CO provient spécialement des véhicules 4a 
essence; Fig. 10. 


ANHYDRIDE SULPHUREUX (S02) 

La mesurage a été effectué avec un appareil semi- 
automatique de gazes RAC, Modéle PV. Dans l'analyse on 
emploie la méthode colorimétrique de la p-roseaniline. 
L'echantillonage est permanent dans les 24 heures, deux 
fois par semaine, selon calendrier établi au commencement 
de l'année. 

La Norme de Qualité de l'Air au Chili pour le S02 est 
80 ug/m3 (moyenne annuelle) ou, 365 ug/m3 pour 24 heures 
firtors: parvaniy: 

Dans la période 1977-1986 on ne registre pas des va- 
leurs, moyenne annuelles, sur la norme. La Station 2 regis- 
trait 51 ug/m3, ~L'indice “le’*’plus’ haut, ‘correspondamoe 
l'aire centrale de la ville. Les Stations 2, S eG@me ue 
registraient indices entre 35 et 42 ug/m3 dans des secteurs 
de haut flux véhiculaire et probablement avec des batiments 
au chauffage a mazut. Les indices les plus hauts se regis- 
trent en hiver. 

Source: Combustion de mazut provenant des chauffages, 
quelques véhicules et fours; Fig. 11. 


POUSSIERE EN SUSPENSION 

L'obtention de la poussiére en suspension est effectué 
avec l'instrument High Volume, RAC, Modéle GMW-2000. Les 
échantillons sont analysés gravimétriquement. L'obtention 
est continue pendant les 24 heures, deux fois par semaine, 
selon calendrier établi au commencement de l'année. 

La Norme de Qualité de l1'Air au Chili pour la Poussié- 
re en Suspension est de 260 ug/m3 (moyenne annuelle) pour 
24 heures (1 fois par an) ou, 75 ug/m3 moyenne geométrique 
annuelle. 

Les Stations 2, 11 et 12 ont dépassé la norme avec une 
moyenne pour la période 1977-1986, de 330, 290 et 320 ug/m3 
respectivement. La Station 2 est situé dans l'aire centrale 


417 


wena ville ou il ywalun- haut. fiux véhiculaire.,. Les ‘Sta- 
moisesul 6t i2 sont loin. du centre ville. et. proche aux 
espaces ouvertes et aux centres miniéres exposés a l'action 
e6lique. Les Stations 2' et 6 sont dans des aires de haut 
flux véhiculaire et la Station 13 proche aux aires inha- 
bitées et a l'action edlique. 

pource. BACUIVi tes: industrielles; dey la. constrution, 
l'usage des pneumatiques, mouvement des autobus et des 


Camions. Pig. 12. 


POUSSIERE SEDIMENTAIRE 

Pour l'echantillonage de la poussiére sédimentaire on 
utilise un depét de materiel polimérique de 14 cm de lumié- 
re et 40 cm de haut par des périodes de 30 jours. Le depét, 
auquel on a mis de la protection contre les oiseaux, est 
exposé a 3 m hauteur. La poussiére sédimentée est extraite 
du cylindre avec de l'eau distilée et avec de l'eau démi- 
neralisée. Les paticules sont separés au moyen du papier 
filtre, san insectes ou des morceaux des plantes ou d'au- 
tres matiéres. Le papier filtre est analysé par fluores- 
cence des rayons X dans un spectrométre EDAX/PHILIPS. 

La Norme de Qualité de l1'Air recommendée par le WHO, 
adoptée au Chili pour la Poussiére Sédimentaire est 0.5 
ug/cem2/30 jours (moyenne annuelle). 

Dans*les Stations: 8, 9, 10; 11, .12- et: 15, la moyenne 
dans la période 1977-1986, la poussiére sédimentaire de- 
passait la norme. La Station 12 registrait une moyenne de 
0.85 ug/cm2, l'indice le plus haut. La plus petite quantité 
de poussiére sédimentaire fit registrée dans les Stations 
‘ere oe avec 0.4 ug/em2. Les indices les plus. hauts se 
trouvent dans les mois d'Octobre a Mars, les mois moins 
pluvieux. 

Source: Activités industrielles, combustion de dispo- 
sitif domestiques, activités de la construction, activités 
miniéres, mouvement des autobus et camions et le vent qui 
transportent la poussiére; Fig. 13. 


DIVISION GEOGRAPHIQUE DE LA PROV. DE SANTIAGO 
La Province de Santiago a une surface de 2.000 km2 et 
une population de 3.672.374 habitants. Géographiquement, 
a été divisé en 32 comunes, dans 11 desquelles on a récol- 
tée des lichens (Fig. 14), dans les autres comunes fit im- 
possible effectuer des récoltes lichéniques. 


418 


90 


60 


ug/m3 


30 


O 
2225) 610 t, 12 01s 1415 


IPS tb GiGi] Gm es 


Fig. 9.- Concentration moyenne 
annuelle de Oxide de Nitrogéne, 
selon Station. Période 1977-1986 


60 


O 
DED 5 aS OM! Ieatie wlan 40 


eB te Gi ene s 


Fig. 11.- Concentration moyenne 


78 79 80 81 82 83 84 85 86 


Anos 


Fig. 10.- Concentration maxime de 
Monoxide de Carbone par heure, mo- 
yenne annuelle, dans le Station 1. 


Période 1978-1986 
400 


Ag/m3/ 24hrs 


22) 5..6. 10011) ieuianee 


E Sateavce-Onniens 


Fig. 12.- Concentration moyenne 


annuelle de Anhydride Sulphureux, annuelle de Poussiére en Suspension, 


selon Station. Période 1977-1986 


1.0 


0.8 


ug/cm2/30 dias 
ie) 
a 


O 


selon Station. Période 1978-1986 


OARS 67) COO OMIA alo TI 4a 1omI6 


ErsuteacrOnnnems 


Fig. 13.- Poussiére Sédimentaire selon Station. Période 1977-1986 


419 


Fig. 14.- Province de Santiago. Les lichens firent récoltées dans 
les comunes hachées: 1.- Comuna de Huechuraba; 2.- Comuna de Recoleta; 
3.- Comuna de Quinta Normal; 4.- Comuna de Santiago; 5.- Comuna de Pro- 
videncia; 6.- Comuna de Las Condes; 7.- Comuna de Nufioa; 8.- Comuna 
de La Reina; 9.- Comuna de Lo Espejo; 10.- Comuna de La Cisterna; 11.- 
Comuna de La Granja. 


DISTRIBUTION DES LICHENS DANS LES DIFFERENTES 
COMUNES ETUDIEES 


BUTE tet fie U'S CUA AN Yi eA BR. 


COMUNA DE HUECHURABA: Ave. Circunvalacién Américo Ves- 
pucio, sur l'écorce de Robinia, 1984. COMUNA DE RECOLETA: 
Estadio Recoleta, sur l'écorce de Robinia, 1984. COMUNA 
DE LA GRANJA: La Castrina, sur un poteau en Nothofagus, 
Poof rig. 15a. 


PALOPLACASCERINA .QHHRHe ier HEDWs|), TH. ER: 


COMUNA DE HUECHURABA: Ave. Circunvalacién Américo Ves- 
pucio, sur l'écorce de Robinia, 1984. COMUNA DE LA CIS- 
TERNA: Gran Avenida, Paradero 28 1/2, sur un poteau en 


420 


Nothofagus, 1968; Fig. 15b. 


CANDELARIELLA VITELLINA (HOFFM.) MULL.- ARG. 


COMUNA DE HUECHURABA: Ave. Circunvalacién Américo Ves- 
pucio, sur l'écorce de Robinia, 1984. COMUNA DE LA CISTER- 
NA: Gran Avenida, Paradero 28 1/2, sur un poteau en Notho- 
fagus, 1968; Fig. 15c. 


GASPARRINIA MICROPHYLLA (HUE) DODGE 


COMUNA DE HUECHURABA: Ave. Circunvalacién Américo Ves- 
pucio, sur l'écorce de Robinia, 1984. COMUNA DE RECOLETA: 
Estadio Recoleta, sur l'écorce de Robinia, 1984. COMUNA DE 
QUINTA NORMAL: Quinta Normal, sur 1'écorce de Platanus, 
1984. COMUNA DE SANTIAGO: Cerro San Cristébal, Plaza Ga- 
briela Mistral, sur l'écorce de Robinia, 1987; Rue Nata- 
niel 1777, sur l'écorce de Robinia, 1984; Plaza de Armas, 
sur l'écorce d'Acacia, 1985; Plaza Brasil, sur l'écorce de 
Gymnocladus, 1984; Parque O"HIggins, sur l'écorce de Fra- 
xinus, 1987; Parque Forestal, Ave. Cardenal Caro/Puente 
Loreto, sur l'écorce de Platanus, 1984. COMUNA DE PROVI- 
DENCIA: Rue Clemente Fabres 1040, sur une colonie de Tor- 
tula, 1984. COMUNA DE LAS CONDES: Rue Tomas Moro 261, sur 
l'écorce de Prunus, 1985; Rue San Pascual 59, sur l'écorce 
d'Ulmus, 1984. COMUNA DE NUNOA: Rue Las Dalias/Exequiel 
Fernandez, sur l'écorce de Robinia, 1984; Rue Las Palmeras 
3395, sur l'écorce d'Acer, 1987; Rue Los Alerces 2726, sur 
l'écorce d'Acer, 1984. COMUNA DE LA REINA: Rue Julia Bers- 
tein s/n, sur l'écorce de Robinia, 1984; Rue Alvaro Casa- 
nova s/n, sur l'écorce de Robinia, 1986. COMUNA DE LO ES- 
PEJO: Rue Colén 70, sur l'écorce de Robinia, 1986. COMUNA 
DE LA CISTERNA: Gran Avenida, Paradero 28 1/2, sur un po- 
teau en Nothofagus, 1968; Rue 21 de Mayo O70A, sur 1'écor- 
ce de Prunus, 1986; Fig. 15d. 


LECANIA CHILENA DODGE 


COMUNA DE LA CISTERNA: Gran Avenida, Paradero 28 1/2, 
sur un mur en mortier, 1967; Fig. 15e. 


LECANORA DISPERSA (PERS.) SOMMERF. 


COMUNA DE HUECHURABA: Ave. Circunvalacién Américo Ves- 
pucio, sur l'écorce de Robinia, 1984. COMUNA DE QUINTA 
NORMAL: Quinta Normal, sur l1'écorce de Platanus, 1984; 
Ibid., sur l1'écorce de Magnolia, 1984; Ibid., sur l'écorce 
de Grevillea, 1984; Ibid., Hospital J.A.Rios, sur un mur 
en mortier, 1984; Municipalidad de Quinta Normal, Parque 


421 


Lo Franco, sur l'écorce d'Araucaria, 1984. COMUNA DE SA- 
TIAGO: Parque O"Higgins, sur l'écorce d'Ulmus, 1987; Ibid., 


sur l'écorce de Ligustrum, 1987; Ibid., sur le stipe de 
Washingtonia, 1987; Ibid., sur l'écorce de Fraxinus, 1987; 
Ibid., sur l'écorce de Robinia, 1987; Ibid., sur 1l'écorce 


de Cryptocarya, 1987; Ibid., sur l'écorce de Schinus, 1987; 
Ave. Ricardo Cumming 350, sur l'écorce d'Erythrina, 1984; 
Parque Forestal, Ave. Cardenal Caro/Puente Loreto, sur 1'é- 
corce de Platanus, 1984; Ibid., au coté Nord du Museo de 
Bellas Artes, sur des racines de Washingtonia, 1984; Cerro 
Santa Lucia, sur un mur en mortier, 1968; Plaza Hermanos 
Matte, sur l'écorce de Robinia, 1984; Plaza de Armas, sur 
l'écorce d'Acacia, 1985; Plaza Brasil, sur l'écorce d'Ery- 
thrina, 1984; Ibid., sur le stipe de Phoenix, 1984; Ibid., 
sur l'écorce de Populus, 1984; Plaza Santa Lucia, sur 1'é- 
corce de Cedrus, 1984; Parque Bustamante 1001, sur 1'écor- 
ce d'Ulmus, 1988; Ave. Matta 336, sur l'écorce d'Ulmus, 
1988. COMUNA DE PROVIDENCIA: Plaza Guillermo Frencke, 
pune ecorce d'Acacia; 1987; Rue Roman Diaz 1935, sur 1'é- 
corce de Platanus, 1984. COMUNA DE LAS CONDES: Rue San 
Pascual 59, sur l1'écorce d'Ulmus, 1984. COMUNA DE NUNOA: 
Rue Palqui 2990, sur un mur en mortier, 1984; Rue Dr. Jo- 
how/Ave. Grecia, sur des branches d'Acer, 1984; Rue Exe- 
quiel Fernandez/Los Alerces, sur des branches d'Acer, 1984; 
Rue Las Palmeras 3395, sur l'écorce d'Acer, 1987; Rue Vir- 
ginio Arias 1350, sur un mur en mortier, 1984; Rue Pedro 
Torres 775, sur des branches d'Acer, 1984; Rue Las Dalias 
1809, sur l'écorce de Robinia, 1984; Rue Los Jardines 207, 
sur l'écorce de Citrus, 1988; Fig. 15f. 


LEMPHOLEMMA SP. 


COMUNA DE HUECHURABA: Ave. Circunvalacién Américo Ves- 
pucio, sur l'écorce de Robinia, 1984. COMUNA DE SANTIAGO: 
Cerro Santa Lucia, sur un mur en mortier, 1984. COMUNA DE 
NUNOA: Rue Virginio Arias 1350, sur un mur en mortier, 
1984; Ibid., 1308, sur l'écorce d'Acer, 1988. COMUNA DE 
LA CISTERNA: Gran Avenida, Paradero 28 1/2, sur un poteau 
en Nothofagus, 1968; Fig. 16a. 


PARMELIA SP. 


COMUNA DE RECOLETA: Estadio Recoleta, sur l'écorce de 
Robinia, 1984. COMUNA DE SANTIAGO: Parque Forestal, Ave. 
Cardenal Caro/Puente Loreto, sur 1'écorce de Platanus, 1984 
Cerro San Cristébal, sur 1'écorce de Robinia, 1987. COMUNA 
DE NUNOA: Rue Las Palmeras 3395, sur l'écorce d'Acer, 1987. 
COMUNA DE LA REINA: Rue Alvaro Casanova s/n, sur 1'écorce 


423 


de Robinia, 1984; Fig. 16b. 


Poeseca A ADSCENDENS (TH. FR.) OLIV. ‘emend.: BITT. 


COMUNA DE HUECHURABA: Ave. Circunvalacién Américo Ves- 
pucio, sur l'écorce de Robinia, 1984. COMUNA DE RECOLETA: 
Estadio Recoleta, sur des branches de Robinia, 1984. COMUNA 
DE QUINTA NORMAL: Quinta Normal, sur l1'écorce de Platanus, 
1984. COMUNA DE SANTIAGO: Plaza Brasil, sur l'écorce de 
Gymnocladus, 1984; Plaza de Armas, sur l'écorce d'Acacia, 
1985. Parque Forestal, Ave. Cardenal Caro/Puente Loreto, 
sur l'écorce de Platanus, 1984; Parque O"Higgins, sur 1'é- 
corce de Fraxinus, 1987; Ave. Viel 1462, sur des branches 
de Vitis, 1967. COMUNA DE NUNOA: Rue Dr. Johow/Ave. Grecia, 
sur des branches d'Acer, 1984; Rue Las Dalias/Los Alerces, 
sur l'écorce de Robinia, 1984; Rue Pedro Torres 775, sur 
l'écorce d'Acer, 1984; Rue Las Palmeras 3395, sur l'écorce 
d'Acer, 1987. COMUNA DE LA REINA: Rue Alvaro Casanova s/n, 
sur l'écorce de Robinia, 1984. COMUNA DE LO ESPEJO: Rue 
Colén 70, sur l'écorce de Robinia, 1985. COMUNA DE LA CIS- 
TERNA: Gran Avenida, Paradero 28 1/2, sur un poteau en No- 
thofagus, 1967; Rue 21 de Mayo O70A, sur l'écorce de Pru- 
mus 006. Fig: 16c: 


RAMAGINA  ECKLGONIT (SPRENG.) MEY. et FLOT. 


COMUNA DE RECOLETA: Estadio Recoleta, sur 1l'écorce de 
Robinia, 1984. COMUNA DE LA REINA: Rue Alvaro Casanova s/n, 
sur l'écorce de Robinia, 1984. COMUNA DE LA CISTERNA: Gran 
Avenida, Paradero 28 1/2, sur un poteau en Nothofagus, 
1968; Fig. 16d. 


Mero SCCHT Siw oOeCHRYSOORPHTHALMUS “(D)” TRE EER: 
COMUNA DE LA REINA: Rue Alvaro Casanova s/n, sur 1'é- 
corce de Robinia, 1984; Fig. 16e. 
USNEA IGNIARIA MOT. 
COMUNA DE LA REINA: Rue Alvaro Casanova s/n, sur 1'é- 
corce de Robinia, 1984; Fig. 16f. 
AKANE HOR LAY CARL BANA a{ ee) the PRS 


COMUNA DE SANTIAGO: Cerro San Cristébal, Plaza Gabrie- 
la Mistral, sur 1'écorce de Robinia, 1986. COMUNA DE NUNOA: 
Rue Las Palmeras 3395, sur l'écorce d'Acer, 1987. COMUNA 
DE LA REINA: Rue Alvaro Casanova s/n, sur l1'écorce de Ro- 
binia, 1984; Fig. 17a. 


425 


AANTHORTA?) POLYCARPA’ (HOFEM JV OLEV: 


COMUNA DE HUECHURABA: Ave. Circunvalacién Américo Ves- 
pucio, sur l'écorce de Robinia, 1984. COMUNA DE RECOLETA: 
Estadio Recoleta, sur l'écorce de Robinia, 1984. COMUNA 
DE NUNOA: Rue Rodrigo de Araya/Exequiel Fernandez, sur 
1'écorce de Prunus, 1972; Rue Dr. Johow/Ave. Grecia, sur 
l'écorce d'Acer, 1984; Rue Pedro Torres 775, sur l1'écorce 
d'Acer, 1984. COMUNA DE LA REINA: Rue Alvaro Casanova s/n, 
sur l'écorce de Robinia, 1984. COMUNA DE LA CISTERNA: Rue 
21 de Mayo 070A, sur l'écorce de Prunus, 1986; Fig. 17b. 


a) Xanthoria parietina b) Xanthoria polycarpa 


Fig. 17a-b.- Distribution des espéces lichéniques dans les diffé- 
rentes comunes de la ville de Santiago. Les points noirs indiquent les 
endroits de récolte de chaque espéce. 


Un résumé de la distribution des lichens et ses phoro- 
phytes est exposé dans le Schéme 1. 


D'APRES LA POLLUTION ET LA DISTRIBUTION DES LICHENS 
RECOLTEES DANS LA VILLE DE SANTIAGO, ON OBSERVE QUE: 


1.- Comme conséquence de la situation géographique 
de la ville de Santiago, dans une dépression entourée par 
des cordons montagneux hauts, les contaminants restent 
retenus d'une maniére permanente sur la ville. L'effet 
du vent est inefficace pour transporter la pollution a 
d'autres endroits. 


2.- Les contaminants mesurés sont, en général, moin- 
dres que la norme. La poussiére en suspension et la pous- 
siére sédimentaire dépassait la norme, en moyenne, dans 
3 et 6 endroits respectivement. 


426 


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ee a a OO ee eS 


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Fi 
a 


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ct (ag ct yp el) Gees [tel eG ee ccd ee te ean | her em me eS Vee) a OPT a Te) Se) 
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be | Q) aolol<{|a};}on!;o}; O;C | xX) e| aprile; c}l]r-|o 7/0] 0 
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4) 3 OF Ol OS Worl |S O 3/Q/q0 VQIdnIe-|}O};aO;CI]C!]o® 
(e DIJDIeR | an Ol Oye | Ssye|}]o;}al;a (e Q <elLIL© OF ik 
Fa =} Hiei hie | oO; ClTOIO]LII/DI axl KlOI aTSsIToOos]oa; oa; aQalra!l|o;alo 
Oo je) VIM ODIO rl CI /SI/XInxilie|}oal;oatc Oue@rliet SO} Sl OS | Oris 
ct — a);0O/0/Q KTaO OInI I DII/OLSQ rol aie |/AI;O anlar 
3 oO O ay = * 5 24 su Ged Heyl baa ar olaloloal™ o = S3LAHdONOHd 
le ry 
h oye Qo a MOIS) AlCl]e|4t+]}/oa}oa]o +] O 
i) cir a) —1OleK|S;o;oa];a] 1s m1 > 
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Cc +| O @) O;o|] 74 Q) (1) @ 
09) Qo}; 5 te Q) te im 
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427 


3.- Buellia fuscula, Caloplaca cerina, Candelariella 
vitelina, Lecania chilena, Teloschistes chrysophthalmus 
et Usnea igniaria, etaient dans des aires é@loignées du 


centre ville, aqui est l'aire avec la plus intense 
pollution. 
4,.- Gasparrinia microphylla, Lecanora dispersa et 


Physcia adscendens, sont les espéces les plus répandues 
Gans,ia ville. 


5.- Des 14 espéces observées, 9 se trouvent dans la 
Comune, cde La Reina, vers le limit suburbain a 260 m plus 
haut que les plus grand des autres altitudes régistrées. 
C'est un endroit avec une atmosphére plus propre et plus 
humide. 


6.- Dans l1'écorce de Robinia pseudo-acacia, on a trou- 
vé 13 des 14 espéces lichéniques récoltées (92.9%) et 3 
espéces sont trouvées, chacune, dans un seule type de subs- 
tratum. 


7.- Dans toutes les comunes revisées on a trouvé des 
lichens. 
8.- La polltuion atmosphérique de l'aire centrale de 


la ville n'empéche pas l'accroisement des lichens. Les 
thalles de Gasparrinia microphylla récoltées dans la Plaza 
de Armas (centre ville) se présentent granuleuses et pres- 
que détruites. 


REMERCIEMENTS 
Je remerci’ le Prof. Marcelo Campos, la revision du 
texte et la lecture critique du manuscript. 


BIBLIOGRAPHIE 

1.- Comisién Nacional de la Reforma Administrativa. 
1981. Reformulaci6én Comunal en la Regidén Metropolitana 
de Santiago, 119 pp. 

2.- Comité Panamericano de Geografia Urbana, Instituto 
Panamericano de Geografia e Historia. 1985. Metrdpoli de 
Santiago. Documento preparado en ocasiédn del Seminario 
Internacional de Métodos Geograficos para el Analisis Ur- 
DanOnr i Dp. 

3.- Direcci6én Meteorolégica. Ministerio de Defensa 
Nacional. 1975. Informe Climatolégico del Area Metropoli- 
tana de Santiago, 49 pp. 

Pe eembouce iC. Sy Mee Mahu. 1972. Flora Liquénica”™ de 
Chile, Manuscrito, 1405 pp. 

pee unk anewe ekee o/O.n Introductiom tom pritishn Li- 


428 


chene.) - Tlie Buncle. & Co.) Ltd.; Printers. and *Pupiiienere 
202) pps), be Srp. . 

6.- Hale, M. E. 1970. The Biology of Lichens. Edward 
Arnold (Publishers) Ltd. London 176 pp. 

7.-?Hawksworth;: °D.. L. & F. Rose. 1970. \QUaniataeae 
scale for estimating sulphur dioxide air pollution in 
England and Wales using epiphitic lichens. Nature 227: 
145-148. 

8.- Looney, J/ Al, CC.  E.. Weber, E..' Nieboer sama 
Stetsko & K. A. Kershaw. 1968. Interrelationship between 
concentration of Cs and various stable elements in three 
lichens species. Halth Physics Official Journal of the 
Heath Physics Society 50(1): 148-152. 

9.-..Préndez, M., J. Lb. Ortiz, E. Cortés ‘andayageaeeeee 
la. 1984. Elemental Composition of Airborne Particulate 
Matter. ,from Santiago City, Chile, .1976.. Journaiieeseen= 
Air Pollution Control Association 34(1) 54-56. 

10.- Servicio de Salud del Ambiente. Ministerio de 
Salud. 1980-1986. Informe Anual de Contaminacién Atmos- 
férica’(1980;.. 11-1473" 1981: 1-7/1; 1982: 2-20 gee 
1-127; 1984: 1-82; 1985: 1-134; 1986: 1-71). 

11.- Thompson, J. W. 1963. The Lichen genus Physcia 
in North America. Nova Hedwigia 7: 1-172, 47 maps and 
25 pl. Verlag von Kramer, West Germany. 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 429-440 January-March 1989 


STUDIES ON XYLOPHILOUS FUNGI FROM ARGENTINA. IV. 
ANAMORPHS OF BASIDIOMYCETES ON EUCALYPTUS VIMINALIS (MYRTACEAE )* 


A.I. Romero, D. Cabral & S.E. Lopez 


Dto. de Ciencias Biolégicas, Facultad de Ciencias Exactas y Naturales, 
Universidad de Buenos Aires, 1428 Buenos Aires, Argentina. 


ABSTRACT 


Eight anamorphs of lignicolous Basidiomycetes are described. Among 
them three new genera: Fibulochlamys, Corticomyces and Glomerulomyces 
are proposed with their respective species: F. ferruginosa, C. xenasma— 
toides and G. fibulosus. One new species: Allescheriella cyanea is pro- 
posed. Haplotrichum curtisii, H. gracile and Sporotrichum aurantiacum 
are first records for Argentina. The position of “Sporotrichum" azureum 
is open to discussion. 


INTRODUCTION 


During an ecological and systematic survey of fungal communities 
on standing trees, logs and stumps of Eucalyptus viminalis Labill. in 
the province of Buenos Aires (Lopez, 1983; Romero, 1983; Lopez, 1984; 
Lopez & Wright, 1985) a number of anamorph samples were collected which 
revealed either the presence of clamp-connections on the vegetative 
or conidiogenous hyphae, or a close affinity with Basidiomycetes as 
recorded in the literature. 

In spite of the increasing interest showed in the last years on 
the mitospores in Basidiomycetes (Kendrick & Watling, 1979; Stalpers, 
1984; Buchalo et al., 1985) most workers are basidiomycetologists more 
interested with the teleomorph than the anamorph, describing it poorly. 
Nevertheless, the number of new basidiomycetous anamorphs increases 
year by year (Nag Raj, 1978; Dyko & Sutton, 1979; Sivanesan & Watling, 
1980; Gochenaur, 1981; Wright, 1983; Stalpers & Vlug, 1983; Pantidou 
et al., 1983; Udagawa et al., 1986) resulting in the need of descrip- 
tions on the basis of modern conidiogenic concepts and recognition of 
the generic and specific anamorphic taxa in order to allow an orderly 
systematic disposition of those forms which may lead to a better under- 
standing of the group as a whole. 


*Research funded by the Consejo Nacional de Investigaciones Cientifi- 
cas y Técnicas (CONICET), Argentina. Publication N2 29 of the PRHIDEB 


430 


DESCRIPTIONS OF THE TAXA 
1. Fibulochlamys Romero & Cabral, gen. nov. 


Hyphis hyalinis, fibulatis. Conidiophoriis absentibus vel microne- 
matosis, simplicibus vel rarae ramosis. CellTulis conidiogenis cylindri- 
cis, terminalibus vel subinde intercalaribus, hyalinis, fere semper 
in fibula basali consociatis. Conidiis holothallici, globosis vel 
ellipsoideis vel pyriformibus, aseptatis, pallide flavis, laevibus cra- 
ssissime tunicatis, umbonatis, interdum pedicellatis. 


Species typica: Fibulochlamys ferruginosa Romero & Cabral 


Etymology: Fibula, Latin for clamps; Chlamys, Greek for "mantle" enve- 
loping conidia. 


Hyphae hyaline, with clamps. Conidiophora absent or micronematous 
simple or seldom branched. Conidiogenous cells cylindric, terminal or 
intercalary, hyaline, almost always associated with a basal clamp. 
Conidia holotallic, globose ellipsoidal or pyriform, sometimes pedi- 
cellate, O-septate, pale yellow, very thick-walled. 


Type species: Fibulochlamys ferruginosa Romero & Cabral 


l.a. Fibulochlamys ferruginosa Romero & Cabral, sp. nov. (Fig. 1: A-C) 


Coloniae ad corticem et lignun 2-3 cm in longitudine, resupina- 
tae, adpressae, aspectu crystallino sub lente, in sicco ferrugineo 
pallido. Mycelio hyalino, fibulato, hyphis cylindricis, 3-4 um diam. 
et hyphis e cellulis brevibus et irregularibus composito. Conidiopho- 
riis absentibus vel micronematosis, simplicibus vel rarae ramosis. 
CelTulis conidiogenis cilindricis, terminalibus vel rarae intercalari- 
bus, fere semper consociatis in fibula basali. Conidibus holothallicis, 
pallide flavis, globosis, 12-18 um diam, ellipsoideis, l0-l2Z=xe20=23 
um, vel globosis pedicellatis, umbonatis, aseptatis, continentibus gra- 
nulosis cum 2-3 gutulae, dextrinoidae. 


HOLOTYPUS: Argentina, Buenos Aires, Ramallo; leg.: Romero, Cabral & 
Lopez, VIII-1982, ad partem amputatam trunci Eucalypti viminalis, in 
Herb. BAFC N° 30826 conservatum est. 


Etymology: ferruginosus Latin for the rusty colour of the colonies 
Teleomorph: unknown 


Conidiomata on bark and wood, 2-3 cm in length, resupinate, tight, 
like crystals of sugar under a lens, light ferrugineous when dry (M 
& P, Pl.12, 10 K, "Chinese Gold"). Hyphae hyaline, with clams present 
at all prymary septa, composed of cylindric hyphae 3 um diam and irre- 
gular hyphae consisting of short cells. Conidiophores absent or micro- 
nematous, not branched or rarely so. Conidiogenous cells cylindric, 
terminal, generally associated with a basal clamp, or intercalary. 
Conidia holotallic, light yellow, globose, 12-18 um diam, pyriform 
or ellipsoidal, 10-12 x 20-23 um or sometimes pedicellate, often with 
an umbo at both end as a result of a subapical widening of the conidio- 


431 


genous cell, O-septate, thick-walled, wall 2-3 um thick when mature, 
dextrinoid, with 1-3 oildrops. 


Material studied: the holotype. 


Remarks: Despite a number of Basidiomycetes having "chlamydospore" type 
of conidia related with the basidiocarp or described in culture, it 
is rare to find only chlamydospores in nature, without any associated 
basidiocarp. Because no teleomorph was detected in this material, we 
think it is necessary to describe as an independent new form-genus. 

With the date available we are not be able to state if Fibulochla- 
mys is the anamorph of other Basidiomycetes with chlamydospore-like 
structures but this should be a subject of further studies. 

This specimen has been illustrated and described by Lopez (1984) 
who included it in the Allescheriella type, according to Kendrick & 
Watling (1979). But on account of its holothallic ontogeny, we consider 
it a new genus. 

On the other hand, it is closely related to the genus Pagidospora 
Drechsler, but it differs because of the lack of the trapping struc- 
tures which is the main characteristic of this genus (Drechsler,1960). 


2. Corticomyces Romero & Lopez, gen. nov. 


Hyphis hyalinis, fibulatis, paucis. Conidiophoriis absentibus. 
Cellulis conidiogenis blasticis, cylindricis vel indistinctis. Conidiis 
holoblasticis, rhexolyticis, ellipsoidis, hyalinis, aseptatis, tenui- 
tunicatis, laevibus. 


Species typica: Corticomyces xenasmatoides Romero & Lopez. 


Etymology: Cortex Latin for bark; myces Greek for fungus. 


Hyphae hyaline, with clamps, scanty. Conidiophores absent. Coni- 
diogenous cells blastic, cylindric or undifferentiated from the vege- 
tative cells. Conidia holoblastic, rhexolytic secession, ellipsoidal, 
hyaline, O-septate, thin-walled, smooth. 


Type species: Corticomyces xenasmatoides Romero & Lopez 


2.a. Corticomyces xenasmatoides Romero & Lopez, sp. nov. (Fig.1: D-F) 


Coloniae ad corticem et lignum, resupinatae, araneosae et prui- 
nosae, aspectu xenasmatoide, albae vel griseae. Mycelio inconspicuo, 
e hyphis tenuis, 0.5-1.5 um diam, fibulatis, paucis composito. Conidio- 
phoriis absentibus. Cellulis conidiogenis blasticis, cylindricis vel 
indistinctis, 10-20 um longitudine. Conidiis holoblasticis, rhexo- 
lyticis, ellipsoideis, hyalinis, aseptatis, tenuitertunicatis, laevi- 
bus, interdum guttulatis, 6-9 x 4-7 um. 


; i i leg.: Romero, 
HOLOTYPUS: Argentina, Buenos Aires, Gobernador Castro, ome 
Cabral & <a V-1983, ad partem amputatam trunci Eucalyptus vimina- 
lis, in Herb. BAFC N° 30827 conservatus est. 


Etymology: xenasmatoides Latin for resembling the basidiocarp of the 


432 


genus Xenasma. 
Teleomorph: unknown. 


Conidiomata on bark and wood, resupinate, effuse, arachnoid and 
pruinose, with xenasmatoid aspect, greyish white. Mycelium scanty, 
hyphae thin, 0.5-1.5 um diam, with clamps. Conidiophores absent. Coni- 
diogenous cells blastic, cylindrical or undifferentiated from the vege- 
tative cells, 10-20 um long. Conidia holoblastic, rhexolytic secession, 
ellipsoidal,hyaline, O-septate, thin-walled, smooth, 6-9 x 4-7 um, 
sometimes guttulate, always terminal. 


Material studied: the holotype. Argentina, Buenos Aires, Gobernador 
Castro, JTeg.: Romero, Cabral & Lopez, II-1982 (BAFC N° 30828); ibid, 
Ramallo, V-1982 (BAFC N° 31272). 


Remarks: it grows on ME with similar characteristics as in nature. 

One could find some similarities between Corticomyces conidia and 
the chlamydospores produced in culture by some Basidiomycetes .However, 
such conidia do not have the typical chlamydospore wall and cytoplasma- 
tic content to be considered a resting spore. 


3. Glomerulomyces Romero & Lopez, gen nov. 


Hyphis hyalinis, fibulatis. Conidiophoriis semimacronematosiis, 
simplicibus vel ramosis, fibulatis. Cellulis conidiogenis mono-poly- 
blasticis, ellipsoideis, terminalibus, hyalinis. Conidiis holoblas- 
ticis, globosis, hialinis, aseptatis, tenuitertunicatis, Taevibus gem- 
miferis, in capituliis globosis compactis. 


Species typica: Glomerulomyces fibulosus Romero & Lopez 


Etymology: Glomerulus Latin for the clusters of conidia into a head; 
myces Greek for fungus. 


Hyphae hyaline, with clamps. Conidiophores semimacronematous, 
simple or branched, with clamps. Conidiogenous cells mono-polyblastic, 
ellipsoidal, terminal, hyaline. Conidia holoblastic, spherical, hya- 
line, aseptate, smooth, budding, in globose compact heads, thin-walled, 


Type species: Glomerulomyces fibulosus Romero & Lopez 


3.a. Glomerulomyces fibulosus Romero & Lopez, sp. nov.(Fig. 1: G-K) 


Coloniae ad corticem et lignum, effusae, pelliculosae, farinaceae, 
albae calcareae vel cremeae. Mycelio hyalino, fibulato, hyphis 2.5-5 


FIGURE 1 : A-C : Fibulochlamys ferruginosa. A: Conidiogenous cell in 
different stages of development. B: Conidiophores with a young coni- 
dium. C: Conidia. D-F; Corticomyces xenasmatoides. D: Conidiogenous 
cells. E: Mycelium with clamps. F: Conidia. G-K: Glomerulomyces fibu- 
losus. G: Conidiophores with conidia in compact , globose heads. H:Co- 
nidiogenous cell with one budding conidium., I: Budding conigias 
J: Cluster of mature conidia. K: Conidium. 


434 


um diam composito. Conidiophoriis semimacronematis, simplicibus et ra- 


mosis, fibulatis, 30-40 x 2-4 um. Cellulis conidiogenis mono-polyblas- 
ticis, ellipsoideis, hyalinis, 8-10 x 4-5 um. Conidiis holoblasticis, 


globosis, hyalinis, aseptatis, 6-10 um diam, gemmiferis, in capituliis 
globosis compactis, 20-25 um diam. 


HOLOTYPUS: Argentina, Buenos Aires, Ramallo; leg.: Romero, Cabral & 
Lopez, VIII-1982, aa partem amputatam trunci Eucalyptus viminalis, in 
Herb. BAFC N° 30829 conservatus est. 


Etymology: fibula Latin for clamps 
Teleomorpn: unknown. 


Conidiomata on bark and wood, effuse, pelliculose, farinaceous, 
chalkly white to cream. Hyphae hyaline, with clamps, 2.5-5 um diam. 
Conidiophores semimacronematous, simple or branched, with clamps, 
30-40 x 2-4 um. Conidiogenous cells mono-polyblastic, ellipsoida, hya- 
line, 8-10 x 4-5 um. Conidia holoblastic, spherical, hyaline, aseptate, 
6-10 um diam, budding in several directions to produce compact globose 
heads, 20-25 um diam. 


Material studied: the holotype. 


Remarks: When first looked at, this fungus resembles Papulospora, 
Burgoa and allied genera, because the cluster of conidia appears as 
a bulbil. We rejected Papulospora because this is restricted to the 
anamorph of some Ascomycetes. On the other hand, the conidial structure 
in Glomerulomyces is not a bulbil as in Burgoa and allied genera. 
Budding and development in the former results in a compact conidial 
head with different number of cells and colour, whereas in the latter 
genus budding originates a pseudoparenchymatous structure called a 
"bulbil" (Weresub et al., 1971). 

In the slides, the heads of Glomerulomyces can be easily separeted 
in cells proving they are conidia rather than pseudoparenchymatous 
cells. 

We should take in account another bulbillous species, Aegerita 
state, which form a cluster of apically enlarged and radially arranged 
hyphae aS a prosenchymatous propagule (Kendrick & Watling,1979), but 
this is not our case. 


4.a. Allescheriella cyanea Cabral & Romero, sp. nov. (Fig. 2: A-D) 


Coloniae ad corticem et lignum, e parvis maculis conformatae, 
adpressae, interdum coalescentes in magna extensione, resupinatae, 
marginibus indefiniti, paginis iridiscentes, laeves vel tuberculatae, 
sub lente aspectu crystallino, resentes azureae virellae (M & P, Pl 
38 L-1 vel L-7) et margine albo, siccae virides. Mycelio paucis, hyphis 
tenuitertunicatis afibulatis e quibus conidiophora enascentibus, et 
hyphis tenuis, fibulatis, plus minusve parallelibus ad substratum, com- 
posito. Conidiophoriis mononematosis, micro vel semimacronematosis, 
simplicibus vel ramosis,flexuosis, hyalinis. Cellulis conidiogenis 
monoblasticis, cylindricis, terminalibus, interdum intercalaribus. 
Conidiis globosis, 13-20 um diam, subglobosis vel late ellipsoideis, 
13-15 x 17-22 (28) um, laeviter pigmentiferis, crassitunicatis 3-4 um 
crassis, stratosis, continentibus granulosis et guttulosis, dextri- 


435 


noideis. 


HOLOTYPUS: Argentina, Buenos Aires, Gobernador Castro; leg.: Romero, 
Cabral & Lopez, II-1982, ad partem ainputatam trunci Eucalyptus vimi- 
nalis, in Herb. BAFC N° 30830 conservatus est. 


Etymology: cyaneus Latin for the blue colour of the colony in nature. 
Teleomorph: unknown. 


Conidiomata on bark and wood, composed of small compact spots, 
coalescing into large areas, irregular, with indistinct margin, surface 
smooth or tuberculate, under a lens with the aspect of sugar crystals 
because of the conidial mass; when fresh dark greenish-blue (M & P, 
Pl. 38 L-7) and white margin, when dry green "Juniper" (M & P, Pl. 31 
H-4). Hyphae scanty, more or less parallel to the substrate, composed 
of thin walled hyphae without clamps from which the conidiophores 
arise, interlaced with thin clamped hyphae. Conidiophores mononematous, 
micro to seminematous, simple or branched, flexuose, hyaline. Conidio- 
genous cells monoblastic, integrate, discrete, terminal, occasionally 
intercalary, cylindrical. Conidia solitary, spherical 13-20 um diam, 
subspherical or widely ellipsoidal 13-15 x 17-22 (28) um, slightly 
pigmented, thick-walled 3-4 um thick, with several layers, contents 
granulose and with 2-3 large drops, with strong dextrinoid reaction. 


Material studied: the holotype. Argentina, Buenos Aires, Ramallo; leg.: 
Romero, Cabral & Lopez, I1-1983 (BAFC N° 30831); ibid, Gobernador Cas- 
tro, 1y-1982 (BAFC, N® 31273). 


remarks: this material has been previously illustrated and partially 
described by Lopez (1984). Despite the lack of the sporodochium and 
the presence of clamps it was included in the genus Allescheriella 
sensu Hughes (1951) and Ellis (1971) because of the conidial ontogeny- 
type and conidial morphology. 


5.a. "Sporotrichum" azureum Wright & v. Arx apud v. Arx, Persoonia 
TA 2G ACV 285 191 SeR RIG a2 ReE-Ge 


Teleomorph: unknown 


Material studied: Argentina, Buenos Aires, Gobernador Castro; leg.: 


Romero, Cabral & Lopez, V-1982 (BAFC N° 30832). 


Remarks: it was originally described from Argentina by Wright & v. Arx 
(von Arx, 1973). Kendrick & Watling (1979) suggested this anamorph 
would be better placed in Allescheriella. According to Stalpers (1984) 
this species belongs neither to Sporotrichum nor to Allescheriella. 
He believes it can be better placed in a genus of its own (pers. 
comm.), thus we have decided to maintain this genus name until 
Stalpers'revision is published. 


436 


6.a. Sporotrichum aurantiacum (Bull.: Fr.) Fr., Syst. Mycol. 3:423, 
1832. hig “2s Hed. 


Teleomorph: Pycnoporellus metamorphosus (Fuck.) Stalpers 


Material studied: Argentina, Buenos Aires, Gobernador Castro; leg.: 
Romero, Cabral & Lopez, V-1982, (BAFC N° 30833). 

Remarks: the species has been recently described by Stalpers (1984) 
giving a complete diagnosis of both anamorph and teleomorph states. 
Our sample agrees with Stalpers'description (op.cit.) except for the 
absence of chlamydospores and crystals. We also observed dextrinoid 
reaction whereas Stalpers mentioned red reaction in KOH. 


7.a. Haplotrichum curtisii (Berk.) Hol. Jechova, Ceska Mykol. 30: 3-4, 
1976, APiG.3.(A-D)2 


Conidiomata on bark, resupinate, velvety, elastic, margin indis- 
tinct, rusty paler towards the margin. Mycelium of hyphae up to 10 um 
diam, thick-walled, wall up to 1 um, from which arise the conidiopho- 
res. Conidiophores erect, branched in right angles, pale brown, sligh- 
tly tapering towards the tip. Conidiogenous cells polyblastic, inflated 
terminal or intercalary, bearing several denticles. Conidia holoblas- 
tic, spherical or subspherical 10-15 um diam, solitary or rarely in 
very short chains, pale golden brown, thick-walled, slightly orna- 
mentated. 


Teleomorph: Botryobasidium vagum (Berk. & Curt.) Rogers. 


Material studied: Argentina, Buenos Aires, Ramallo; leg.: Romero, Ca- 
bral & Lopez, II-1982 (BAFC N° 30834); ibid, Gobernador Castro, XI-1982 
(BAFC N° 30835). 


Remarks: it is the first record for Argentina. 


7.b. Haplotrichum gracile (Linder) Hol. Jechova, Ceska Mycol. 30: 3-4, 
HU7O AP Geo bale 


Conidioma on bark, effuse, cotton or velvety, loose, margin irre- 
gular, pale yellow. Mycelium scanty of hyphae with simple septa, 6-9 um 
diam, slightly yellow, wall rather thick. Conidiophores erect, few bran- 
ched. Conidiogenous cells polyblastic, inflated, terminal and interca- 
lary, bearing cylindrical denticles up to 1.5 um high. Conidia holoblas- 


tic, ellipsoidal, smooth to slightly ornamentated, pale yellow, trunca- 
te at the base, 8-9 x 10-14 um. 


Teleomorph: unknown 


FIGURE 2 : A-D : Allescheriella cyanea. A: Mycelium, branched conidio-— 
phores, conidiogenous cells and young conidia. B: Attached mature coni- 
dium. C: Mature conidia. D: Clamped hyphae. E-G : "“Sporotrichum™ azu— 
reum. —: Conidiophores. F: Young and mature conidia. G: Clamped hyphae. 
H-I : Sporotrichum aurantiacum. H: Conidiophores. I: Conidia. 


437 


438 


Material studied: Argentina, Buenos Aires, Gobernador Castro,leg, 
Romero, Cabral & Lopez, XI-1981 (BAFC N° 30836); ibid, XI-1982 (BAFC 
N° 30837). 


Remarks: it is the first record for Argentina. 


ACKNOWLEDGEMENTS 


We thank Dr. J.A. Stalpers, C.B.S., Dr. R. Watling, Royal Botanic 
Gardens, Edinburgh, and u.E. Wright, University of Buenos Aires for 
their critical review of this manuscript. 


REFERENCES 


Arx, J.A. von. 1973. Further observations on Sporotrichum and some si- 
milar fungi. Persoonia 7(2): 127-136. 


Buchalo, A.S., V. Sasek & O.A. Zacordonec. 1985. Scanning electron 
microscopic study of anamorphs of some Basidiomycetes in cul- 
ture. Folia Microbiol. 30: 506-508. 


Drechsler, C. 1960. A clamp-bearing fungus using stalked adhesive 
young chlamydospores in capturing amoebae. Sydowia Ann. 
Mycol..ser.Il> 1432246-257. 


Dyko, B.J. & B.C. Sutton. 1979. Two new and unusual Deuteromycetes. 
Irnansee pre Mycols soc. 7243)6.4-4ive 


Ellis, M.B. 1971. Dematiaceous Hyphomycetes. Ist. ed. Commonw. Mycol. 
Inst., Kew, 50/ pp. 


Gochenaur, S.E. 1981. Cyrenella elegans gen. et sp. nov., a dikaryotic 
anamorph. Mycotaxon 13(1): 267-277. 


Hughes, S.J. 1951. Studies on Micro-Fungi. VII. Allescheriella crocea, 
Oidium simile and Pellicularia pruinata. Mycol. Pap. 41. 


Kendrick, B. & R. Watling. 1979. Mitospores in Basidiomycetes. In The 
Whole Fungus II. Ed. Kendrick, B. Nat. Mus. of Nat. Scien. 
Nac. Mus. of Canada & the Kananaski Found. Canada. 


Lopez, S.E. 1983. Sucesion Fungica en madera de Eucalyptus viminalis 
(Myrtaceae). I. Basidiomycetes sobre arbol vivo y troncos 
en estiba. Bol. Soc. Argent. Bot. 22(1-4): 21-39. 


1984. Estudio floristico y ecolodgico de Basidiomycetes 
xilofilos en plantaciones de Eucalyptus viminalis. Unpu- 
blished Thesis. Fac. de Cs. Exactas y Naturales, Univ. de 
Buenos Aires. 


& J.E. Wright. 1985. A new species of Hypochnicium (Corticia- 
ceae} from Argentina. Mycotaxon 23: 439-443. 


Maerz, A. & M.R. Paul. 1930. A dictionary of color. New York, Mc Graw- 
Bl ec ee DOr 


439 


Conidiophores with den- 


Paclime 3 = A-D = Haplotrichum curtisii-. A, B 
Young conidia. 


ticled conidiogenous cells and catenulate eonidia.< 1G 
D : Mature conidia. E-G : Haplotrichum gracile. E£, F : Conidiophores 
with denticled conidiogenous cells. G : Conidia. 


440 


Nag Raj, T.R. 1978. Genera coelomycetum. XVI. Fibulocoela form-gen. 
nov. a coelomycetes with basidiomycetous affinities. Can. 
J. Bot. :56(13): 1485-1491. 


Pantidou, M., R. Watling & Zacharoula, G. 1983. Mycelial characters, 
anamorph and teleomorph in genera and species of various fa- 
milies of Agaricales in culture. Mycotaxon 17: 409-432. 


Romero, A.I. 1983. Contribucio6n al estudio de los hongos xiléfilos de 
la Argentina. I. Deuteromycotina en Eucalyptus viminalis 
(Myrtaceae). Bol. Soc. Argent. Bot. 22(1-4): 57-79. 


Sivanesan, A. & R. Watling. 1980. A new mitosporic Basidiomycetes. 
Trans.) BeawMycoh. $00 14/4902) 424-4315 


Stalpers, J.A. 1984. A revision of the genus Sporotrichum. Stud. Mycol. 
2A HOS: 
& J. Viug. 1983. Confistulina, the anamorph of Fistulina 
hepatica. Can. J. Bot. 61: 1660- 


Udagawa, S., T. Awao & S.M. Adullah. 1986. Thermophymatospora, a new 
thermotolerant genus of  Basidiomycetous Hyphomycetes. 
Mycotaxon 27: 99-106. 


Weresub, L.K. & P.M. LeClair. 1971. On Papulospora and bulbilliferous 
Basidiomycetes Burgoa and Minimedusa. Can. J. Bot. 49: 2203- 
£213. 


Wright, J.E. 1983. Hirschioporus aculeifer, a polypore with anamorphic 
pileus processes. Rev. Biologia 12: 131-134. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 441-460 


January-March 1989 


SYNOPSIS OF A REVISED CLASSIFICATION 
FOR THE ENTOMOPHTHORALES 
(ZYGOMYCOTINA) 


RICHARD A. HUMBER 
USDA-ARS Plant Protection Research Unit 
US Plant, Soil, and Water Laboratory 
Tower Road 
Ithaca, New York 14853-0331, USA 


A revised classification of the Ento- 
mophthorales (Zygomycotina) based on 
expanded sets of familial and generic 
characters recognizes six families: Ento- 
mophthoraceae, Completoriaceae (fam. 
nov.), Ancylistaceae, Meristacraceae (fam. 
nov.), Neozygitaceae, and Basidiobol- 
aceae. Characters with familial signifi- 
cance include several aspects of nuclear 
cytology, the modes of formation and 
germination of resting spores, and the 
nature of vegetative growth and develop- 
ment. 

Primary generic characters all involve 
diverse aspects of the primary conidia: 
overall conidial and papillar morphology, 
nuclear number, uni- or bitunicate state of 
the wall, mode of discharge, and morph- 
ology of conidiophores and conidiogen- 
ous cells. Significant secondary generic 
characters include such correlated factors 
as the formation and morphology of any 
rhizoids and/or cystidia, the types of sec- 
ondary conidia formed, and characteristics 
of resting spores (e.g., color or decora- 
tion), vegetative cells (whether cells are 
walled or protoplastic, etc.), and general 
pathobiology (mode of action, host affini- 
ties, general habit, etc.). 


Major changes in this revision include 
the following: The four Batkoan subgen- 
era of Zoophthora are raised to genera as 
Zoophthora Batko, Erynia (Nowakowski 
ex Batko) Remaudiére & Hennebert, Furia 
(Batko) Humber, and Pandora Humber 
[= Erynia (Neopandora) Ben-Ze'ev & 
Kenneth]. Entomophaga Batko is re- 
stricted to species with pyriform conidia 
and forming no rhizoids, and Batkoa 
Humber is segregated for species with 
globose conidia formed on conidiogenous 
cells with a narrowed apical extension 
and that may form rhizoids with discoid 
holdfasts. An emended description of 
Conidiobolus (Ancylistaceae) provides the 
first morphological and developmental 
separation of this genus from Entomo- 
phaga and Batkoa (Entomophthoraceae). 
The Completoriaceae (fam. nov.) is de- 
scribed for Completoria complens, an 
obligate, intracellular parasite of fern 
prothallia. The Meristacraceae (fam. nov.) 
includes genera forming several conidia 
on erect, unbranched conidiogenous axes. 
The descriptions of the four previously 
accepted families are modified to include 
all characters accepted here as having 
familial value. 


442 


The description of the Neozygitaceae and the segregation of Thaxtero- 
sporium from Neozygites (Ben-Ze'ev et al., 1987) seemed to have modified 
the taxonomic criteria used to delimit entomophthoralean taxa (Humber, 
1981, 1982, 1984a; Ben-Ze'ev & Kenneth, 1982a). While this new family 
and new genus are appropriate additions to entomophthoralean 
systematics, their acceptance suggested the need to re-evaluate and to 
expand the range of taxonomic criteria accorded generic and familial 
value. As a result of the ensuing taxonomic analysis, several new familial 
and generic characters are recognized. The uniform application of these 
augmented taxonomic filters to all entomophthoralean fungi has indicated 
the further need to adjust the classification presented by Ben-Ze'ev & 
Kenneth (1982a,b) and Humber (1984a). The revised classification is 
presented in Table 1. 

The review of all taxa within the order has revealed some nomen- 
clatural problems. To the greatest extent possible, these problems are 
remedied here. The most notable among them is the discovery that 
Erynia Nowakowski (1881), which has been accepted by many authors as 
a valid generic name, is an invalidly published nomen nudum. The 
ramifications of this finding are discussed in some detail below, and 
should help guarantee the early acceptance of generic status for the four 
Batkoan subgenera of Zoophthora. 


REVISED FAMILIAL AND GENERIC CRITERIA 


A comprehensive treatment of the revised classification presented in 
Table 1 is being presented for publication elsewhere (Humber, 1989). This 
monograph includes extended discussions about (1) the emergence of the 
current systematics of the Entomophthorales, (2) the application of 
characters accorded familial and generic value (with special emphasis on 
critical nuclear characters which have never been discussed in depth in 
previous publications and on other aspects of development of resting 
spores and vegetative cells introduced’in this new classification), (©) 
some of the means now being adopted to distinguish species and species 
complexes, and (4) the evolutionary position of the Entomophthorales 
among the Zygomycetes. The systematic part of the full monograph 
provides detailed descriptions of all families, genera, and subgenera, com- 
plete bibliographic citations and synonymies of all taxa, information about 
the types of all species, and includes new combinations and new species 
not included in this synopsis. A synoptic key to the families, genera, and 
subgenera (Humber, 1989) is the first comprehensive key to use the post- 
Batkoan classification of this order. 

Families or other higher level taxonomic ranks should never be 
defined by single characters or small set of narrowly related characters; 
good systematics always depends upon the correlation of the largest 
possible number of fundamentally diverse characters. Humber (1981, 
1984a) provided bases for the use of nuclear cytological characters to de- 


443 


Table 1. Families and Genera of the Entomophthorales. This revised 
classification is fully elaborated in Humber (1989). It incorporates larger sets of 
familial and generic characters than those accepted by Humber (1981, 1982, 
1984), Tucker (1981), and Ben-Ze'ev & Kenneth (1982a-b). Correction of 
nomenclatural errors have required the adoption of author attributions for some 
taxa that differ from those previously accepted as correct. 


ENTOMOPHTHORACEAE Winter ANCYLISTACEAE Fisher 


Batkoa Humber, gen. nov. 

Entomophaga Batko 

Entomophthora Fresenius 

Erynia (Nowakowski ex Batko) 
Remaudiére & Hennebert 

Eryniopsis Humber 

Furia (Batko) Humber, stat. nov. 

Massospora Peck 


Ancylistes Pfitzer 
Conidiobolus Brefeld 
subgenus Conidiobolus 
subgenus Capillidium Ben- 
Ze'ev & Kenneth 
subgenus Delacroixia (Saccardo 
& Sydow) Tyrrell & MacLeod 
Macrobiotophthora Reukauf 


Pandora Humber, gen. nov. 

Strongwellsea Batko & Weiser NEOZYGITACEAE Ben-Ze'ev & 

Tarichium Cohn Kenneth in Ben-Ze'ev, Kenneth & 

Zoophthora Batko Uziel 

| Neozygites Witlaczil 

Thaxterosporium Be-Ze'ev & 
Kenneth in Ben-Ze'ev, Kenneth 
& Uziel 


COMPLETORIACEAE Humber, fam. nov. 
Completoria Lohde 


MERISTACRACEAE Humber, fam. nov. 


Ballocephala Drechsler BASIDIOBOLACEAE Claussen in 
Meristacrum Drechsler Engler & Gilg 
Zygnemomyces Miura Basidiobolus Eidam 


fine families within the Entomophthorales, but the application of these 
characters has been open to some misinterpretation. These significant 
cytological characters include the relative nuclear size and appearance and 
the quantity and distribution of any condensed chromatin during inter- 
phase, the morphology of mitotic chromosomes, the relative placement of 
the spindle at metaphase, and fate of the nuclear envelope during mitosis. 

This current revision adds several new characters Cinvolving resting 
spores and vegetative cells) to those cytological characters useful for 
circumscribing families. The resting spores characters include (1) the 
mode of formation (either axially aligned with the parental cell or budded 
through a narrow isthmus), (2) the presence or absence of a round of 
preconjugational gametangial mitosis, and (3) the mode of germination 
(with germ conidiophores forming directly from the spore or indirectly on 
a limited germ mycelium). The nature of vegetative cells seems to be 
useful at both familial and generic ranks. These cells may be walled or 
protoplastic; they may be multi-, pluri-. or uninucleate; their shape may be 
as hyphae forming a recognizable mycelium, as variably shaped hyphal 
bodies, as yeasts, or as botryose plasmodia; and, in relation to the host 


444 


cells, fungal growth and development may be intracellular or matrical (in 
the hemocoel or other intercellular spaces). 

Primary and secondary sets of characters are recognized for delimiting 
entomophthoralean genera. The primary characters elaborated previously 
(Humber 1981, 1984a; Ben-Ze'ev & Kenneth, 1982a) focus on primary 
conidia: the number of conidial nuclei, conidial morphology, uni- or 
bitunicate nature of the conidial wall, the mode of conidial discharge, and 
the morphology of the conidiophore and/or conidiogenous cell. One or 
more secondary characters should be invoked for the separation of genera 
only if they show strong correlations to variations in at least one of the 
primary generic characters. In the absence of significant variations in one 
or more primary characters, variations in secondary characters may serve 
as a reasonable basis for distinguishing subgenera. 

Secondary generic characters include (1) the presence (and mor- 
phology) of any rhizoids and/or cystidia, (2) the types of secondary coni- 
dia formed, (3) resting spore characters (e.g., unusual decorations, full 
separation of endospore and epispore, melanic or other characteristic 
colors, etc.), (4) vegetative characters (which are listed above), and (5) 
pathobiology (e.g., host range, mode of pathogenesis, habitat). Further 
secondary characters may yet be recognized, but any such additional 
characters must have legitimate significance and not be accepted merely 
for the convenience of justifying a particular taxonomic opinion. The 
secondary characters listed here are all relatively major features which 
appear to involve considerable portions of the fungal genome. 

The following two examples illustrate the care with which secondary 
generic characters must be used: In the first major post-Batkoan classifi- 
cation (Remaudiére & Hennebert, 1980; Remaudiére & Keller, 1980), the 
formation of secondary capilliconidia was the only unique character which 
distinguished Zoophthora from Erynia; the adoption of these two genera 
was rejected in favor of retaining the species in all of Batko's (1966) sub- 
genera of Zoophthora in one genus (Humber & Ben-Ze'ev, 1981; Ben- 
Ze'ev & Kenneth, 1982b; Li & Humber, 1984). There are subtle but 
reasonably consistent differences in conidial morphology among these 
subgenera, and some minor variation in the degree of branching of the 
conidiophores; these subtle differences in primary characters cannot, in 
themselves, justify the elevation of these subgenera to generic rank. 
These variations in primary characters are, however, strongly correlated 
with marked differences among all subgenera in the morphologies of 
rhizoids and cystidia, in the overall host range and habitat, and, appar- 
ently, in the walled or protoplastic nature of the vegetative cells. None of 
these differences in secondary characters is individually sufficient to 
warrant a change from subgeneric to generic rank, but when the entire 
range of primary and secondary characters are taken together, there is 
ample basis to raise the Batkoan subgenera of Zoophthora to generic 
status. The three subgenera of Conidiobolus (Ben-Ze'ev & Kenneth, 
1982a) provide a second valuable example for the application of second- 
ary generic characters. These subgenera are separated by only one 


445 


secondary character, their types of secondary conidia formed. Because 
there are no significant differences among the subgenera in any primary 
generic characters, these secondary differences cannot be recognized 
reasonably as taxonomically different at any rank higher than the 
subgenus. 


NEW TAXA, COMBINATIONS, AND EMENDATIONS 


The formal descriptions and combinations required to validate the 
classification in Table 1, and to allow the immediate adoption of this 
classification are provided in this section. For most users, the major 
changes proposed herein include (1) the subgenera of Erynia sensu 
Humber & Ben-Ze'ev (1981; Ben-Ze'ev & Kenneth, 1982b) are elevated to 
generic rank, (2) notice that the use of Erynia in the broad sense (Humber 
& Ben-Ze'ev, 1981) was nomenclaturally incorrect, and (3) the segregation 
of Batkoa from Entomophaga for species with globose conidia borne on 
narrowed extensions of the conidiogenous cell and which may also 
produce rhizoids with discoid holdfasts. 

Except where specifically listed below, the species accepted in any 
other genus in this present classification generally agree with those listed 
by Tucker (1981), Ben-Ze'ev & Kenneth (1982a-b), and Ben-Ze'ev et al. 
(1987). Further new species not treated in this synopsis will appear in the 
full monographic treatment of this revised classification (Humber, 1989). 


ENTOMOPHTHORACEAE Winter 


Early vegetative stage mycelial, or hyphal bodies with or without distinct 
cell walls, or fusoid to catenate or irregularly shaped amoeboid proto- 
plasts. Nuclei typically 5-12 um (but as little as 2.5-3.0 um) diameter 
during interphase, containing much condensed chromatin that usually 
stains readily with aceto-orcein or bismarck brown; no prominent central 
nucleolus; usually arranged in single file in coenocytic to infrequently 
septate hyphae. Nuclei remain distinctly visible throughout mitosis; 
mitosis intranuclear (with nuclear envelope remaining intact and closed 
throughout mitosis); chromosomes more or less distinct and vermiform 
during late mitotic stages. Conidiophores simple or dichotomously to 
digitately branched; conidiogenous cells apical on each conidiophore 
branch. Primary conidia unitunicate or bitunicate; forcibly discharged by 
papillar eversion or on a propulsive jet of conidiophore contents, or 
passively dispersed from numerous conidiogenous cavities interspersed in 
an easily dissociated fungal mass. Resting spores form as lateral or 
terminal buds connected to parental cell by narrow isthmus. Obligate 
pathogens of insects or other arthropods. 


Type genus: Entomophthora Fresenius 


446 


Batkoa Humber, gen, nov. 


[= Entomophaga subgenus Lichia Weiser & Batko in Batko & Weiser, 1965, J. Invertebr. 
Pathol. 7: 461. (nomen nudum, without diagnosis)] 


Cellulae vegetativae hyphae vel corpora hyphalia filamentosa. Nuclei grandiusculi, 
granulis heterochromatis et prompte per "aceto-orcein" vel "Bismarck brown" coloratis 
repleto; nucleolo parvo eccentrico. Apparatus mitoticus parvus fusiformis eccentricus per 
metaphasem; chromosomata vermiformes spatium nuclei subtotum adsument per 
metaphasem. Conidiophora simplicia; apice cellulis conidiogenis plerumque attenuato 
exserto. Cystidia hymenica desunt. Conidia primaria globosa subglobosa, unitunicata, 
multinucleata; vehementer eversione papillae basalis expulsa. Rhizoidea aliquando prae- 
sentia, cellula conidiogena multo crassius, singularia (nonfasciculata), haptero apicali 
disciformi vel irregulariter ramoso ut in speciebus Pandorae. Sporae perdurantes ex 
cellula parentali pullulantes et mycelium germinale formantes per germinationem. 
Species Entomophthoracearum pathogeniae obligatae hospites insectorum. 


Vegetative cells hyphal or in thread-like hyphal bodies, apparently with a 
cell wall in natural hosts. Nuclei relatively large, with prominent granules 
of condensed chromatin during interphase, staining readily with aceto- 
orcein, Bismarck brown, or other nuclear stains; nucleolus small, irregular, 
randomly located. Mitotic spindle small and eccentric during metaphase, 
chromosomal arms filling nuclear volume; chromosomal arms radiating 
from chromatin-free zone around spindle poles, arranged in dome-like 
fashion during anaphase. Conidiophores simple, conidiogenous cells with 
tendency for neck-like apical narrowing immediately below conidium. 
Cystidia absent from hymenium. Primary conidia globose to subglobose, 
unitunicate, multinucleate; forcibly discharged by papillar eversion. 
Rhizoids occasionally present; if so, distinctly thicker than vegetative 
hyphae or conidiophores, occurring singly, with discoid terminal holdfast 
(resembling that of Pandora species). Resting spores arising by budding 
from parental cell, and assumed to germinate indirectly by formation of 
germ mycelium. Obligately entomopathogenic, with most species 
affecting Homoptera, Hemiptera, or other insects in moist environments. 
Type species: Batkoa apiculata (Thaxter) Humber, comb. nov. Basionym: Empusa 
apiculata Thaxter, 1888, Mem. Boston Soc. Nat. Hist. 4: 163. Lectotype: FH 4331. [= 
Entomophaga domestica Keller, 1987, Sydowia 40: 141.] 
Other included species: 


Batkoa daysderci (Viégas) Humber, comb. nov. Basionym: Empusa dysderci 
Viégas, 1939, Jorn. Agron. 2: 251. 


Batkoa gigantea (Keller) Humber, comb. nov. Basionym: Entomophthora 
gigantea Keller, 1978, Sydowia, Ann. Mycol. (Ser. ID 31: 89. 


Batkoa major (Thaxter) Humber, comb. nov. Basionym: Empusa apiculata var. 
major Thaxter, 1888, Mem. Boston Soc. Nat. Hist. 4: 163. Lectotype: FH 4306. 
[= Entomophaga limoniae Keller, 1987, Sydowia 40: 146.] 


Batkoa papillata (Thaxter) Humber, comb. nov. Basionym: Empusa papillata 
Thaxter, 1888, Mem. Boston Soc. Nat. Hist. 4: 166. Neotype: FH 3316. 


Thaxter (1888) designated no types for the new species described in 
his monograph. The correct disposition of Empusa apiculata, E. major, 


447 


and E. papillata among the Entomophthoraceae (Ben-Ze'ev & Kenneth 
1982a) or the Ancylistaceae (Remaudiére & Keller, 1980; Keller, 1987) 
depends wholly upon the familial affinities of the nomenclatural type 
specimens that must be designated from among Thaxter's authentic 
collections. Virtually all specimens of E. apiculata and E. major used to 
prepare Thaxter's (1888) monograph proved to have entomophthoraceous 
nuclei. Because a thorough search of the entomophthoralean specimens 
at the Farlow Herbarium (FH) disclosed no authentic specimens of B. 
papillata, | neotypified E. papillata with the only other FH collection of 
this species, a specimen whose nuclei were clearly entomophthoraceous. 
A lectotype for B. papillata must be chosen to supercede this neotype if 
Thaxter's authentic specimens are ever located. 

The typifications of B. apiculata, B. major, and B. papillata fix the 
placements of these species among the Entomophthoraceae. The ancy- 
listaceous species identified by Keller (1987) as Conidiobolus apiculatus 
(Thax.) Remaud. & Keller and as Conidiobolus major (Thax.) Remaud. & 
Keller must be described as new Conidiobolus species while their ento- 
mophthoraceous "pairs", Entomophaga domestica Keller and Entomo- 
phaga limoniae must be synonymized with the Thaxterian species here 
placed in Batkoa. 


Entomophaga Batko emend. Humber, emend. nov. 


Vegetative cells as hyphoid or more or less beaded hyphal bodies to 
individual fusoid cells; usually wall-less protoplasts during early stages of 
development. Nuclei relatively large, with prominent granules of con- 
densed chromatin during interphase, staining readily with aceto-orcein or 
other nuclear stains; nucleolus small, irregular, randomly located. Mitotic 
spindle small and eccentric during metaphase, chromosomal arms filling 
nuclear volume; chromosomal arms radiating from chromatin-free zone 
around spindle poles, arranged in dome-like fashion during anaphase. 
Conidiophores simple, conidia produced directly on apex without arising 
on a narrower neck-like extension. Cystidia absent from hymenium. 
Primary conidia pyriform, unitunicate, multinucleate, with more or less 
prominent papilla; forcibly discharged by papillar eversion; on water- 
logged specimens, (coronate) primary conidia with 2+ distal projections 
may be formed and passively dispersed. Rhizoids absent. Resting spores 
arise by budding from parental cell, and germinate indirectly by formation 
of germ mycelium. Obligately entomopathogenic. 
Type species: Entomophaga grylli (Fresenius) Batko, 1964, Bull. Acad. Polon. Sci., 

Ser oci--Biol., 12: 325. 
Other included species: 

E. aulicae (Reichardt in Bail) Humber, 1984, Mycotaxon 21: 270 

E. batkoi (Balazy) Keller, 1987, Sydowia 40: 138. 

Entomophaga calopteni (Bessey) Humber, comb. nov. Basionym: Entomophthora 

calopteni Bessey, 1883, Amer. Nat. 17: 1280. 
E. conglomerata (Sorokin) Keller, 1987, Sydowia 40: 138. 


448 


E. kansana (Hutchison) Batko, 1964, Bull. Acad. Polon. Sci., Sér. Sci. Biol., 12: 404. 
E. saccharina (Giard) Batko, 1964, Bull. Acad. Polon. Sci., Sér. Sci. Biol., 12: 404. 
E. tabanivora (Anderson & Magnarelli) Humber, 1984, Mycotaxon 21: 270. 

E. tenthredinis (Fresenius) Batko [Bull. Acad. Polon. Sci., Sér. Sci. Biol., 12: 404. 


Entomophaga tipulae (Fresenius) Humber, comb. nov. Basionym: Entomoph- 
thora tipulae Fresenius, 1858, Abhandl. Senckenb. Naturf. Ges. 2: 206. 


This emended generic description recognizes the ability of some 
Entomophaga species to form multiradiate primary or secondary conidia 
on waterlogged specimens. Descals & Webster (1984) noted the forma- 
tion of such conidia by E. tipulae (as the invalidly published synonym, 
Entomophthora thaxteri Brumpt); such conidial behavior had been 
known previously only from species of Erynia (sensu Stricto). 

Entomophaga calopteni was originally described as a pathogen of the 
differential grasshopper, Melanoplus (= Caloptenus) differentialis. This 
name is adopted for the North American member of the E. grylli species 
complex that attacks spur-throated grasshoppers (Melanoplus spp.; 
Orthoptera: Acrididae), and produces resting spores but no conidia 
under normal conditions either in the host or in culture. E. calopteni is 
more widely known in the literature as the North American pathotype II 
of E. grylli (Soper et al., 1983; Humber 1984b). The North American 
pathotype I of E. grylli (Soper et al., 1983; Humber 1984b) freely pro- 
duces conidia and attacks hosts from genera other than Melanoplus; this 
fungus and other members of this species complex from non-European 
locations will be described later as new species (Humber, 1989). 


Entom ra Freseni 


The complex of species which includes E. muscae (Cohn) Fresenius, 
the type species of Entomophthora, now also includes E. israelensis Ben- 
Ze'ev & Zelig (= E. brevinucleata Keller & Wilding), E. scatophagae 
Giard (Steinkraus and Kramer, 1988), E. schizophorae Keller & Wilding in 
Keller (1987, p. 160), and E. trinucleata Keller (1987, p. 161), Other 
Entomophthora species include E£. culicis (Braun) Fresenius; E. erupta 
(Dustan) Hall; FE. helvetica Keller & Ben-Ze'ev in Ben-Ze'ev, Keller & 
MacEwen; E. planchoniana Cornu; E. thripidum Samson, Ramakers & 
Oswald; and E. weberi Lakon ex Samson in Samson, Ramakers & Oswald. 


Erynia (Nowakowski ex Batko) Remaudiére & Hennebert 


emend. Humber, emend. nov. 


Zoophthora subgenus Erynia Nowakowski ex Batko, 1966, Acta Mycol. 2: 18. 
Erynia (Nowakowski ex Batko) Remaudiére & Hennebert, 1980, Mycotaxon 11: 
301. 

Vegetative cells as hyphal bodies, apparently with cell walls in natural 
hosts. Nuclei medium to large, with prominent granules of condensed 


sll 


449 


chromatin during interphase, staining readily with aceto-orcein or other 
nuclear stains; nucleolus small, irregular, randomly located. Mitotic 
spindle small and eccentric during metaphase, chromosomal arms filling 
nuclear volume; chromosomal arms radiating from chromatin-free zone 
around spindle poles, arranged in dome-like fashion during anaphase. 
Conidiophores simple, arising from more or less spherical hyphal bodies; 
conidiogenous cells slightly clavate. Cystidia present in hymenia of all 
known species, at least twice as thick as conidiophores, occasionally 
branched, standing out well above layer of conidiophores. Primary 
conidia pyriform to elongate or fusoid, often strongly curved to lunate, 
with rounded to broadly conical basal papilla; bitunicate; uninucleate; 
forcibly discharged by papillar eversion, or, in waterlogged specimens, by 
passively dispersed (coronate) primary conidia with 2+ distal projections. 
Secondary conidia globose or resembling primary and forcibly discharged 
by papillar eversion, or, in waterlogged specimens, with 2+ proximal 
projections ("stellate secondary conidia") and passively dispersed. 
Rhizoids at least twice as thick as conidiophores or vegetative cells; 
without distinct terminal holdfast. Resting spores arise by budding from 
parental cell; assumed to germinate indirectly by formation of germ 
mycelium. Obligately entomopathogenic. 
Type species: Erynia ovispora (Nowakowski) Remaudiére & Hennebert, 1980, 

Mycotaxon 11: 301. 
Other included species: 

E. aquatica (Anderson & Anagnostakis) Humber. 1981. Mycotaxon 13: 213] 

E. conica (Nowakowski) Remaudiére & Hennebert, 1980, Mycotaxon 11: 302]. 

E. curvispora (Nowakowski) Remaudiére & Hennebert, 1980, Mycotaxon 11: 302. 

E. delpiniana (Cavara) Humber, 1981, Mycotaxon 13: 212] 

E. erinacea (Ben-Ze'ev & Kenneth) Remaudiére & Hennebert, 1980, Mycotaxon 11: 

302. 

E. gracilis (Thaxter) Remaudiére & Hennebert, 1980, Mycotaxon 11: 302. 

E. henrici (Molliard) Humber & Ben-Ze'ev, 1981, Mycotaxon 13: 509. 

E. plecopteri Descals & Webster, 1984, Trans. Brit. Mycol. Soc. 83: 669. 

E. rhizospora (Thaxter) Remaudiére & Hennebert, 1980, Mycotaxon 11: 302. 

E. sepulchralis (Thaxter) Remaudiére & Hennebert, 1980, Mycotaxon 11: 302. 

E. variabilis (Thaxter) Remaudiére & Hennebert, 1980, Mycotaxon 11: 302. 


The taxonomic authorities and synonymy accepted here for the name 
Erynia diverges markedly from that recognized in recent years. This 
generic name first appeared in a published report of a meeting of Polish 
physicians and biologists (Nowakowski, 1881). In this report, no 
characters were given to distinguish Erynia from Empusa, Entomophthora, 
or Lamia; Erynia Nowakowski (1881) is, therefore, an invalidly published 
nomen nudum. Nowakowski later rejected this genus since he did not 
mention Erynia and treated its intended species (E. ovispora, E. conica, 
and E. curvispora) as Entomophthora species in his monograph 
(Nowakowski, 1883); if Erynia had been validly published in 1881, this 
later rejection would have had no effect on the nomenclatural priority of 


450 


the generic name. Batko (1966) did characterize, typify, and validate 
Erynia Nowakowski as a subgenus of Zoophthora despite his mistaken 
belief about the reasons for the invalidity of the generic name. 

Remaudiére & Hennebert (1980) were the first authors to use the 
validated Erynia at the generic rank even though they had accepted 
Nowakowski's (1881) mention of Erynia to be nomenclaturally valid. If 
the true status of Erynia Nowakowski (1881) had been known, Humber & 
Ben-Ze'ev (1981) and Ben-Ze'ev & Kenneth (1982a,b) would have 
recognized Zoophthora Batko (1964) as the oldest valid and nomenclatur- 
ally correct name for the genus in their broad sense rather than accepting 
Erynia. It is important to note that the taxonomy proposed in this 
revision is unaffected by these past nomenclatural errors since the type 
species of Zoophthora (Z. radicans) and Erynia (E. ovispora) are once 
again placed in separate genera as they had been by Remaudiére & 
Hennebert (1980) and Remaudiére & Keller (1980). 


Furia (Batko) Humber, stat. gen. nov. et emend. nov. 


Zoophthora subgenus Furia Batko, 1966, Acta Zool. 2: 20. 
Erynia subgenus Furia (Batko) Li & Humber, 1984, Canad. J. Bot. 62: 662. 


Vegetative growth in vivo as hyphoid protoplasts or walled hyphal bodies. 
Nuclei large, generally >5 um diameter, with prominent condensed 
chromatin during interphase, staining readily with aceto-orcein and other 
nuclear stains. Mitotic metaphase plate small, lateral, and occupied by 
kinetochores while chromosomal arms occupy remaining nuclear volume. 
Conidiophores with low order of apical branching (not prominently 
digitate), intertwined in dense hymenia. Cystidia as thick as conidio- 
phores, unbranched, tapering apically. Primary conidia ovoid to 
cylindrical, obpyriform to obclavate or fusoid, with no strong tendency to 
show bilateral symmetry (basal papilla may be displaced laterally from 
spore axis); uninucleate; bitunicate (outer wall layer may separate in 
liquid); forcibly discharged by papillar eversion. Secondary conidia more 
or less resembling primary conidia, produced singly on primary conidia; 
forcibly discharged by papillar eversion; capilliconidia absent. Rhizoids 
monohyphal or fasciculate or (rarely?) pseudorhizomorphic; no thicker 
than conidiophores; with no strongly differentiated terminal holdfast, or 
with small, sucker-like expansions near apices, or with few, irregular, thin, 
short terminal branches spreading on substrate. 
Type species: Furia virescens (Thaxter) Humber, comb. nov. Basionym: Empusa 

virescens Thaxter, 1888, Mem. Boston Soc. Nat. Hist. 4: 178. 
Other included species: 

Furia americana (Thaxter) Humber, comb. nov. Basionym: Empusa (Ento- 

mophthora) americana Thaxter, 1888, Mem. Boston Soc. Nat. Hist. 4: 179. 
Furia creatonoti (Yen in Humber) Humber, comb. nov. Basionym: Erynia 
creatonoti Yen in Humber, 1981, Mycotaxon 13: 474. 


451 


Furia crustosa (MacLeod & Tyrrell) Humber, comb. nov. Basionym: Ento- 
mophthora crustosa MacLeod & Tyrrell, 1979, Canad. Entomol. 111: 1138. 


Furia ellisiana (Ben-Ze'ev) Humber, comb. nov. Basionym: Erynia ellisiana Ben- 
Ze'ev, 1986, Mycotaxon 27: 266. 


Furia ithacensis (Kramer) Humber, comb. nov. Basionym: Erynia ithacensis 
Kramer, 1981, Mycopathologia 75: 160. 


Furia montana (Thaxter) Humber, comb. nov. Basionym: Empusa montana 
Thaxter, 1888, Mem. Boston Soc. Nat. Hist. 4: 180. 


Furia neopyralidarum (Ben-Ze'ev) Humber, comb. nov. Basionym: Erynia 
neopyralidarum Ben-Ze'ev, 1982, Mycotaxon 16: 276. 

Furia pieris (Li & Humber) Humber, comb. nov. Basionym: Erynia pieris Li & 
Humber, 1984, Canad. J. Bot. 62: 656. 


Furia sciarae (Olive) Humber, comb. nov. Basionym: Empusa sciarae Olive, 
1906, Bot. Gaz. (Crawfordsville) 41: 196. 


Furia vomitoriae (Rozsypal) Humber, comb. nov. Basionym: Zoophthora 
vomitoriae Rozsypal, 1966, Acta Mycol. 2: 24. 


Furia zabri (Rozsypal ex Ben-Ze'ev & Kenneth) Humber, comb. nov. Basionym: 
Erynia zabri Rozsypal ex Ben-Ze'ev & Kenneth (as E. zabrii), 1982, Mycotaxon 14: 
465. 


The placement of so many species in Furia may rightfully be 
questioned since Batko (1966) and Li & Humber (1984) regarded this to 
be a limited group and since Ben-Ze'ev & Kenneth (1982b) declined to 
place several of the species included here in any Erynia subgenus. The 
morphology of primary conidia in Furia species is virtually indistin- 
guishable from that of Pandora species, but is distinct from the elongate 
or to prominently cylindrical conidia of many species of Erynia s.str. and 
Zoophthora s.stt.. Furia species are readily distinguished from Pandora 
species, however, by their rhizoids and/or cystidia. In Furia, the rhizoids 
are no thicker than vegetative hyphae; these structures may be solitary, 
fasciculate, or even bundled into pseudorhizomorphs (ike those of some 
Entomophthora ot Zoophthora species), but have either no differentiated 
terminal holdfasts, rudimentary suckers or sparse, open systems of thin 
branches (which may even be arranged in a discoid fashion). The rhi- 
zoids of Furia species are neither so thick as those of Pandora species, 
nor do they ever possess the massive, solid discoid holdfast of the latter 
genus (in which rhizoids are solitary rather than fasciculate). In the 
erynioid genera, the thickness of the cystidia matches that of the rhizoids; 
in Furia, cystidia are no thicker than vegetative hyphae or conidiogenous 
cells; as in other genera, they taper apically. 


Pandora Humber, gen, nov, 


= Zoophthora subgenus Pandora Batko [pro parte], 1966, Acta Mycol. 2: 18. [as described 
but excluding the designated type, Entomophthora aphidis Hoffmann in Fres., which is a 
species of Zoophthora sensu stricto] 

= Erynia subgenus Neopandora Ben-Ze'ev & Kenneth, 1982, Mycotaxon 14: 460. 


Cellulae vegetativae plerumque filamentosae in vivo protoplastes sive parietatae. Nuclei 
magni, > 5 um diam., granulis heterochromatis et prompte per "aceto-orcein" vel "Bis- 


452 


marck brown" coloratis repleto; nucleolo parvo eccenrico. Apparatus mitoticus parvus 
fusiformis eccentricus per metaphasem; chromosomata vermiformes spatium nuclei 
subtotum adsument per metaphasem. Conidiophora digitata apicale, in hymeniis densis 
intertextis. Cystidia hymenica attenuata plerumque praesentia, cellula conidiogena 2-3plo 
crassius. Conidia primaria ovoidea subcylindrica, papilla basali axiali vel laterali, 
uninucleata, bitunicata; vehementer eversione papilla basali expulsa. Conidia secundaria 
solitaria, vehementer eversione papilla expulsa; capilliconidia desunt. Rhizoidea solitaria, 
cellula conidiogena 2-3plo crassius, vacuolata, haptero apicali disciformi vel irregulariter 
ramoso terminantia. Sporae perdurantes ex cellula parentali pullulantes et mycelium 
germinale formantes per germinationem. Species Entomophthoracearum pathogeniae 
obligatae hospites insectis arachnidis. ! 


Vegetative growth in vivo as hyphoid protoplasts or walled hyphal bodies. 
Nuclei large, generally =5 tm diameter, with prominent condensed 
chromatin during interphase, staining readily with aceto-orcein and other 
nuclear stains. Mitotic metaphase plate small, lateral, and occupied by 
kinetochores while chromosomal arms occupy remaining nuclear volume. 
Conidiophores digitately branched at apices (rarely bifurcate or simple), 
intertwined in dense hymenia. Cystidia usually present, 2-3 times thicker 
than conidiophores, tapering apically. Primary conidia ovoid to 
cylindrical, obpyriform to obclavate or fusoid, with no strong tendency to 
show bilateral symmetry (basal papilla may be displaced laterally from 
spore axis); uninucleate; bitunicate (outer wall layer may separate in 
liquid); forcibly discharged by papillar eversion. Secondary conidia more 
or less resembling primary conidia, produced singly on primary conidia; 
forcibly discharged by papillar eversion; capilliconidia absent. Rhizoids 
monohyphal, 2-3 times thicker than conidiophores, highly vacuolate, 
terminating in discoid or irregularly branched and spreading holdfast. 
Type species: Pandora neoaphidis (Rem. & Henn.) Humber, comb. nov. Basionym: 
Erynia neoaphidis Remaudiére & Hennebert, 1980, Mycotaxon 11: 307. 
Other included species: 
Pandora blunckii (Lakon ex Zimmermann) Humber, comb. nov. Basionym: 
Entomophthora blunckii Lakon ex Zimmermann, 1978, Entomophaga 23: 182. 
Pandora brahbminae (Bose & Mehta) Humber, comb. nov. Basionym: Ento- 
mophthora brahminae Bose & Mehta, 1953, Trans. Brit. Mycol. Soc. 36: 55. 
Pandora bullata (Thaxter & MacLeod in Humber) Humber, comb. nov. Basionym: 
Erynia bullata Thaxter & MacLeod in Humber, 1981, Mycotaxon 13: 472. 
Pandora calliphborae (Giard) Humber, comb. nov. Basionym: Entomophthora 
calliphorae Giard, 1879, Bull. Sci. Dep. Nord, Sér. 2, 2: 356. 
Pandora dacnusae (Balazy) Humber, comb. nov. Basionym: Erynia dacnusae 
Balazy, 1981 (1982), Bull. Acad. Polon. Sci., Sér. Sci. Biol. 29: 229. 
Pandora delphacis (Hori) Humber, comb. nov. Basionym: Entomophthora 
delphacis Hori, 1906, Entomol. Mag. (Tokyo) 3: 81. 
Pandora dipterigena (Thaxter) Humber, comb. nov. Basionym: Empusa dip- 
terigena Thaxter, 1888, Mem. Boston Soc. Nat. Hist. 4: 177. 
Pandora echinospora (Thaxter) Humber, comb. nov. Basionym: Empusa 
echinospora Thaxter, 1888, Mem. Boston Soc. Nat. Hist. 4: 180. 
Pandora formicae (Humber & Balazy in Humber) Humber, comb. nov. Basionym: 
Erynia formicae Humber & Balazy in Humber, 1981, Mycotaxon 13: 475. 


453 


Pandora gammae (Weiser) Humber, comb. nov. Basionym: Tarichium gammae 
Weiser, 1965, Ceska Mykol. 19: 203. 


Pandora gloeospora (Vuillemin) Humber, comb. nov. Basionym: Entomophthora 
gloeospora Vuillemin, 1886, Bull. Soc. Sci. Nat. Nancy, Sér. 2, 19: 34. 


Pandora kondoiensis (Milner in Milner, Mahon & Brown) Humber, comb. nov. 
Basionym: Erynia kondoiensis Milner in Milner, Mahon & Brown, 1983, Austral. J. 
Bot. 31: 183. 


Pandora nouryi (Remaudiére & Hennebert) Humber, comb. nov. Basionym: 
Erynia nouryi Remaudiére & Hennebert, 1980, Mycotaxon 11: 313. 


Pandora phalangicida (Lagerheim) Humber, comb. nov. Basionym: Entomopb- 
thora phalangicida Lagerheim, 1898, Bihang Svensk. Vet. Akad. Handl. 24: 12. 


Pandora suturalis (Ben-Ze'ev) Humber, comb. nov. Basionym: Erynia (Neo- 
pandora) suturalis Ben-Ze'ev, 1987, Mycotaxon 28: 405. 


Zoophthora Batko 


Species included in this genus, which is typified by Z. radicans 
(Brefeld) Batko, follow the listing accepted in Erynia (Zoophthora) by 
Ben-Ze'ev & Kenneth (1982b) together with two further species either 
added to this group or described as new. New combinations are provided 
for these two species: 

Zoophthora anglica (Petch) Humber, comb. nov. Basionym: Entomophthora 
anglica Petch, 1944, Trans. Brit. Mycol. Soc. 27: 89. 


Zoophthora anbuiensis (Li) Humber, comb. nov. Basionym: Erynia anhuiensis Li, 
1986, Acta Mycol. Sin. 5: 2. 


Z. aphidis (Hoffmann in Fresenius) Batko, 1964, Bull. Acad. Polon. Sci., Sér. Sci. Biol. 
12: 405. 


Z. canadensis (MacLeod, Tyrrell & Soper) Remaudiére & Hennebert, 1980, Mycotaxon 
ase 1. 


Z. crassitunicata Keller, 1980, Sydowia Ann. Mycol., Ser. 2, 33: 170. 


Zoophthora geometralis (Thaxter) Batko, 1964, Bull. Acad. Polon. Sci., Sér. Sci. Biol. 
12: 404. 


Z. lanceolata Keller, 1980, Sydowia Ann. Mycol., Ser. 2, 33: 167. 

Z. occidentalis (Thaxter) Batko, 1964, Bull. Acad. Polon. Sci., Sér. Sci. Biol. 12: 404. 
Z. orientalis Ben-Ze'ev & Kenneth, 1981, Phytoparasitica 9: 35. 

Z. petchii Ben-Ze'ev & Kenneth (as Z. petchi), 1981, Entomophaga 26: 140. 

Z. phalloides Batko, 1966, Acta Mycol. 2: 7. 

Z. phytonomi (Arthur) Batko, 1964, Bull. Acad. Polon. Sci., Sér. Sci. Biol. 12: 404. 
Z. radicans (Brefeld) Batko, 1964, Bull. Acad. Polon. Sci., Sér. Sci. Biol. 12: 323. 


COMPLETORIACEAE Humber, fam. nov. 


Cellulae vegetativae parvae botryosae irregulares probabiliter protoplastes in cellulis 
prothallorum filicium. Nuclei comparate magni chromatino granulato condensato per 
interphasem. Mitoses non vidi. Conidiophora simplicia brevia, ex cellulis vegetativis 
directe sine cellulis conidiogenis specialibus formantibus. Conidia primaria unitunicata, 
subglobosa, papilla pusilla rotundata; vehementer eversione papillae basalis expulsa. 
Sporae perdurantes globosae et parvae; in axe cellula parentali formare et per 


454 


germinatione conidii germinalis formationibus directis adsumptae sunt. Genus 
monotypicus Entomophthoralium pathogeniae obligatae prothallis filictum. 


Vegetative growth as small, botryose, irregular hyphal bodies inside cells 
of fern gametophyte; appearance and development suggest that no cell 
walls are present. Nuclei relatively large, filled with condensed chromatin 
at interphase (giving granular appearance to nucleoplasm); no mitoses 
observed. Conidiophores simple, short, arising directly from vegetative 
cells; no separate conidiogenous cell formed. Conidia unitunicate, sub- 
globose, with small, rounded papilla; forcibly discharged by papillar ever- 
sion. Resting spores globose, but neither mode of formation nor of 
germination has been confirmed; resting spores apparently form in axis of 
parental cell; germination is probably by direct production of germ coni- 
diophore and forcibly discharged conidium. Obligate intracellular parasite 
of fern gametophytes. 


Type genus: Completoria Lohde 


The inclusion of a monotypic family in this classification may seem 
odd, especially when so many key characters have been inferred rather 
than confirmed by observation. Completoria complens appears to be a 
rather rare fungus and has been observed only from greenhouse cultures 
where it grows as an intracellular parasite of fern gametophytes. I am 
unaware of any report of this fungus affecting field populations of 
gametophytes. The missing characters that have been "filled in" for this 
familial description were done so by choosing the most probable character 
states of characters for such a highly specific parasite. The accuracy of 
these guesses will provide a substantial test for the ability to use this 
classification to predict character states. Completoria was previously 
included in the Entomophthoraceae (Tucker, 1981; Ben-Ze'ev & Kenneth, 
1982a; Humber, 1984a); removing C. complens leaves only obligately 
entomopathogenic species the Entomophthoraceae. 


ANCYLISTACEAE Fisher 


Vegetative growth as irregularly septate or coenocytic mycelium, tending 
to become disjointed with evacuation of cytoplasm from some cells and 
fragmentation of wall. Nuclei small, usually 3-5 um long during interphase, 
with prominent central nucleolus; no significant quantity of condensed 
chromatic granular material in nucleoplasm during interphase (nuclei not 
staining strongly with aceto-orcein or bismarck brown). Nuclei usually 
difficult to observe during interphase or mitosis; mitosis intranuclear Cwith 
nuclear envelope remaining intact throughout mitosis), with nucleolus 
persistent and laterally displaced during mitosis; chromosomes tiny and 
indistinct. Conidiophores simple or infrequently branched and bearing a 
single terminal conidium per branch. Primary conidia unitunicate, forcibly 
discharged by papillar eversion. Resting spores typically zygospores 
formed from conjugations of adjacent cells in hypha or by scalariform 


455 


conjugations between two hyphae; resting spores formed in axis of 
parental hypha or conjugation tube. Saprobes in soil or pathogens of 
insects or other soil invertebrates or desmid algae, or as facultative 
mycotic agents of vertebrates (including humans). 


Type genus: Ancylistes Pfitzer 


Conidiobolus Brefeld emend. Humber, emend. nov. 


Mycelium inconspicuous when young; sparse to luxuriant when mature, 
often infrequently septate; aerial mycelium nonexistent to abundant; 
vegetative growth always as walled cells; often forming variably shaped 
hyphal bodies by pinching off or dissociation of intercalary cells devoid of 
cytoplasm; growing readily on nutritionally simple culture media but 
unable to utilize nitrate or nitrite. Nuclei 2.5-5 um in diameter, usually 
inconspicuous by light microscopy; nucleoplasm with little or no 
condensed chromatic stainable by aceto-orcein or other chromatin stains; 
nucleolus central, ovoid; mitotic stages cryptic with light microscopy; 
mitotic chromosomes tiny, indistinct. Conidiophores simple or (rarely) 
branched dichotomously, positively phototropic; conidiogenous cells 
usually undistinguished in diameter or appearance from vegetative hyphae 
(but occasionally clavate and/or distinctly thicker than vegetative hyphae), 
with a basal septum, producing a single apical conidium. Primary conidia 
multinucleate, unitunicate (outer wall layer not separating from conidial 
surface), pyriform to obovoid or globose; papilla rounded to apiculate; 
forcibly discharged toward light source by papillar eversion against the 
conidiogenous cell. Secondary conidia usually with shape of primary 
conidia but smaller; formed singly on short secondary conidiophore and 
forcibly discharged toward light source by papillar eversion or ovoid to 
cylindrical capilliconidia dispersed passively from apices of capillary 
secondary conidiophores, produced singly from a primary conidium or 
producing 3+ small, forcibly discharged microconidia on spike-like 
sterigmata; species capable of forming capilliconidia or microconidia also 
produce single forcibly discharged secondary conidia; capilliconidia and 
microconidia never formed by the same species. Rhizoids usually absent; 
if present, thicker than conidiophores, terminating in a plate-like holdfast, 
occurring singly, not forming pseudorhizomorphs. Cystidia absent. 
Resting spores globose, with two thickened wall layers; colorless to cream, 
yellowish, or amber in mass; zygospores produced inside larger of two 
(sometimes inconspicuous) conjugating cells, remaining in axis of these 
cells or only slightly displaced laterally from it at maturity (but not budded 
laterally through a narrow isthmus); germinating to produce a germ 
_ conidiophore and germ conidium or limited germ mycelium with several 
germ conidiophores; germ conidia resemble primary conidia and are 
forcibly discharged by papillar eversion. Other resting spore types include 
chlamydospores (usually irregular in shape, with a single thickened wall 
layer) or villose conidia (produced by decoration of primary conidial wall 


456 


with numerous short to long, hair-like appendages). Habit saprobic in soil 
or decaying plant detritus, facultatively (or possibly obligately) pathogenic 
for insects or other arthropods, or weakly (facultatively) pathogenic for 
vertebrates. 


Type species: Conidiobolus utriculosus Brefeld 


This is the family whose membership has been most changed in this 
revision. Macrobiotophthora had been placed in the Entomophthoraceae 
(Ben-Ze'ev & Kenneth, 1982a; Humber 1984a), but is here reassigned to 
the Ancylistaceae on the basis of its nuclear cytology which was studied in 
detail by Tucker (1984). It is also notable that the odd secondary 
capilliconidia of Macrobiotophthora vermicola (Tucker, 1981) are 
orthotropically attached (Humber, 1984a); the only other entomoph- 
thoralean capilliconidia which are so attached are those formed by within 
the subgenus Capillidium of Conidiobolus. Meristacrum, Ballocephala, 
and Zygnemomyces are removed from the Ancylistaceae (Tucker, 1981; 
Ben-Ze'ev & Kenneth, 1982a; Humber 1984a) to a new family, Merista- 
craceae, on the basis of their obligately zoophilic habits and formation of 
multiple conidia on unbranched erect conidiogenous axes. 


MERISTACRACEAE Humber, fam. nov. 


Hyphae vegetativae parietatae coenocyticae primum septatae tandem; cellulae ad septum 
dissociantes et gametangia aut axes conidiogenas externas serviunt. Nuclei parvi, 3-5 um 
diam. per interphasem, nucleolo centrali ovoideo; sine granulis heterochromatis et per 
"aceto-orcein" vel "Bismarck brown" noncoloratis. Mitoses non vidi autem verosimilter 
ancylistoides. Axes simplices multiconidiogenae rectae vel torsivae apicale erecte 
corpore hospitis emergentes. Conidia primaria unitunicata, vehementer eversione 
papillae basalis aut expulsioni retrorsa protoplasmis celluaris tumidae subconidiae 
dispersa aut indehiscentia sterigmatibus angustis. Conidia secundaria conidia primaria 
simulantia expulsa vehementer vel indehiscentia e conidiophoris angustatis aut destitutis. 
Sporae perdurantes plerumque zygosporae, axiale in cellulis parentalis vel ponticuli 
conjugentibus scalariformibus conjugationum diclinearum. Species familiae Ento- 
mophthoralium pathogeniae obligatae hospites invertebratarum solo, nematodarum 
tardigradarum praesertim et insectorum raro. 


Vegetative growth as walled hyphae in host body, coenocytic at first then 
becoming septate and cells becoming disjointed, each segment able to 
give rise to an external hypha bearing conidia or serving as a 
gametangium for zygosporogenesis. Nuclei small, usually 3-5 um long 
during interphase, with prominent central nucleolus; no significant 
quantity of condensed chromatic granular material in nucleoplasm during 
interphase, so nuclei not staining strongly with aceto-orcein or Bismarck 
brown. Nuclei generally difficult to observe; mitosis not observed but 
assumed to be intranuclear (with nuclear envelope intact throughout 
mitosis), with nucleolus persistent and laterally displaced during mitosis; 
chromosomes tiny and indistinct. Conidiophore an unbranched sporo- 
genous axis bearing several conidia. Primary conidia unitunicate, forcibly 


457 


discharged by papillar eversion, by fluid discharge from swollen lateral 
cell interposed between conidium and conidiophore, or passively dis- 
charged from short sterigmatic extension of conidiophore. Secondary 
conidia either resembling primary, forcibly discharged; passively dispersed 
from strongly narrowed, elongate conidiophore; or not formed. Resting 
spores typically zygospores formed from conjugations of adjacent cells in 
hypha or by scalariform conjugations between two hyphae; resting spores 
formed in axis of parental hypha or conjugation tube. Obligately 
pathogens of soil invertebrates, particularly nematodes and tardigrades, 
but also possibly insects. 


Type genus: Meristacrum Drechsler 
Other included genera: Ballocephala Drechsler, Zygnemomyces Miura 


NEOZYGITACEAE Ben-Ze'ev & Kenneth 
in Ben-Ze'ev, Kenneth & Uziel 


Vegetative growth as globose to rod-shaped hyphal bodies; cell walls 
absent or present. Nuclei small, ca. 3-5 um diameter; nucleolus ovoid, 
more or less central; condensed chromatin inconspicuous (by light 
microscopy) during interphase so nuclei stain poorly with aceto-orcein or 
other nuclear stains; mitoses are synchronous in cell, nuclear envelope 
remains intact; metaphase spindle occupies entire nuclear volume, and 
small, vermiform chromosomes are aligned on a conspicuous, central 
plate. Conidiophores simple, cylindrical to slightly clavate; usually pro- 
ducing terminal conidiogenous cell before conidiogenesis or (rarely) 
conidia are produced on and discharged from short conidiophores 
emerging directly from hyphal bodies. Conidia unitunicate, 4 (5) or 7-11 
nucleate; papilla truncate or small; discharged (weakly) by papillar 
eversion; with strong tendency to produce secondary capilliconidia in 
Neozygites. Secondary conidia resembling primary in shape and forcibly 
discharged by papillar eversion or amygdaliform capilliconidia Cusually 
with terminal mucoid hapteron on short peg-like extension) passively dis- 
persed from capillary conidiophores. All spore types (primary, secondary, 
and resting) are melanized (pale grey to smoky grey to dark brown of 
black). Resting spores (zygospores) bud from point of conjugation of two 
hyphal bodies following round of mitosis in both cells; zygospore receives 
one nucleus from each gametangium; epispore of mature zygospores 
strongly melanized and readily detached from endospore; ovoid and 
smooth surfaced or globose to subglobose and roughened; germinating 
directly to produce secondary-type capilliconidium on capillary germ 
conidiophore. Obligately pathogenic for insects (especially Homoptera) 
and mites. 


Type genus: Neozygites Witlaczil 


458 


BASIDIOBOLACEAE Claussen 
in Engler & Gilg 


Vegetative growth as regularly septate mycelium with uninucleate cells or 
as uninucleate yeast form; tending to proliferate by mitosis and plasmo- 
tomy inside cell wall ("Darmform" of Levisohn, 1927). Nuclei large, 
typically longer than 10 um during interphase, with a prominent central 
nucleolus; no condensed chromatic granular material obviously present in 
nucleoplasm during interphase, so nuclei not staining strongly with aceto- 
orcein or bismarck brown. Nuclei becoming indistinct during mitosis (as 
nuclear envelope breaks down and nuclear region is surrounded by 
endoplasmic reticulum); chromosomes tiny, indistinct, and numerous. 
Conidiophores (if present) simple or infrequently branched with a swollen 
apex subtending conidium. Primary conidia Gf present) unitunicate, 
uninucleate; forcibly discharged by backward ejection of conidiophore 
contents upon circumscissile rupture of conidiophore swelling. Resting 
spores and/or chlamydospores formed in axis of parental hypha; resting 
spores typically zygospores resulting from fusion of gametangia after 
formation of short, paired projections which persist on mature spores. 
Occurring as saprobes in soil, colonizers of guts of amphibia or reptiles, or 
as facultative mycotic agent in vertebrates (including humans). 


Type genus: Basidiobolus Eidam 


The genera included in the Neozygitaceae, and Basidiobolaceae are 
unchanged from those previously recognized (Ben-Ze'ev et al., 1986; Ben- 
Ze'ev & Kenneth, 1982a). The familial descriptions of these and all other 
families of the Entomophthorales include all of the characters now 
recognized to be significant. 

A second genus can be attributed to the Basidiobolaceae (Humber, 
1989). The single species in this still undescribed new genus causes fatal, 
disseminated mycoses of snakes (Ippen, 1980; Jessup & Seely 1981; 
Kaplan et al., 1983). An infection of a corn snake originally attributed to a 
colorless alga, Prototheca sp. (Crispens & Marion, 1973), was almost 
certainly caused by this basidiobolaceous fungus. The fungus grows as a 
yeast whose globose cells cleave internally to release 2 or 4 cells upon the 
rupture of the mother cell's wall. Some globose cells may produce an 
simple or infrequently branched hypha of determinate length. This hypha, 
which is nearly always uninucleate, usually forms an ovoid intercalary 
chlamydospore (or zygospore?). Neither forcibly or passively discharged 
conidia are known from this fungus. Interphasic nuclei are large and have 
a prominent central nucleolus. As in Basidiobolus, the light microscopic 
image of nuclei of the snake fungus is less distinct during mitosis than 
during interphase because the nuclear envelope dissociates during mitosis. 
The spindle fills the nucleus at metaphase, and the small chromosomes 
are arranged on a central metaphase plate. 


459 


The taxonomic changes proposed here represent a further maturation 
of the phylogenetically based systematics proposed earlier and provide a 
still greater degree of biological predictability for the classification of the 
Entomophthorales. The acceptance of generic status for each of the four 
Batkoan subgenera of Zoophthora (Batko, 1966; Ben-Ze'ev & Kenneth, 
1982b; Li & Humber, 1984) should remove all remaining differences 
between the adherents of classifications based on phenetics (Remaudiére 
& Hennebert, 1980; Remaudiére & Keller, 1980) or on probable 
phylogenetic relationships (Humber, 1981, 1982, 1984a; Tucker, 1981; 
Ben-Ze'ev & Kenneth, 1982a,b). This change should now enable the 
universal adoption of a single, comprehensive post-Batkoan classification 
for the Entomophthorales. 


ACKNOWLEDGMENT 


I thank Drs. R.P. Korf (Cornell University), G.L. Benny (University of 
Florida, Gainesville), and N. Wilding (Rothamsted Experimental Station, 
U.K.) for their constructive presubmission reviews and comments. Many 
thanks is also due to Dr. T.J. Poprawski (USDA-ARS, European Parasite 
Laboratory) for obtaining and translating a copy of Nowakowski (1881). 


REFERENCES 


Batko, A. 1964. On the new genera: Zoophthora gen. nov., Triplosporium (Thaxter) 
gen. nov., and Entomophaga gen. nov. (Phycomycetes: Entomophthoraceae). Bull. 
Polon. Acad. Sci., Sér. Sci. Biol. 12: 323-326. 

Batko, A. 1966. On the subgenera of the fungus genus Zoophthora Batko 1964 (Ento- 
mophthoraceae). Acta Mycol. 2: 15-21. 

Ben-Ze'ev, I., and R. G. Kenneth. 1982a. Features-criteria of taxonomic value in the 
Entomophthorales: I. A revision of the Batkoan classification. Mycotaxon 14: 393-455. 
Ben-Ze'ev, I., and R. G. Kenneth. 1982b. Features-criteria of taxonomic value in the 
Entomophthorales: II. A revision of the genus Erynia Nowakowski 1881 (= Zoophthora 

Batko 1964). Mycotaxon 14: 456-475. 

Ben-Ze'ev, I., R.G. Kenneth, and A. Uziel. 1987. A reclassification of Entomophthora 
turbinata in Thaxterosporium gen. nov., Neozygitaceae fam. nov. (Zygomycetes: 
Entomophthorales). Mycotaxon 28: 313-326. 

Crispens, C. G., Jr., and K. R. Marion. 1975. Algal infection in a corn snake (Elaphe 
guitata guttata). Lab. Anim. Sci. 25: 788. 

Descals, E., and J. Webster. 1984. Branched aquatic conidia in Erynia and Ento- 
mophthora sensu lato. Trans. Brit. Mycol. Soc. 83: 669-682. 

Humber, R. A. 1981. An alternative view of certain taxonomic criteria used in the Ento- 
mophthorales (Zygomycetes). Mycotaxon 13: 191-240. 

Humber, R. A. 1982. Strongwellsea vs. Erynia: the case for a phylogenetic classification 
of the Entomophthorales (Zygomycetes). Mycotaxon 15: 167-184. 

Humber, R. A. 1984a. Foundations for an evolutionary classification of the Entomoph- 
thorales (Zygomycetes). In: Fungus/insect relationships: perspectives in ecology and 
evolution (Q. Wheeler and M. Blackwell, eds.), pp. 166-183. Columbia University Press, 
New York. 


460 


Humber, R. A. 1984b. The identity of Entomophaga species (Entomophthorales: 
Entomophthoraceae) attacking Lepidoptera: Mycotaxon 21: 265-272. 

Humber, R. A. 1989. The Entomophthorales (Zygomycotina): augmented perspec- 
tives and systematics. In preparation for Mycologia Memoirs. 

Humber, R. A., and I. Ben-Ze'ev. 1981. Erynia (Zygomycetes: Entomophthorales): 
emendation, synonymy, and transfers. Mycotaxon 13: 506-516. 

Ippen, R. 1980. Ein Beitrag zu den Mykosen der Schlangen. Milu (Berlin) 5: 386-396. 

Jessup, D. A., and J. C. Seely. 1981. Zygomycete fungus infection in two captive 
snakes: gopher snake (Pituophis melanoleucos); copperhead (Agkistrodon contor- 
trix). J. Zoo Anim. Med. 12: 54-59. 

Kaplan, W., F. W. Chandler, A. A. Padhye, and T. E. Hamm, Jr. 1983. A zygo- 
mycotic infection in captive snakes. Sabouraudia 21: 85-91. 

Keller, S. 1987. Arthropod-pathogenic Entomophthorales of Switzerland. I. Conidio- 
bolus, Entomophaga, and Entomophthora. Sydowia 40: 122-167. 

Li, Z.. and R. A Humber. 1984. Erynia pieris (Zygomycetes: Entomophthoraceae), 
a new pathogen of Pieris rapae (Lepidoptera: Pieridae): description, host range, and 
notes on Erynia virescens. Canad. J. Bot. 62: 653-663. 

MacLeod, D. M., and E. Miller-Kégler. 1970. Insect pathogens: species originally 
described from their resting spores mostly as Tarichium species (Entomophthor- 
ales: Entomophthoraceae). Mycologia 62: 33-66. 

Nowakowski, L. 1881. O grupie owadomork6w (Entomophthoreae). Dziennik III 
Zjazdu Lekarzy i Przyrodnikow Polskich w Krakowie 6: 67-68. 

Remaudiére, G., and G. L. Hennebert. 1980. Révision systématique de Entomoph- 
thora aphidis Hoffm. in Fres. Description de deux nouveaux pathogénes d'aphides. 
Mycotaxon 11: 269-321. 

Remaudiére, G., and S. Keller. 1980. Révision systématique des genres d'Ento- 
mophthoraceae 4 potentialité entomopathogéne. Mycotaxon 11: 323-338. 

Soper, R. S., B. May, and B. J. Martinell. 1983. Entomophaga gryili enzyme poly- 
morphism as a technique for pathotype identification. Envir. Entomol. 12: 720-723. 

Steinkraus, D. C., and J. P. Kramer. 1988. Entomophthora scatophagae desc. 
ampl. (Zygomycetes: Entomophthorales), a fungal pathogen of the yellow dung fly, 
Scatophaga stercoraria (Diptera: Anthomyiidae). Mycotaxon 32: 105-113. 

Thaxter, R. 1888. The Entomophthoreae of the United States. Mem. Boston Soc. 
Nat. Hist. 4: 133-201. 

Tucker, B. E. 1981. A review of the nonentomogenous Entomophthorales. Myco- 
taxon 13: 481-505. 


Tucker, B. E. 1984. Aspects of the biology and ultrastucture of the nematode- 
destroying fungus Macrobiotophthora vermicola (Zygomycetes: Entomophthor- 
ales). Ph.D. dissertation, University of Washington, Seattle. 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 461-473 January-March 1989 


MNEMONIC THREE-LETTER ACRONYMS 
FOR THE NAMES OF FUNGAL FAMILIES 


JOSEPH Gh LE ERE LERE: 


Department of Ecology and Evolutionary Biology 
University of Arizona, Tucson AZ 85721 USA 


In the appendices of this article are listed three- 
letter abbreviations for the names of 629 families of 
fungaw  finis: (work Was inspired by a similar article 
Suggesting standardized three-letter acronyms’ for the 
Families of vascular plants CWeber 1982). Appendix I lists 
the families alphabetized according to the full name of the 
Family, while in Appendix II the same information is 
alphabetized according to the abbreviation. 

‘There are several advantages to such a system of 
Standardized abbreviations. The acronyms can be printed 
directly on herbarium labels, conveying a considerable 
amount of taxonomic information in a minimal amount of 
Space. A display containing the codes and their meanings 
can be posted at conspicuous locations in the herbarium to 
allow users to interpret the codes rapidly. The 
abbreviations can also be used to save storage space Cand 
data-entry time) in computerized databases. Programs can 
easily be written which can translate the abbreviations 
into full names if the need arises. Standardization of such 
names will enable workers from different institutions to 
share information more readily. 

As with the listing of plant families, the codes 
presented here were chosen mnemonically to make rapid 
memorization and interpretation possible. Many of the codes 
represent the first three letters of the actual name. In 
most other cases, they were constructed From the first 
letter of the name plus two consonants chosen from the 
other letters in the name. 

Compiling the list of currently-used family names 
proved difficult given the state of Flux in which Fungal 
taxonomy finds itself at the present time. Several 
Classification systems are being used by different workers, 
and new Families are still being described rather 
Frequently. The intent here is not to pass judgment on the 
merits of any particular classification scheme but rather 
to supply an indexing system useful to mycologists 
belonging to several schools of thought. The list was 
compiled primarily from Ainsworth et al. €1973), Bessey 
Poo) EcCiksson s&sHawksworth Cisse7o;. Jdbich c1gs4o. and 
Alexopoulos (1962), supplemented with information drawn 
From recent monographs on various taxa. Form-families of 
Fungi Imperfecti were included in the interests of 


462 


completeness. Some older families were included under the 
assumption that these names are still being used by some 
workers, but very old families which appear to have totally 
disappeared from use have been omitted. 

The original dBase II files containing family names 
and abbreviations are available from the author for $10.00. 


Dl LER CORES ei) 


Ainsworth, G.C., Frederick K. Sparrow, & Alfred S. Sussman 
Ceds.3 €1973) The Fungi: an advanced. treatise, vol. 
IV. New York: Academic Press. 

Bessey, Ernst Althearn €1950) Morphology and taxonomy of 
Fungi. Philadelphia: Blakiston. 

Eriksson, Ove E., & David L. Hawksworth €1987) Outline of 
the Ascomycetes - 1987. Systema Ascomycetum 6C2):eS9- 
33.4% 

JUilich, Walter (1984) Kleine Kryptogamenflora, Band IIb/1: 
Basidiomyceten, erste Teil: Die Nichtblatterpilze, 
Gallertpilze, und Brauchpilze. Stuttgart: Gustav 
Fischer. 

Weber, William A. €1982) Mnemonic three-letter acronyms for 
the families of vascular plants: a device for more 
effective herbarium curation. Taxon 61(€1):74-6B. 


ACKNOWLEDGEMENTS 
I would like to thank Ove’ Eriksson,” Walter =J0grcw 
William Weber, and Harry Thiers for helpful comments on an 


earlier draft of this manuscript. 


APPENDIX I: FAMILIES AND ACRONYMS ALPHABETIZED BY FAMILY 


Acarosporaceae ACA Anisolpidiaceae ANI 
Acetabulaceae ACE Antennulariellaceae ANN 
Achaetomiaceae ACH Anthostomataceae ANT 
Achlyogetonaceae ACG Anziaceae ANZ 
Acrasiaceae ACR Aphelariaceae APH 
Acrocordiaceae ACC Aporpiaceae APO 
Acrospermaceae ACS Arachniaceae ARA 
Actinoplacaceae ACT Arctomiaceae ARC 
Acytosteliaceae ACY Arcyriaceae ARR 
Agaricaceae AGA Argunnaceae ARG 
Aguriaceae AGY Arthoniaceae ART 
Albuginaceae ALB Arthopyreniaceae ATP 
Alectoriaceae ALE Arthrodermataceae ADR 
Aleuriaceae ALR Arthrorhapidaceae ATR 
Aleuriomycetaceae ANC Arundinulaceae ARU 
Allantosphaeriaceae ALL Ascobolaceae ASB 
Amanitaceae AMA Ascocortiaceae ASC 
Amauroascaceae ANR Ascodichaenaceae ADC 
Amoebidiaceae AMB Ascohymeniaceae AHM 
Amorphothecaceae AMO Ascoideaceae ASD 
Amphisphaeriaceae AMP Ascosphaeraceae ASP 
Amylariaceae ANY Asellariaceae ASE 
Anapyreniaceae ANA Aspergillaceae APG 


Ancylistidaceae ANC Aspidotheliaceae ATH 


Asterinaceae 
Asterothyriaceae 
Astraeaceae 
Astrogastraceae 
Astrotheliaceae 
Atichiaceae 
Aulographaceae 
Auriculariaceae 
Auriscalpiaceae 
Bacidiaceae 
Bactrosporaceae 
Baeomycetaceae 
Balsamiaceae 
Bankeraceae 
Basidiobolaceae 
Biatoraceae 
Blastocladiaceae 
Bolbitiaceae 
Boletaceae 
Boliniaceae 
Bondarzewiaceae 
Botryosphaeriaceae 
Brachybasidiaceae 
Brefeldiellaceae 
Brigantiaeaceae 
Broomeiaceae 
Buelliaceae 
Cainiaceae 
Caliciaceae 
Caloplacaceae 
Calosphaeriaceae 
Calostomataceae 
Calycidiaceae 
Candelariaceae 
Cantharellaceae 
Capnodiaceae 
Carbomycetaceae 
Catabotryudaceae 
Catenariaceae 
Catillariaceae 
Cavosteliaceae 
Cephaloascaceae 
Cephalothecaceae 
Ceratiomyxaceae 
Ceratobasidiaceae 
Ceratomycetaceae 
Ceratostomataceae 
Chadefaudiellaceae 
Chaetomiaceae 
Chaetothyriaceae 
Chiodectonaceae 
Choanephoraceae 
Chryusotrichaceae 
Chytridiaceae 
Cladochytriaceae 
Cladoniaceae 
Clastodermataceae 


ASN 
AST 
ASR 
ASG 
ASL 
ATI 

AUL 
AUR 
ARS 
BCD 
BAC 
BAE 
BAL 
BNK 
BAS 
BIA 
BLA 
BLB 
BE 
BLN 
BDZ 
BOT 
BRA 
BEL 
BRI 
BRO 
BUE 
CAI 

GEG 
Ge 
Gis 
Gish 
CAL 
CND 
CNT 
CAP 
CAR 
CITB 
CAT 
Gi 
CAV 
CES 
CER 
CMxX 
EES 
Eve 
Goa 
CDF 
Sn 
opine 
Git 
CHO 
CHR 
CHY 
CDC 
CLA 
CDR 


Clathraceae 
Claustulaceae 
Clavariaceae 
Clavariadelphaceae 
Clavicipitaceae 
Clavicoronaceae 
Clavulinaceae 
Clypeosphaeriaceae 
Coccocarpiaceae 
Coccodiniaceae 
Coccoideaceae 
Coccotremataceae 
Cochlonemaceae 
Coelomomycetaceae 
Coenogoniaceae 
Coleosporiaceae 
Collemataceae 
Coniochaetaceae 
Coniocybaceae 
Coniophoraceae 
Cookellaceae 
Coprinaceae 
Corneromycetaceae 
Coronophoraceae 
Corticiaceae 
Cortinariaceae 
Coryneliaceae 
Crepidotaceae 
Cribbiaceae 
Cribrariaceae 
Crocyniaceae 
Cronartiaceae 
Cryptococcaceae 
Cryptomycetaceae 
Cucurbitariaceae 
Cunninghamellaceae 
Cyuanocephalaceae 
Cypheliaceae 
Cyphellaceae 
Cyustobasidiaceae 
Cyttariaceae 
Dacampiaceae 
Dacryumycetaceae 
Dematiaceae 
Dendrosphaeraceae 
Dentinaceae 
Dermateaceae 
Dermatocarpaceae 
Dexteriaceae 
Dianemaceae 
Diaporthaceae 
Diatrypaceae 
Dictyopeltinaceae 
Dictyosteliaceae 
Didymiaceae 
Didymosphaer iaceae 
Dimargaritaceae 


463 


CTR 
ESE 
CUR 
CDL 
CVUC 
EGR 
COL 
CLY 
EP 
CDN 
cco 
CCL 
ECE 
GUE 
CNG 
Bok 
GLE 
CNC 
CON 
CNS 
COO 
COP 
CMT 
Oe 
CRT 
CTN 
CRN 
CRE 
CRB 
CRI 
CRC 
CRO 
GRE 
CRM 
CUC 
CUN 
CYA 
GE 
CEE 
EY 
[Eig 8 
DCM 
DCR 
DEM 
DND 
DNT 
DRM 
DMC 
DEX 
DNM 
DEL 
Ler 
DRL 
DIC 
DID 
DDM 
DMG 


464 


Dimeriaceae 
Diploschistaceae 
Dipodascaceae 
Discellaceae 
Dothideaceae 
Dothioraceae 
Ecchynaceae 
Eccrinaceae 
Echinodontiaceae 
Echinosteliaceae 
Ectolechiaceae 
Ectrogellaceae 
Figleriaceae 
Elaeomyxaceae 
Elaphomycetaceae 
Elasmomyucetaceae 
Elsinoaceae 
Endochytriaceae 
Endogonaceae 
Endomycetaceae 
Englerulaceae 
Enteridiaceae 
Entolomataceae 
Entomophthoraceae 
Entophlyuctaceae 
Eocronartiaceae 
Eoterfeziaceae 
Ephebaceae 
Eremascaceae 
Eremomycetaceae 
Erysiphaceae 
Euantennariaceae 
Fuceratomycetaceae 
Eurotiaceae 
EFurychasmaceae 
Excipulaceae 
Exidiaceae 
Exobasidiaceae 
Fassariaceae 
Favolaschiaceae 
Fenestellaceae 
Filobasidiaceae 
Fimetariaceae 
Fistulinaceae 
Fuckeliaceae 
Fuscideaceae 
Galeropsidaceae 
Ganodermataceae 
Gasterellaceae 
Gastrosporiaceae 
Gautieriaceae 
Geastraceae 
Gelatinodiscaceae 
Gelopellidaceae 
Geneaceae 
Genistellaceae 
Geoglossaceae 


DMR 
DIP 
DPD 
DIS 
DID 
DTR 
ECN 
ECC 
ECD 
ECS 
Eee 
ECG 
EIG 
EMX 
ELA 
ESI 
ELS 
ENC 
END 
a 
ENG 
ENR 
ENL 
ENM 
ENP 
Ee 
EDEL 
EPH 
ERE 
ERM 
ERY 
EUA 
EUG 
EUR 
ERC 
EAC 
EX 

EXO 
FAS 
FUL 
FEN 
pd Ge 
Ban 
mis 
FUC 
rus 
GAL 
GAN 
G55 
GSP 
GAU 
GEA 
GLT 
Beye! 
GEN 
GNS 
GEO 


Glaziellaceae 
Glischrodermataceae 
Gnomoniaceae 
Gomphaceae 
Gomphidiaceae 
Gomphillaceae 
Gonapodyaceae 
Graphidaceae 
Graphiolaceae 
Guttulinaceae 
Gualectaceae 
Gyumnoascaceae 
Gyrophoraceae 
Haematommataceae 
Haliphthoraceae 
Halosphaeriaceae 
Haplosporaceae 
Harpellaceae 
Harpidiaceae 
Harpochytriaceae 
Helicocephalidaceae 
Helocarpaceae 
Helotiaceae 
Helvellaceae 
Hemiascaceae 
Hemiascosporiaceae 
Hemigastraceae 
Hemiphacidiaceae 
Hemisphaeriaceae 
Heppiaceae 
Hericiaceae 
Herpomycetaceae 
Herpotrichiellaceae 
Heterodeaceae 
Heterosctyphaceae 
Hoehnelomyucetaceae 
Huiliaceae 
Humariaceae 
Hyalographaceae 
Hyaloriaceae 
Hyaloscyphaceae 
Hydnaceae 
Hyudnangiaceae 
Hygrophoraceae 
Hymeneliaceae 
Hymenochaetaceae 
Humenogastraceae 
Hyphochytriaceae 
Hypocreaceae 
Hypodermataceae 
Hypogyumniaceae 
Hypomycetaceae 
Hyponectriaceae 
Hysterangiaceae 
Hysteriaceae 
Imbricariaceae 
Ixechinaceae 


GL2 
GLI 
GNO 
GMP 
GOM 
GMP 
GON 
GRD 
GRL 
GUT 
GYA 
GYM 
GYR 
HAE 
HLP 
HES 
HPS 
HAR 
HRP 
HCH 
NnGr 
HEL 
HLT 
HLU 
HSC 
HMC 
HGS 
HPC 
HMS 
HEP 
HRC 
HRM 
HiT 
HTD 
HTS 
HOE 
HUI 
HUM 
HLG 
HYA 
HLC 
HDN 
HNG 
HYG 
HML 
HCT 
HYM 
HPH 
HCR 
HDR 
HGM 
He 
HNC 
HSG 
HYS 
IMB 
IXE 


Jugasporaceae 
Karstenellaceae 
Kickxellaceae 
Koerberiellaceae 
Koralionastetaceae 
Kriegeriellaceae 
Laboulbeniaceae 
Labyrinthulaceae 
Laccariaceae 
Lachnocladiaceae 
Lagenidiaceae 
Lahmiaceae 
Lamprodermaceae 
Lanopilaceae 
Lasiolomataceae 
Lasiosphaeriaceae 
Lecanactidaceae 
Lecanidiaceae 
Lecanoraceae 
Lecideaceae 
Lecidomataceae 
Lecotheciaceae 
Leger iomycetaceae 
Leotiaceae 
Lepiotaceae 
Leptolegniellaceae 
Leptomitaceae 
Leptopeltidaceae 
Leptosphaeriaceae 
Leptostromataceae 
Letrouitiaceae 
Leucogastraceae 
Leveillellaceae 
Liceaceae 
Lichenotheliaceae 
Lichinaceae 
Limboriaceae 
Lipomycetaceae 
Lithographaceae 
Lobariaceae 
Lopadiaceae 
Lophiaceae 
Lophiostomataceae 
Lophiotrichaceae 
Lophophytomaceae 
Loramycetaceae 
Lycogalaceae 
Lycoperdaceae 
Macrocystidiaceae 
Marasmiaceae 
Massarinaceae 
Mastodiaceae 
Medeolariaceas 
Megachytriaceae 
Megalariaceae 
Megalosporaceae 
Melampsoraceae 


JGS 
KAR 
KXL 
KOE 
KOR 
KRI 
MERE} 
LBR 
LCR 
LAC 
LAG 
LAH 
LAM 
LNP 
LLM 
LAS 
LEG 
LCN 
LNR 
LCD 
LDM 
LET 
LEG 
LEO 
EPt 
LLG 
LPM 
LPL 
Lor 
HES he 
Oe 8 
LSU 
LEV 
LiG 
LTH 
LGH 
LIM 
LeeLee 
Paha 
LOB 
eis 
LPH 
LUE 
LIR 
CPE 
LRM 
LGE 
LCP 
MAC 
MAR 
MAS 
MST 
MED 
MEG 
MGL 
MSR 
MLM 


Melanconiaceae 
Melanconidaceae 
Melanogastraceae 
Melanommataceae 
Melanosporaceae 
Meliolaceae 
Melogrammataceae 
Meruliaceae 
Mesnieraceae 
Mesophelliaceae 
Metacapnodiaceae 
Metschnikowiaceae 
Micareaceae 
Microascaceae 
Microglaenaceae 
Micropeltidaceae 


Microtheliopsidaceae 


Microthyriaceae 
Miltideaceae 
Mixiaceae 
Mollisiaceae 
Monascaceae 
Moniliaceae 
Monoblepharidaceae 
Montagneaceae 
Montagnellaceae 
Morchellaceae 
Moriolaceae 
Mortierellaceae 
Mucoraceae 
Munkiellaceae 
Mycenastraceae 
Mycobilimbiaceae 
Mycoblastaceae 
Myucocaliciaceae 
Mycoporaceae 
Mycosphaerellaceae 
Mycotyphaceae 
Myriangiaceae 
Mytilinidiaceae 
Myxotrichaceae 
Nectaromycetaceae 
Nectriaceae 
Nectrioidaceae 
Nematosporaceae 
Neolectaceae 
Nephromataceae 
Nidulariaceae 
Nitschkiaceae 
Octavianinaceae 
Odontotremataceae 
Olpidiaceae 
Olpidiopsidaceae 
Omphalotaceae 
Onygenaceae 
Opegraphaceae 
Ophioparmaceae 


465 


MLC 
MLD 
MGS 
MLN 
HLS 
MEL 
MGR 
MER 
MSN 
HSe 
MET 
MKW 
MCR 
MIC 
MCG 
NCP 
MTL 
MTR 
MPL 
MIX 
MOL 
MNS 
MNL 
MBL 
MNT 
MTG 
MRC 
MRL 
MRT 
MUC 
MNK 
MCS 
MBB 
MCB 
MCL 
MYC 
MSL 
MCT 
MYR 
MYT 
MXT 
NMC 
NEC 
NCD 
NEM 
NEO 
NEP 
NID 
NIT 
Gil 
ODO 
OLP 
ODP 
OMP 
ONY 
OPE 
UE 


466 


Ophiostomataceae 
Oplotheciaceae 
Orbiliaceae 
Orphniosporaceae 
Ostropaceae 
Otideaceae 
Pachyascaceae 
Palavasciaceae 
Pannariaceae 
Parataeniellaceae 
Parmeliaceae 
Parmulariaceae 
Parodiellaceae 
Parodiopsidaceae 
Paropsidaceae 
Patellariaceae 
Paxillaceae 
Peltigeraceae 
Peltulaceae 
Perichaenaceae 
Pericystaceae 
Peridiaceae 
Perisporiaceae 
Peronophythoraceae 
Peronosporaceae 
Pertusariaceae 
Peyritschiellaceae 
Pezizaceae 
Phacidiaceae 
Phaeomonomeraceae 
Phaeophragmiaceae 
Phaeosphaeriaceae 
Phaeotrichaceae 
Phallaceae 
Phaneromycetaceae 
Phillipsiellaceae 
Phleogenaceae 
Phlyuctidiaceae 
Phragmopelthecaceae 
Phylaciaceae 
Phyllachoraceae 
Phyllobatheliaceae 
Physalaciaceae 
Physaraceae 
Physciaceae 
Physodermataceae 
Physosporellaceae 
Piedraiaceae 
Pilobolaceae 
Pilocarpaceae 
Piptocephalidaceae 
Pisolithaceae 
Pithoascaceae 
Placolecidaceae 
Placynthiaceae 
Plasmodiophoraceae 
Plectascaceae 


OPH 
OPL 
ORB 
ORP 
Opi 
OTI 
RES 
PLU 
PAN 
PIN 
PRM 
GHEE 
Bue 
Bur 
Res 
Pad 
BAL 
Be 
ind Oe 
Pen 
PASE 
PRD 
RRS 
PEE 
PRN 
PRT 
REY 
REZ 
Rog 
PHM 
BEG 
oh 
Pry 
PHA 
PNM 
PHI 
REG 
Ree 
PHR 
PILY 
PEG 
Rib 
Boe 
PSR 
Bas 
PDR 
Bos 
Ge 
BREE 
PGR 
itis 
RIS 
Pst 
PLA 
PCN 
PMR 
PLE 


Pleomassariaceae 
Pleosporaceae 
Pleurotaceae 
Pluteaceae 
Podaxaceae 
Podoscyphaceae 
Polyporaceae 
Polystigmataceae 
Polystomellaceae 


Polystomellopsidaceae 


Porinaceae 
Porpidiaceae 
Protogastraceae 
Protomycetaceae 
Protophallaceae 
Protosteliaceae 
Pseudeurotiaceae 
Pseudorhizinaceae 
Pseudosphaer iaceae 
Pseudovalsaceae 
Psoraceae 
Psorulaceae 
Pterulaceae 
Pucciniaceae 
Punctulariaceae 
Pycnothyriaceae 
Pyrenidiaceae 
Pyrenophoraceae 
Pyrenopsidaceae 
Pyrenotrichaceae 
Pyrenulaceae 
Pyronemataceae 
Pythiaceae 
Pyxidiophoraceae 
Radiomyucetaceae 
Ramalinaceae 
Rechingeriellaceae 
Requienellaceae 
Reticulariaceae 
Rhipidiaceae 
Rhizidiaceae 
Rhizidiomycetaceae 
Rhizinaceae 
Rhizocarpaceae 
Rhizopogonaceae 
Rhizothyriaceae 
Rhodataceae 
Rhodophyllaceae 
Rhodotorulaceae 
Rhopalosporaceae 
Rhytismataceae 
Richoniellaceae 
Rimulariaceae 
Roccellaceae 
Roccellinastraceae 
Russulaceae 
Saccardiaceae 


PLM 
PLS 
PLR 
PLU 
PDX 
PDS 
PLP 


es pe | 


PSM 
Pot 
POR 
PRU 
PGS 
PMC 
PPH 
Row 
PEU 
Pre 
Foe 
PUL 
PSO 
PRL 
Pipe 
PUG 
PUN 
PYC 
PND 
Led cf ee 
PNS 
PTR 
PNL 
RI 
Pxt 
BY 
RAD 
RAM 
RCG 
REQ 
RNC 
RPD 
RZD 
R2M 
RZN 
RCR 
R2P 
R2T 
RDT 
RPL 
RHO 
RSP 
RHY 
RIC 
RIM 
ROC 
RLN 
RUS 
SRD 


Saccardinulaceae 
Saccharomycetaceae 
Saccharomycodaceae 
Saccharomycopsidaceae 
Sakseneaceae 
Sappiniaceae 
Saprochaetaceae 
Saprolegniaceae 
Sarcoscyphaceae 
Sarcosomataceae 
Sarrameanaceae 
Saturnopichiaceae 
Schadoniaceae 
Schaereriaceae 
Schenellaceae 
Schizophyilaceae 


Schizosaccharomycetaceae 


Heures ociiaaceac 
Sclerodermataceae 
Sclerophoraceae 
Sclerotiniaceae 
Scoliciosporaceae 
Scutigeraceae 
Secotiaceae 
Septobasidiaceae 
Seuratiaceae 
Sirobasidiaceae 
Sirolpidiaceae 
Solorinaceae 
Sordariaceae 
Sparassidaceae 
Spathulosporaceae 
Spermophthoraceae 
Sphaeriaceae 
Sphaerioidaceae 
Sphaerobolaceae 
Sphaerophoraceae 
Sphaerophoropsidaceae 
Sphaeropsidaceae 
Sphinctrinaceae 
Spilomataceae 
Spizellomycetaceae 
Sporobolomycetaceae 
Sporormiaceae 
Squamar inaceae 
Stegillaceae 
Stemonitaceae 
Stephanosporaceae 
Stephanothecaceae 
Stereaceae 
Stereocaulaceae 
Stictaceae 
Stictidaceae 
Stigmataceae 
Stigmateaceae 
Stilbaceae 
Stilbellaceae 


SDN 
SAC 
SMD 
SCM 
SAK 
SPN 
Site 
SiG 
‘Sop e 
bl 
SAR 
SL 
ser 
SCR 
SGN 
ey we 
S25 
Spal 
SDR 
Sil 
SIN 
SCO 
Seu 
ey ay B 
Sen 
SEU 
Spo 
SRL 
SOL 
SOR 
Sou 
SPA 
Site 
sot Sa 
2) AB) 
BBL 
SER 
ah ed i 
SPS 
SC 
SPI 
SEZ 
SPB 
SRM 
SQU 
S6r 
STM 
SoR 
Sieh 
STR 
Sn 
oe ® 
SiO 
SGM 
SMT 
STB 
Sit 


Stomatogenel laceae 
Strigulaceae 
Strobilomycetaceae 
Strophariaceae 
Suncephalastraceae 
Syunchytriaceae 
SuzZzugosporaceae 
Taeniellaceae 
Taphrinaceae 
Teloschistaceae 
Tephromelataceae 
Terfeziaceae 
Testudinaceae 
Tetragoniomycetaceae 
Thamnidiaceae 
Thelebolaceae 
Thelephoraceae 
Thelotremataceae 
Thermoascaceae 
Thielaviascaceae 
Thraustochytriaceae 
Thrombiaceae 
Thryptosporaceae 
Thyuridiaceae 
Tilletiaceae 
Torrendiaceae 
Torulopsidaceae 
Trapeliaceae 
Traustochytriaceae 
Tremellaceae 
Tremoleciaceae 
Triblidiaceae 
Trichiaceae 
Trichocomaceae 
Tricholomataceae 
Trichopeltaceae 
Trichopeltidaceae 
Trichopeltinaceae 
Trichosphaeriaceae 
Trichotheliaceae 
Trichothyriaceae 
Triposporiopsidaceae 
Trybilidiaceae 
Trypetheliaceae 
Tuberaceae 
Tuberculariaceae 
Tubeuf iaceae 
Tulasnellaceae 
Tulostomataceae 
Umbilicariaceae 
Uredinaceae 
Usneaceae 
Ustillaginaceae 
VYValsaceae 
Venturiaceae 
Verrucariaceae 
Vezdaeaceae 


467 


SGN 
STG 
SBM 
pee to 
SER 
SNC 
IVS 
CAE 
TAP 
TES 
ate 
MSs 
Es 
NS) Sel 
THA 
PEE 
LP 
He Shy 
TRM 
THYV 
TCH 
THR 
LAG 
EY 
Ji a0 w. 
TRN 
TRE 
ERP 
TRA 
TRE 
LEG 
TBD 
TRE 
TEE 
fee 
vis 
Lee 
TPN 
TRS 
Tike 
TRI 
Dor 
TRB 
ray 
TUB 
BG 
TBF 
TSN 
LES 
UMB 
URE 
USN 
BS 
VAL 
VEN 
VER 
VEZ 


468 


Vialaeaceae VIA Zerovaemycetaceae ZRYU 
Vizellaceae VIZ Zodiomycetaceae ZOD 
Woroninaceae WOR Zoopagaceae 200 
Xanthopsorellaceae XAN Zopf iaceae ZOP 
Xerulaceae XER Zythiaceae Ke 39 
Xylariaceae XYL 


APPENDIX II: FAMILIES AND ACRONYMS ALPHABETIZED BY ACRONYM 


ACA Acarosporaceae ASP Ascosphaeraceae 
ACC Acrocordiaceae ASR Astraeaceae 

ACE Acetabulaceae AST Asterothyriaceae 
ACG Achlyogetonaceae ATH Aspidotheliaceae 
ACH Achaetomiaceae ATI Atichiaceae 

ACR Acrasiaceae ATP Arthopyreniaceae 
ACS Acrospermaceae ATR Arthrorhapidaceae 
ACT Actinoplacaceae AUL Aulographaceae 
ACY Acytosteliaceae AUR Auriculariaceae 
ADC Ascodichaenaceae BAC Bactrosporaceae 
ADR Arthrodermataceae BAE Baeomycetaceaes 
AGA Agaricaceae BAL Balsamiaceae 

AGY Agyuriaceae BAS Basidiobolaceae 
AHM Ascohymeniaceae BCD Bacidiaceae 

ALB Albuginaceae BDZ BondarZzewiaceae 
ALE Alectoriaceae BFL Brefeldiellaceae 
ALL Allantosphaeriaceae BIA Biatoraceae 

ALR Aleuriaceae BLA Blastocladiaceae 
AMA Amanitaceae BLB Bolbitiaceae 

ANB Amoebidiaceae BLN Boliniaceae 

ANC Aleuriomycetaceae BL Boletaceae 

ANO Amor phothecaceae BNK Bankeraceae 

AMP Amphisphaeriaceae BOT Botryosphaeriaceae 
ANR Amauroascaceae BRA Brachybasidiaceae 
ANY Amylariaceae BRI Brigantiaeaceae 
ANA Anapyreniaceae BRO Broomeiaceae 

ANC Ancylistidaceae BUE Buelliaceae 

ANI Anisolpidiaceae CAI Cainiaceae 

ANN Antennulariellaceae CAL Calycidiaceae 

ANT Anthostomataceae CAP Capnodiaceae 

ANZ Anziaceae CAR Carbomycetaceae 
APG Aspergillaceae CAT Catenariaceae 

APH Aphelariaceae CAV Cavosteliaceae 
APO Aporpiaceae ESS Ceratobasidiaceae 
ARA Arachniaceae a Wy Coccoideaceae 

ARC Arctomiaceae EGis Cochlonemaceae 
ARG Argynnaceae OF we Coccocarpiaceae 
ARR Arcyriaceae CCR Clavicoronaceae 
ARS Auriscalpiaceae Cor Coccotremataceae 
ART Arthoniaceae des We Cladochytriaceae 
ARU Arundinulaceae CDF Chadefaudiellaceae 
ASB Ascobolaceae COL Clavariadelphaceae 
ASC Ascocortiaceae CDN Coccodiniaceae 
ASD Ascoideaceae CDR Clastodermataceae 
ASE Asellariaceae CEP Cephalothecaceae 
ASG Astrogastraceae CHI Chiodectonaceae 
ASL Astrotheliaceae CHO Choanephoraceae 


ASN Asterinaceae CHR Chrysotrichaceae 


CHY 
CLA 
GLC 
one 
CLP 
CLS 
CLT 
CLY 
Cnc 
CNT 
cnx 
CNC 
CND 
CNG 
CNS 
CNT 
COE 
CON 
coo 
coP 
CPC 
CPL 
CPS 
CRB 
CRC 
CRE 
CRI 
CRM 
CRN 
CRO 
CRP 
CRT 
CSL 
CSP 
CST 
CIB 
OTL 
CIM 
CIN 
CTR 
Gir 
cuc 
CUN 
CUC 
CUL 
CUR 
CYA 
CYP 
cys 
CYT 
pcm 
DCR 
pom 
DEM 
DEX 
DIC 
DID 


Chytridiaceae 
Cladoniaceae 
Caliciaceae 
Collemataceae 
Caloplacaceae 
Calosphaeriaceae 
Calostomataceae 
Clypeosphaeriaceae 
Ceratomycetaceae 
Corneromycetaceae 
Ceratiomyxaceae 
Coniochaetaceae 
Candelariaceae 
Coenogoniaceae 
Coniophoraceae 
Cantharellaceae 
Coelomomycetaceae 
Coniocybaceae 
Cookellaceae 
Coprinaceae 
Cryptococcaceae 
Cyphellaceae 
Cephaloascaceae 
Cribbiaceae 
Crocyniaceae 
Crepidotaceae 
Cribrariaceae 
Cryptomycetaceae 
Coryneliaceae 
Cronartiaceae 
Coronophoraceae 
Corticiaceae 
Claustulaceae 
Coleosporiaceae 
Ceratostomataceae 
Catabotrydaceas 
Catillariaceae 
Chaetomiaceae 
Cortinariaceae 
Clathraceae 
Chaetothyriaceae 
Cucurbitariaceae 
Cunninghamellaceae 
Clavicipitaceae 
Clavulinaceae 
Clavariaceae 
Cyanocephalaceae 
Cypheliaceae 
Cystobasidiaceae 
Cyttariaceae 
Dacampiaceae 
Dacrymycetaceae 
Didymosphaer iaceae 
Dematiaceae 
Dexteriaceae 
Dictyosteliaceae 
Didymiaceae 


UU ee 
DIS 
DNC 
DMG 
DMR 
DND 
DNM 
DNT 
DPD 
DPL 
DPT 
DRM 
DID 
DLP 
DIR 
EEG 
EGU 
ECG 
ECN 
EGS 
Eats 
ELA 
ELG 
EES 
EMC 
EMX 
ENC 
END 
ENG 
ENL 
ENM 
ENP 
ENR 
EOC 
EOT 
EPH 
ERG 
ERE 
ERM 
Pay 
ESM 
EUA 
EUG 
EUR 
EXC 
EAL 
EXO 
FAS 
FEN 
Fee: 
FIM 
PLS 
Fug 
EUS 
RUE 
GAL 
GAN 


469 


Diploschistaceae 
Discellaceae 
Dermatocarpaceae 
Dimargaritaceae 
Dimeriaceae 
Dendrosphaeraceae 
Dianemaceae 
Dentinaceae 
Dipodascaceae 
Dictyopeltinaceae 
Diaporthaceae 
Dermateaceae 
Dothideaceae 
Diatrypaceae 
Dothioraceae 
Eccrinaceae 
Echinodontiaceae 
Ectrogellaceae 
Ecchynaceae 
Echinosteliaceae 
Eigleriaceae 
Elaphomycetaceae 
Ectolechiaceae 
Elsinoaceae 
Endomycetaceae 
Elaeomyxaceae 
Endochytriaceae 
Endogonaceae 
Englerulaceae 
Entolomataceae 
Entomophthoraceae 
Entophlyctaceae 
Enteridiaceae 
Eocronartiaceae 
Foterfeziaceae 
Ephebaceae 
Eurychasmaceae 
Eremascaceae 
Eremomycetaceae 
Erysiphaceae 
Elasmomycetaceae 
Euantennariaceae 
Euceratomycetaceae 
Eurotiaceae 
Excipulaceae 
Exidiaceae 
Exobasidiaceae 
Fassariaceae 
Fenestellaceae 
Filobasidiaceae 
Fimetariaceae 
Fistulinaceae 
Fuckeliaceae 
Fuscideaceae 
Favolaschiaceae 
Galeropsidaceae 
Ganodermataceae 


470 


GAU 
GEA 
GEN 
GEO 
el 
GLP 
GLE 
GL2Z 
GMP 
GMP 
GNO 
GNS 
GOM 
GON 
GRD 
GRL 
GSL 
B58 
GUT 
GYA 
GYM 
GYR 
HAE 
HAR 
HCH 
HCP 
HCR 
Tesh 
HDN 
HDR 
HE 
HEP 
HGM 
HGS 
HEC 
HLG 
HED 
HES 
HLT 
HLU 
HMC 
HML 
HMS 
HNC 
HNG 
HOE 
Hee 
HPH 
ins 
HED 
HRC 
HRM 
HRP 
Hoc 
HSG 
HTD 
hg 


Gautieriaceae 
Geastraceae 
Geneaceae 
Geoglossaceae 
Glischrodermataceae 
Gelopellidaceae 
Gelatinodiscaceae 
Glaziellaceae 
Gomphaceae 
Gomphillaceae 
Gnomoniaceae 
Genistellaceae 
Gomphidiaceae 
Gonapodyaceae 
Graphidaceae 
Graphiolaceae 
Gasterellaceae 
Gastrosporiaceae 
Guttulinaceae 
Gyualectaceae 
Gyumnoascaceae 
Gyrophoraceae 
Haematommataceae 
Harpellaceae 
Harpochytriaceae 
Helicocephalidaceae 
Hypocreaceae 
Hymenochaetaceae 
Hydnaceae 
Hypodermataceae 
Helocarpaceae 
Heppiaceae 
Hypogymniaceae 
Hemigastraceae 
Hyaloscyphaceae 
Hyalographaceae 
Haliphthoraceae 
Halosphaeriaceae 
Helotiaceae 
Helvellaceae 
Hemiascospor iaceae 
Hymeneliaceae 
Hemisphaeriaceae 
Hyponectriaceae 
Hydnangiaceae 
Hoehnelomyucetaceae 
Hemiphacidiaceae 
Hyphochytriaceae 
Haplosporaceae 
Herpotrichiellaceae 
Hericiaceae 
Herpomycetaceae 
Harpidiaceae 
Hemiascaceae 
Hysterangiaceae 
Heterodeaceae 
Heteroscyphaceae 


HUI 
HUM 
HYA 
HYG 
HYM 
HYP 
HY S 
IMB 
IXE 
JGS 
KAR 
KOE 
KOR 
KRI 
KXL 
LAC 
LAG 
LAH 
LAM 
LAS 
LBL 
LBR 
Oye: 
eer 
LCN 
eer 
LCR 
Eee 
LDM 
EG 
PEG 
EEG 
ed lS 
CEU 
LEV 
LGL 
ogy e 
LIM 
ji a 
Pash 
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MY COTAXON 


Vol. XXXIV, No. 2, pp. 475-487 January-March 1989 


CANALISPORIUM, A NEW GENUS OF 
LIGNICOLOUSTHY PHOMY CETES FROM MALAYSIA 


A. NAWAWI AND A.J. KUTHUBUTHEEN 


Department of Botany, University of Malaya 
59100 Kuala Lumpur, Malaysia 


Canalisporium gen.nov. is erected to accommodate 
Berkleasmium caribense and B.pulchrum. 
Canalisporium elegans sp.nov. is described as the 
third species in the genus. 


Holubova-Jechova & Mercado (1984) described two new 
Species of Berkleasmium Zobel, namely B.caribense Hol.-Jech. & 
Mercado and B.pulchrum Hol.-Jech. & Mercado occurring on dead 
branches from Cuba. Both species produce colonies that are 
sporodochial, punctiform, pulvinate, black and_ shining. Their 
conidia are flattened dorsiventrally, comprising a single layer of 
regularly arranged cells, supported by a small, thinner-walled, 
cuneiform, lighter-coloured basal cell. Conidia of B.caribense 
possess a Single longitudinal septum with 3 — 6 equally spaced 
transverse septa, while in B.pulchrum there are two longitudinal 
septa and 4 — 6 transverse septa. B.caribense has since been 
recorded from Kenya (Kirk, 1985), Uganda and Taiwan 
(Matsushima, 1987). In recording their occurrence both authors 
commented that B.caribense is not. closely’ related to 
Berkleasmium. B.pulchrum has also been recorded from Kenya 
(Kirk, 1985) and India (Rao & Hoog, 1986). 


Recent collections of submerged decaying wood and bark 
from a freshwater stream, subsequently incubated in moist 
chambers have yielded B.caribense, B.pulchrum and an undescribed 
species. Having had the opportunity of examining authentic 
specimens of B.caribense (PRM 831526) and B.pulchrum (PRM 
831528) kindly supplied by Dr. Holubova-Jechova and _ the 
collections of Dr. Kirk from Kenya (B.caribense IMI 284792a and 
B.pulchrum IMI 285428a), Rao from India (B.pulchrum CBS-H 3852) 
and Dr. Matsushima from Taiwan (B.caribense MFC-6T 811), we 
are of the opinion that none of these collections are closely 
related to Berkleasmium and that a new genus is required to 
accommodate them. 


Fig.l. Canalisporium caribense. A, habit sketch; B, stages in 
conidium formation arranged in a developmental series from bark; 
C, conidia in surface and side view; D, two conidia from the 
holotype material (PRM 831526). 


477 


B.concinnum(Berk.) S.Hughes the type species of 
Berkleasmium has well formed stromata and its conidia are 
distinctly cylindrical with numerous cells that are randomly 
arranged and without well spaced transverse and _ longitudinal 
septa. The conidia often lack a basal cell, but instead possess a 
protruding hilum. Developmentally the conidia of B.concinnum 
"originate as blown-out hyphal ends, and reach nearly the size of 
mature conidia before septation is initiated" (Goos, 1969). Conidial 
maturation is by meristematic growth of the entire conidium and 
secession is apparently rhexolytic. These differences merit 
separation at the generic level. 


Canalisporium Nawawi and Kuthubutheen gen.nov. 


Sporodochia dispersa, punctiformia, pulvinata, atra, nitida. 
Mycelium in substrato immersum, ex hyphis ramosis, septatis, 
laevibus, pallide brunneis vel brunneis compositum. Stromata nulla 
vel rudimenta. Conidiophora semi-macronemata vel macronemata, 
mononemata, fasciculata, simplicia vel parce ramosa, subhyalina 
vel pallidissima brunnea, leavia, septata. Cellulae conidiogenae in 
conidiophoris incorporatae, terminales, determinatae. Conidiorum 
secessio schizolytica. Conidia acrogena, solitaria, late ellipsoidea, 
obpyriformia vel subglobosa, complanata, muriformia, 
olivaceo-brunnea, rubrobrunnea vel atrobrunnea, laevia, cum septis 
transversis et longitudinalis, atris, crassis praedita; cellula basalia 
cuneiformia, fere tenuitunicata, subhyalina vel _ pallidissima 
brunnea. Lumina cellularum canaliculis tenuibus, pigmento apposito 
connexa. 


Species typica: Berkleasmium_ caribense’ Hol.-Jech. & 
Mercado. Céska Mykologie 38: 89 (1984). 


Sporodochia scattered, punctiform, pulvinate, granular, black, 
shining. Mycelium mostly immersed in the substratum, composed 
of branched, septate, smooth, pale brown to brown hyphae. 
Stromata none or rudimentary. Conidiophores semi-macronematous 
to macronematous, mononematous, fasciculate, simple to sparsely 
branched, subhyaline to pale brown, smooth, septate. 
Conidiogenous cells integrated, terminal, determinate, cylindrical, 
sometimes swollen. Conidial secession. schizolytic. Conidia 
acrogenous, solitary, broadly ellipsoidal, obpyriform to subglobose, 
flattened, muriform, olivaceous brown, brown, reddish brown to 
dark brown, smooth, with dark, thick-walled transverse and 
longitudinal septa; basal cell cuneiform, thinner-walled, subhyaline 
to light brown. Cell lumen connected by narrow canals each 
surrounded by a marked ring of pigmentation, barrel-shaped in side 
view. 


478 


479 


Canalisporium caribense (Hol.-Jech. & Mercado) Nawawi & 


Kuthubutheen comb.nov. 
Ee Somes 2 


Berkleasmium caribense Hol.-Jech. & Mercado. Céska 
Mykologie 38: 99, 1984 (basionmy). 


Sporodochia on the bark scattered, punctiform, pulvinate, 
granular, black, shining, up to 200 um diam. Mycelium mostly 
submerged in the substratum, composed of irregularly branched, 
septate, smooth subhyaline to pale brown hyphae 1.5 — 2.5 um 
wide. Stromata none or rudimentary consisting of irregularly 
branched, short, intertwined hyphae. Conidiophores 
semi-macronematous to macronematous, mononematous, 
fasciculate, erect to ascending, unbranched to sparsely branched, 
hyaline to subhyaline, smooth up to 20 um long x 2 — 3.5 um wide. 
Conidiogenous cells integrated, terminal, determinate, cylindrical 
or swollen. Conidial secession schizolytic. Conidia acrogenous, 
solitary, flattened, one-cell thick, smooth, thick-walled, broadly 
ellipsoidal to obovoid in surface view, fusiform to obclavate in 
lateral view, muriform, with a single, slightly curved longitudinal 
septum and from 3 —- 6 straight transverse septa, slightly 
constricted at the septa, evenly brown to reddish dark brown, 28 — 
41x 21 — 28 wide x 10 — 14 um. Basal cell subhyaline to pale 
Drownwecunelorm, with, thinner’ wall, 5 — 5.5. long x 3°- 3.5 um 
wide. Septa becoming progressively darker with conidial maturity. 
The left and right cell lumen are connected by narrow canals, 
each surrounded by a marked ring of pigmentation, visible in 
surface view as a circular disc 2 — 2.5 um diam, barrel-shaped in 
side view. Canals are also present in the transverse septa, but 
only on one side of the conidium adjacent to the curved side of 
the longitudinal septum. 


Specimen examined: On submerged decaying bark, Gombak 
Brelgeestudy., Centre, Selangor, Malaysia, 12 Sept.,, 1987, A. 
Nawawi, IMI 326601. 


Fig.2. A -— B, Canalisporium caribense. A, three stages in 
conidium formation and maturation from bark; B, mature conidia. 

C — J, Canalisporium pulchrum. C, several developing 
conidia enveloped in mucilage; D, three conidia in surface view; E, 
two conidia in side view with the connecting pores clearly visible; 
F, part of a conidium showing pores along the longitudinal septa; 
G, a bleached conidium; H — I, conidia from the holotype material, 
one bleached (PRM 831528); J, two conidia from the Indian 
collection (CBS-H 3852). 


480 


Fig.3. Canalisporium pulchrum. A, habit sketch; B, stages in 
conidium formation from bark; C — C, conidia in surface and side 
view; E, two conidia from the holotype material (PRM 831528); F, 
a conidium from the Indian collection (CBS-H 3852). 


481 


Conidia germinate very slowly on agar media by the 
production of a germ tube from the basal cell. On CMA linear 
extension of the hyphae is limited, and colonies from a single 
conidium, after 4 weeks at 25 — 27°C consist of a compact mass 
of hyphae rarely exceeding 8 mm in diam. The colonies are at 
first hyaline, progressively turning grey and finally black with 
most of the hyphae becoming moniloid. Sporulation was sparse on 
CMA, but occurred profusely on strips of sterilised banana leaves 
overlying CMA. Sporodochia are: similar to those formed on the 
bark, but tend to coalesce to form an extensive mass of conidia. 


Conidial development from bark and on banana leaves is 
similar. Conidia originate as a blown-out hyphal ends, at first 
hyaline and club-shaped. Cross septa develop quite early followed 
by the formation of longitudinal septa. Development of 
pigmentation occurs soon after all the septa have been formed and 
becomes more intense as the conidia age. 


Comparative studies made on the holotype material from 
Cuba and those collected by Kirk from Kenya with the Malaysian 
isolate show that there is no fundamental difference in conidial 
morphology, size and in the number of cells making up a conidium. 
Conidia from the holotype material, however, appear to be slightly 
reddish brown with less pigmentation and hence the connective 
pores are readily visible. 


Canalisporium pulchrum (Hol.-Jech. & Mercado) Nawawi & 
Kuthubutheen comb.nov. 
FigsieZy-s 


Berkleasmium pulchrum  Hol.-Jech. & Mercado. Céska 
Mykologie 38: 96 — 120. 1984 (basionym) 


Sporodochia scattered, punctiform, pulvinate, granular, black, 
shining, circular to ellipsoidal, up to 300 um diam. Mycelium 
scanty, mostly immersed in the substratum composed of branched, 
smooth, septate, pale olivaceous brown to brown hyphae 2 — 3.5 
um wide. Stromata absent or at most rudimentary, consisting of 
much-branched intertwined hyphae. Conidiophores 
semi-macronematous, mononematous, fasciculate, densely 
aggregated, simple or sparsely branched, short, septate, pale 
brown, smooth, up to 20 um high, 2.5 — 4 um wide. Conidiogenous 
cells integrated, terminal, determinate. Conidial secession 
schizolytic. Conidia arising as terminal blown out ends of 
conidiogenous cells, initially club-shaped and _ hyaline, each 
surrounded by a thick layer of mucilage. Transverse septa formed 
early in development, followed by longitudinal septa, at first 
thin-walled, progressively becoming thicker and darker and finally 


482 


heavily pigmented. Conidia acrogenous, solitary, broadly ellipsoidal 
in surface view, clavate in lateral view, muriform, one-cell thick, 
smooth, evenly olivaceous brown to brown, slightly constricted at 
the. septa, 36 —- 55 xX422 — 927 x12") =) 15) [iiss 
thinner-walled, lighter, cuneiform, 5 — 6 um long x 4 — 5 um wide. 
Longitudinal septa with narrow, barrel-shaped pores, 1.5 — 2 um 
diam connecting each cell lumen laterally; transverse septa with 
pores only in the middle column of cells. 


Specimen examined. On rotten wood, Gombak Field Study 
Centre, Selangor, Malaysia 20 Oct. 1987, A. Nawawi, IMI 326602. 


Conidia germinate slowly on agar by producing a germ tube 
from the basal cell. On CMA the colonies are dark, restricted, 
with little aerial mycelium, attaining a diam of 10 mm in 4 weeks 
at- 25 —' 27°C. Sporulation was ‘Sparse..on agaraybutmocemnnes 
profusely on strips of sterilised banana leaves overlying CMA. 
Conidia formed on agar are slightly smaller but otherwise 
resemble those formed on the natural substratum. 


Comparative studies on conidial morphology, size, colour, 
septation and the number of cells making up a conidium between 
the holotype material from Cuba and those collected from India 
and Malaysia are listed in Table l. 


Table 1. Comparison of Canalisporium pulchrum 
from three localities 


Locality Conidial size No. of No. of cells/ No. of cells 
(um) transverse conidium in apical row 
septa 
Cuba 27-43x23-27x12-15 3-6 ; 13-21 1-3 
(36x25)* 
India 28-50x20-29x15-17 3-6 11-20 mostly 1 
(41-25) 
Malaysia 36-52x22-27x12-15 4-7 14-24 1-3 
(44x24) 


* Figures in brackets are average length x width. 


While there is no_ significant difference in conidial 
morphology and width between the 3 collections, their lengths 
vary slightly. The Malaysian collection has longer conidia, less 
pigmented septa, and contains more cells per conidium. In other 


483 


Fig.4. Canalisporium elegans. A, habit sketch; B, stages in 
conidium formation from bark; C, conidia in surface and side 
view. 


484 


aspects it is very similar to the holotype material. Conidia of the 
Indian collection are darker, heavily pigmented and the majority 
possess only one cell in the apical row. 


Collection from Kenya (IMI 285428a) Figa:> 


The collection from Kenya which Kirk (1985) identified as 
Berkleasmium pulchrum appears to be quite different. The conidia 
are broadly obpyriform and significantly larger, measuring 34 — 56 
x 27 — 34 x 15 — 18 um (avg. 48 x 30 um), reddish brown to brown 
with 5 — 7 transverse septa that are heavily pigmented. The 
number of cells per conidium varies from 17 — 19. The most 
obvious difference is the absence of a basal cell. Instead, a row of 
three, smaller, subhyaline cells is present, joined to the 
conidiophore and conidial secession appears to be rhexolytic. These 
differences have been pointed out by Rao & Hoog (1986). A 
further study of conidiogenesis is necessary before any 
reclassification is proposed. 


Canalisporium elegans Nawawi & Kuthubutheen sp.nov. Figs. 4 — 5 


Sporodochia dispersa, punctiformia, atra, nitida, usque ad 250 
um diam. Mycelium in substrato immersum, ex hyphis ramosis, 
septatis, laevibus, pallide brunneis, 2 — 3.5 um latis compositum. 
Stromata nulla. Conidiophora semi-macronemata, mononemata, 
fasciculata, simplicia vel parce ramosa, subhyalina vel pallidissima 
brunnea, laevia, septata, usque ad 15 um longa x 3 — 4 um crassa. 
Cellulae conidiogenae in conidiophoris incorporatae, terminales. 
Conidiorum  secessio schizolytica. Conidia acrogena, solitaria, 
subglobosa, complanata, muriformia, olivacea brunnea vel brunnea, 
laevia, 32. —°947 x27 — 35 x, 10 — 13) um, Cul 
transversis et 4 (-5) septis longitudinalibus, atris, crassis praedita; 
cellula basali cuneiformia, fere tenuitunicata, subhyalina, 5— 6 um 
longa x 4 — 5 um crassa. Lumina cellularum canaliculis tenuibus, 
pigmento apposito connexa, 1.5 — 2 um diam. 


Ex ramo putrido deiecto trunci ignoti, Gombak Field Study 
Centre, Selangor, Malaysia 20 oct. 1987, A. Nawawi, IMI 326603, 
holotypus. 


Sporodochia scattered, punctiform, pulvinate, granular, black, 
shining, up to 250 um in diam. Mycelium mostly immersed in the 
substratum, composed of branched, septate smooth, pale brown, 2 
- 3.5 um wide hyphae. Stromata none. Conidiophores 
semi-macronematous, mononematous, fasciculate, simple to 
sparsely branched, subhyaline to pale brown, up to 15 um long x 3 
— 4 um wide. Conidiogenous cells integrated, terminal, 


485 


C 


Fig.5. A, four conidia from the Kenyan collection (IMI 
285428a); B — C, Canalisporium elegans. Mature conidia from 
bark. 


486 


determinate. Conidial secession schizolytic. Conidia acrogenous, 
solitary, subglobose, flattened, one-cell thick, muriform, smooth, 
olivaceous brown to brown, 32 — 47 x 27 — 35 x 10 — 13 um with 
dark, thick-walled 5 — 7 transverse septa and 4 (-5) longitudinal 
septa; basal cell cuneiform, 4 — 6 tim ‘long: xX) 4 aoe iimeioes 
thinner-walled, subhyaline. The number of cells per conidium 
varies , from 26 = °35,**‘apical rowa with |1 —%5 \celisme@elietanmne 
connected by narrow canals, 1.5 — 2 um diam, each surrounded by 
a marked ring of pigmentation. Along the transverse septa, Canals 
are only present in the middle column of cells. 


Conidia germinate slowly on agar by producing a germ tube 
from the basal ceil. Colonies are slow growing, restricted, at first 
grey but eventually turning dark, attaining a diam of 12 mm in 4 
weeks at 25 — 27°C. Sporulation was poor on agar but occurred 
profusely on strips of sterilised banana leaf overlying CMA. 
Conidia formed in cultures are slightly smaller, many with only 3 
longitudinal septa. 


The three species of Canalisporium described above have 
several characters in common. They are all lignicolous with black, 
punctiform, non-stromatic  sporodochia. Conidiophores and 
conidiogenesis are similar. Their conidia are  muriform, 
complanate, generally brown with transverse and longitudinal septa 
that are highly pigmented. The basal cell is distin@t@ancams 
delimited from the spore body in pigmentation, size and thickness 
of cell wall. The pore-like connectives of conidial lumina are 
remarkable in all three species. Conidial germination is by a basal 
germ-tube and their colonies are dark, slow growing and 
eka (egere 


ACKNOWLEDGEMENTS 


The authors are much obliged to Professor J. Webster for 
kindly reviewing the manuscript and to Drs. V. Holubova-Jechova, 
G.S.de Hoog, P.M. Kirk, T. Matsushima and the herbarium of CMI 
for the loan of collections mentioned in the text. 


REFERENCES 


Goos, R.D. (1969). Berkleasmium concinum in culture. Canadian 
Journal of Botany &7:503°— 505. 

Holubova-Jechova, V. & Mercado, S.A. (1984). Studies on 
hyphomycetes from Cuba II. Hyphomycetes from the Isle of 
Juventud. Céska Mykologie 38:96 — 120. 

Kirk, P.M.(1985). New or interesting microfungi XIV. 
Dematiaceous hyphomycetes from Mt. Kenya. Mycotaxon 
fie Sie 10 eon Gaye 


487 


Matsushima, T. (1987). Matsushima Mycological Memoirs 5:1 — 
100. Published by the author, Kobe, Japan. 

Rao, V. & Hoog, G.S. de (1986). New or critical hyphomycetes 
from India. Studies in Mycology 28:1 — 84. 


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MYCOTAXON 


Vol. XXXIV, No. 2, pp. 489-495 January-March 1989 


QUADRICLADIUM AQUATICUM GEN. ET SP.NOV., 
AN AQUATIC HYPHOMYCETE WITH 
TETRARADIATE CONIDIA 


A. NAWAWI and A.J. KUTHUBUTHEEN 


Department of Botany, University of Malaya 
59100 Kuala Lumpur, Malaysia 


Quadricladium aquaticum gen. et sp.nov., an aquatic 
hyphomycete with hyaline, tetraradiate conidia from 
submerged decaying bark is described and illustrated. 
The generic characters are briefly compared with 
those of somewhat similar genera: Lemonniera, 
Tetraposporium, Brachiosphaera and Flabellospora. 


The existence of large, hyaline, pearly, tetraradiate conidia 
in foam samples from fresh streams in Malaysia has been known 
to the senior author since 1975, but the identity of the fungus 
forming these conidia remained undetermined, as it has never been 
detected on its natural substrata and details of conidiogenesis 
were unknown. Repeated attempts to germinate and establish the 
fungus in pure culture were unsuccessful. A recent collection of 
submerged decaying bark pieces from a fresh water stream has 
yielded the fungus in question. Having now successfully observed 
the fungus developing on its natural substrate and conidial 
development in pure culture, we are of the opinion that it does 
NOt sit “into any known taxa. A new genus is erected to 
accommodate it. 


Quadricladium gen.nov. 


Coloniae effusae, hyalinae, inconspicuae. Mycelium 
plerumque superficiale, ex hyphis laevibus, ramosis, septatis, 
hyalinis compositum. Conidiophora macronemata, mononemata, 
solitaria, erecta, simplicia, recta vel leviter flexuosa, laevia, 
septata, hyalina. Cellulae conidiogenae in_ conidiophoris 
incorporatae, terminales. Proliferatio holoblastica et sympodialis. 
Conidia acrogena, solitaria, hyalina, tetraradiata, septata; axis 
principalis cum conidiophoris continuis; ramis lateralis 3 — 4, simul 
gemmantia. Conidiorum secessio schizolytica. 


490 


Species typica Quadricladium aquaticum sp.nov. 


Colonies effuse, hyaline, inconspicuous. Mycelium mostly 
superficial, composed of branched, smooth, septate, hyaline 
hyphae. Conidiophores macronematous, mononematous, solitary, 
erect, simple, straight to slightly flexuous to slightly flexuous, 
smooth, septate, hyaline. Condiogenous cells incorporated in the 
conidiophores, terminal. Proliferation holoblastic and sympodial. 
Conidia acrogenous, solitary, hyaline, septate, tetraradiate; 
principal arm continuous with the conidiophores; lateral arms 3 — 
4, produced simultaneously. Conidial secession schizolytic. 


Quadricladium aquaticum sp.nov. Rigs glee 

Coloniae effusae, hyalinae, inconspicuae. Mycelium 
plerumque superficiale, ex hyphis laevibus, ramosis, septatis, 
hyalina, 1.5 — 2:5 um  latis compositUm as COnimiGn ors 
macronemata, mononemata, solitaria, erecta, recta vel leviter 
flexuosa, interdum, leviter geniculata, laevia,) l= oeeseprara. 


hyalina, 16 — 58 um alta x 3— 4 um lata. Cellulae conidiogenae in 
conidiophoris incorporatae, subglobosa vel ellipsoidea, 8 — 10 um x 
4.5 — 6 um, terminales. Proliferatio holoblastica et sympodialis. 
Conidia acrogena, solitaria, hyalina, tetraradiata; parte centralis 
subglobosa 9 — 12 um in diam cum conidiophoris connexa, Cum uno 
axe et 3 — (4) ramis divergentibus, simul gemmantia. Rami et axis 
leviter constricti ad basim, 41 — 67 wm longi x 6.5 —*/2Omiieee 
— 6 septati. Conidiorum secessio schizolytica. 


Habitat saprophytice in cortice decidui trunci in flumine 
immersi, Mimaland, Selangor, Malaysia, 27 Julii, 1988, A. Nawawi, 
IMI 327415 holotypus. 


This fungus was found growing and sporulating on pieces of 
decaying bark taken from a submerged log of an unidentified tree 
in a small fresh water stream at Mimaland, Selangor. On 
incubation at 25 — 27°C in water this fungus sporulated for several 
days. Conidiophores bearing conidia were formed under water and 
also on the moist exposed parts of the substratum. 


Fig. 1. Quadricladium aquaticum. A —- B, Stages in conidium 
formation on bark arranged in a developmental series; C — E, 
conidiophores showing detachment scars and proliferation; F, three 
typical conidia, one viewed from below; G, part of a conidium 
showing germ tubes. 


492 


Colonies effuse, hyaline, inconspicuous. Mycelium mostly 
superficial, composed of smooth, branched, septate, hyaline hyphae 
1.5 — 2.5 um wide. Conidiophores macronematous, mononematous, 
solitary, simple, erect, straight to slightly flexuous, sometimes 
geniculate, 16.— 58 um long x 3 -— 4 pm wide, “SmOOtTiye ian? 
septate, hyaline. Conidiogenous cells integrated, terminal, broadly 
ellipsoidal to ovoid, 8 — 10 pm long x 4.5 — 6 um wide. 
Proliferation holoblastic and sympodial. Conidial development on 
the bark begins with the formation of a small spherical swelling at 
the apex of the conidiogenous cell. This swelling is soon delimited 
by a septum, elongates slightly, becomes tetrahedral and 2-celled. 
The four arms of the conidium develop simultaneously from the 
corners of the tetrahedron, one arm always distal and continuous 
with the conidiophores, the others radiating outwardly forming an 
angle of 100 — 120° with the distal arm. The distal arm elongates 
slightly faster than the others and is usually longer in mature 
conidia. Septa are formed as the arms elongate. When a conidium 
reaches maturity, another conidiogenous cell develops very close 
to the attachment point and soon produces another conidium. This 
process of sympodial proliferation is repeated 3 — 4 times. 
Conidial secession is schizolytic. The conidia, if undisturbed, often 
remain attached to their respective conidiogenous cells. A small, 
circular, collarette-like scar is clearly visible on the conidiogenous 
cell after each conidium is released. Mature detached conidia are 
tetraradiate, occasionally pentaradiate hyaline, pearly, with a 
globose to pentagonal central cell 9 — 12 um in diam; attachment 
scar distinct in side view as a conicotruncate protuberance 2 um 
wide. Arms straight, cylindrical, tapering only slightly, 41 — 67 um 
long x 6.5 — 7.5 um at the widest and 3 um at the rounded apex, 
narrowing slightly at the base. Septa thin-walled, but distinct, up 
to 6 per arm. 


Conidia germinate with difficulty. On CMA _ germination 
occurred after 4 days at 25 — 27°C. Germ tubes are produced 
from the apical cells or anywhere along the arms including the 
central cell. Colonies are slow growing, white, ciruclar with a 
slight mound, attaining a diam of 10 mm in 30 days; reverse 
concolorous. Aerial mycelium compact, hyphae hyaline 1.5 — 2.5 
lum wide, occasionally forming spherical structures 3 — 4 um diam 
terminally or intercalarily. No sporulation occurred on dry agar 
but when strips of the colony were suspended in water conidia 
were formed abundantly. The pattern of conidial formation is 


Fig. 2. Quadricladium aquaticum. A, States in conidium formation 
from bark; B, conidiophores with spent conidiogenous cells; C — D, 
typical conidia. Bars = 20 um. 


493 


4 


494 


similar to those formed on the natural substratum. Conidia, 
however, tend to be slightly smaller in size. 


Conidia of Q.aquaticum are very similar to those of the 
aquatic hyphomycete Lemonniera de Wild., particularly L.terrestris 
Tubaki. However, members of Lemonniera produce their conidia on 
distinct phialides borne on long, often branched conidiophores and 
are widespread in temperate countries (Descals, Webster & Dyko, 
1977). The anamorph genus is apparently not represented in 
Malaysia (Nawawi, 1985). 


Another genus which produces conidia with similar conidial 
morphology and _ configuration to the present fungus is 
Tetraposporium Hughes, but members of this genus are 
dematiaceous and are hyperparasites (Hughes, 1951). Their conidia 
are borne on very. short, determinate conidiophores, often 
undifferentiated from the vegetative hyphae. 


In terms of conidiogenesis, shape and proliferation of the 
conidiogenous cells, Q.aquaticum is similar to Brachiosphaera 
tropicalis Nawawi, a common aquatic hyphomycete occurring on 
submerged decaying branches and twigs. Its conidia are abundant 
in foam samples from Malaysian fresh water streams and widely 
distributed in the tropics (Descals, Nawawi & Webster, 1976). 
However, conidia of B.tropicalis possess a large, spherical central 
cell from which the 4 septate arms radiate out. Q.aquaticum has 
also been considered for inclusion in Flabellospora Alasoadura. The 
type species F.crassa Alasoadura (1968), a rather common aquatic 
hyphomycete in Malaysia, has hyaline, septate, tetraradiate 
conidia with a globose, pedunculate central axis from which the 
obclavate arms radiate out simultaneously. Conidia are borne on 
long, undifferentiated, determinate conidiophores with drooping 
conidiogenous cells. The arms of the conidia are constricted at 
their points of origin and conidial secession is by a break in the 
constriction along the drooping peduncle. 


A leaf-litter hyphomycete described by Matsushima (1981) 
from Alabama, U.S.A. as Flabellospora irregularis Matsushima 
produces conidia with 2 — 4 arms radiating simultaneously from a 
globose, central cell. Conidia are produced on short conidiophores 
and the ovoid conidiogenous cells proliferate sympodially, very 
similar to Q.aquaticum. In recording its occurrence the author 
commented that F.irregularis deviates somewhat from _ other 
members of the genus in its conidial morphology. This fungus has 
also been recorded from Taiwan (Matsushima, 1983), and is 
perhaps congeneric with Q.aquaticum. 


495 


Professor J. Webster of Exeter University continues to be 
the subject of our thanks for his prepublication editorial vetting of 
this manuscript. 


REPERENGES 


Alasoadura, S.O. (1968). Flabellospora crassa n.gen., n.$p., an 
aquatic hyphomycete from Nigeria. Nova Hegwigia 15: 415 — 
418. 

Descals, E.. Nawawi, A. & Webster, J. (1976). Developmental 
studies in  Actinospora and _ three similar aquatic 
hyphomycetes. Transactions of the British Mycological 
mocieLveo/:.20/ — 222° 

Descals, E., Webster, J. & Dyko, B.J. (1977). Taxonomic studies on 
aquatic hyphomycetes. I. Lemonniera de _  Wildeman. 
Transactions of the British Mycological Society 69: 89 — 109. 

Hughes, S.J.(1951). Studies on microfungi. XII. Triposporium, 
Tripospermum, Ceratosporella, and Tetraposporium (gen.nov.). 
Mycological Papers C.M.I. 46: 1 — 3). 

Matsushima, T.(1981). Matsushima Mycological Memoirs 2: | — 68. 
Published by the author, Kobe, Japan. 

Matsushima, T. (1983). Matsushima Mycological Memoirs 3: | — 8&9. 
Published by the author, Kobe, Japan. 

Nawawi, A. (1985). Aquatic hyphomycetes and other water-borne 
fungi from Malaysia. Malayan Nature Journal 39: 75 — 134. 


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MY COTAXON 


Vol. XXXIV, No. 2, pp. 497-501 January-March 1989 


A NEW TAXON IN COLISPORA (HYPHOMYCETES) 
FROM MALAYSIA 


A. NAWAWI and A.J. KUTHUBUTHEEN 


Department of Botany, University of Malaya, 
59100 Kuala Lumpur, Malaysia. 


Colispora curvata sp.nov., an aquatic hyphomycete 
from submerged decaying petioles is described and 
illustrated. 


Staurosporous species are the classic representatives of 
Ingoldian fungi, but scolecosporous forms are also present. A 
number of didymo- and phragmosporous aquatic species have been 
described, all assigned to characteristically terrestrial 
anamorph-genera. Two such examples are Pyricularia aquatica 
Ingold (1943) with a Massarina teleomorphic stage (Webster, 1965) 
and P.submersa Ingold (1944). Both were transferred to Dactylella 
Grove, respectively as D.aquatica (Ingold) Ranzoni (1953) and 
D.submersa (Ingold) Nilsson (1962). Their inclusion in either 
Pyricularia or Dactylella is still debatable. 


An aquatic hyphomycete resembling  P.submersa in 
conidiogenesis and spore morphology, found on _ pieces of 
submerged decaying petioles from a freshwater stream at the 
University of Malaya Field Study Centre in 1976 was left 
undescribed. The senior author was not convinced that it should be 
placed in either Pyricularia or Dactylella. Recently Marvanova 
(1988) erected a new genus of aquatic hyphomycete with a single 
species, Colispora elongata Marvanova, isolated from foam in 
Czechoslovakia with similar characteristics as our undescribed 
fungus, and this is described below as a new species in the genus. 


Colispora curvata sp.nov. AB ESE 


Coloniae effusae, hyalinae, inconspicuae. Mycelium partim 
superficiale, partim in substrato immersum, ex hyphis laevibus, 
ramosis, septatis, hyalinis, 1.5 — 2.5 um _ latis compositum. 
Conidiophora micronematosa vel semi-macronematosa, 
mononemata, solitaria, simplicia, recta, laevia, septata, hyalina, 
Beguemad 18) um. longa, 2.5 — 3.2. um jata, per usque ad .20 
proliferationes percurrentes crescentia. Cellulae conidiogenae in 


498 


conidiophoris incorporatae, terminales, percurrentes. Conidiorum 
secessio schizolytica. Proliferatio enteroblastica et percurrens 
progressiv et annulos’ fingens. Conidia acrogena,  solitaria, 
reniformia, guttulata, hyalina, laevia, 2 — 3 septata, ad apice 
rotundata, basi conico-truncata, excentrica, 25 — 38 (-59) um 
longa, 7.5 — 10 um lata, 3 — 4 ad basim. 


Ex petiolis angiospermis putrescentibus in flumine immersi, 
University of Malaya Field Study Centre, Gombak, Selangor, 
Malaysia, 3 Aug. 1976, A. Nawawi, IMI 327693 holotypus. 


Colonies on the petioles effuse, hyaline, inconspicuous. 
Mycelium partly superficial, partly immersed in the substratum, 
composed of smooth, branched, septate, hyaline hyphae 1.5 — 2.5 
um wide. Conidiophores micronematous to semi-macronematous, 
mononematous, solitary, simple, straight, smooth, septate, hyaline, 
up to 185 wm long, 2.5 — 3.5 wm wide, with up to) 2Z0s0rsmore 
percurrent proliferations. Conidiogenous cells integrated, terminal, 
percurrent. Conidial secession schizolytic. Proliferation 
enteroblastic, percurrent and progressive, leading to the formation 
of annellations. Conidia acrogenous, solitary, reniform, guttulate, 
2 -— .3 septate, rounded at the apex with “aneveccenmic 
conico-truncate base, hyaline, smooth, rich in glycogen, pearly, 25 
— 38 (-59) um long, 7.5 — 10 um wide, 3 — 4 um at the base. 


The course of an individual conidium development on the 
natural substrate was followed in a constant flow chamber 
containing a small piece of the colonised petiole as described by 
Descals, Nawawi & Webster (1976). The conidium starts as a 
small, oval swelling at the end of the conidiophore. It elongates, 
swells considerably, becomes slightly curved and is cut off by a 
cross wall. A septum soon develops near the middle. The rounded 
apex continues to elongate and another septum develops. In the 
meantime several guttules of various sizes appear, rendering the 
conidium pearly and foamy. It takes about 7 — 8 hr for a conidium 
to mature and become detached. As soon as a conidium is 
released, the truncate conidiophore tip resumes growth, proceeds 
for 5 — 9 um before producing the next conidium at a higher level. 
This process is repeated many times and results in a conidiophore 
of up to 480 um long. Annellations on the conidiophore are not 
always prominent as they are thin, and colourless. They appear as 
a faint, transverse band across the broadest part of the beaded 
portion along the conidiophore. In old conidiophores a series of 
these annellations occurs, numbering up to 20. 


Conidia germinate soon after their release. On 2% MA the 
colony is at first white, turning greyish with several concentric 


499 


A// B G D 


Fig.l. Colispora curvata. A — E, a conidiophore drawn at intervals 
over a period of 10 hr in a constant flow chamber showing 
the manner of conidium development. The conidiophore had 
previously produced a conidium at the point A; F, the same 
conidiophore 48 hr later; G, two conidiophores with developing 
conidia; H, typical conidia from petiole. 


Fig.2. Colispora curvata. A — D, a conidiophore showing four 
stages in conidium formation. F — G, conidia and germination. 


zones; aerial mycelium fluffy, reverse concolorous, attaining a 
diam of 22 mm in 14 days at 25 — 28°C. Sporulation was sparse on 
dry agar, but occurred profusely on strips of agar submerged in 
sterile water. It was observed that aeration is essential for normal 
spore formation. Unaerated cultures tend to produce conidia that 
are longer, many with 3 septa. 


C.curvata is essentially similar to C.elongata and P.submersa 
in conidiogenesis and proliferation of their conidiophores and in 


501 


having hyaline, septate, pearly conidia. Conidia of C.elongata and 
P.submersa however, are longer, fusiod to clavate mostly straight 
and with more septa. Colonies on 2% MA were described as 
dark-grey and greenish-grey respectively. 


We are grateful to Professor J. Webster, University of 
Exeter, for kindly reviewing the manuscript and for helpful 
comments. 


REFERENCES 


Descals, E., Nawawi, A. & Webster, J. (1976). Developmental 
studies in  Actinospora and _ three’ similar aquatic 
hyphomycetes. Trans. Br. Mycol. Soc. 67:207 — 222. 

Ingold, C.T. (1943). Further observations on aquatic hyphomycetes 
on decaying leaves. Trans. Br. Mycol. Soc. 26:104 — 115. 
Ingold, C.T. (1944). Some new aquatic hyphomycetes. Trans. Br. 

Mycol. Soc. 27:35 — 47. 


Marvanova, L. (1988). New hyphomycetes from aquatic 
environments in Czechoslovakia. Trans. Br. Mycol. Soc. 
90:607 — 617. 

Nilsson, S. (1962). Second note on Swedish freshwater 


hyphomycetes. Bot. Not. 115:73 — 86. 

Ranzoni, F.V. (1953). The aquatic hyphomycetes of California. 
Farlowia 4:353 — 

Webster, J. (1965). The perfect stage of Pyricularia aquatica. 
Trans. Br. Mycol. Soc. 48:449 — 452. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 503-505 January-March 1989 


ASCOMYCETES OF WESTERN INDIA XIII 


ALAKA PANDE AND V.G. RAO 
Maharashtra Association for the Cultivation of Science 
Law College Road, Pune 411004, INDIA 


The paper communicates 2 new species of bitunicate 


Ascomycetes viz. Didymella capparidis sp.nov. 


and Julella multiloculata sp.nov. 


Didymella capparidis sp.nov. (Fig. 1-4) 


Stroma immersed in the bark, appearing as shining black areas 
through split bark, upto 5 mm long, 2-3 mm wide and 100-200 yam 
deep. Ascocarps separate or aggregated, immersed in stroma, globose 
to subglobose, slightly papillate, 110-180 x 160-200 yim. Wall composed 
of thick-walled polygonal, brown cells. Asci in basal layers, surro- 
unded by pseudoparaphyses, bitunicate, clavate, shortly stalked or 
mearly sessile, 60-90 x 14-18 ym. Ascospores hyaline, 1-septate, 
elliptical, tips rounded, constricted at septum; 27-32 x 6-10 pm. 


Collected on dead twigs of Capparis horida L.f. at Pune (Maha- 
rashtra) 9.X.1981. AMH 7153 (Holotype). 


Stromata anthracina, immersa in corticem 5 mm longa, 2-3 mm 
lata, 100-200 um crassa, exposita per corticem. Ascomata depressa, 
immersa, numerosa, ovoidea-globosa, leviter papillata, magnit. 110- 
180 x 160-200 pm. Paries compositae ex cellulae polygonae, atro- 
brunneae. Asci basalis, paraphysoidibus, bitunicati, clavati, brevi- 
stipitati, 60-90 x 14-18 pm. Ascosporae hyalinae, ellipticae, apicem 
rotundato, uniseptatae, constrictae ad septum, 27-32 x 6-10 ym. 


Ad caulibus mortuis Capparis horida L.f. dt. 9.X.1981. Loco: 
Pune. (Maharashtra). AMH 7153 (Holotypus). 


There is only one stromatic species under the genus Didymella 
viz. D. stromatica Barr & Rogerson (1982). However, the ascomata, 
asci and ascospores in that species are much smaller and also the 
ascospores in D. stromatica are obovoid with obtuse apical end and 
more acute basal tip, whereas in the present collection they are 
elliptical and rounded at both ends. It is described here as a new 
species, the specific epithet being derived from the host genus. 


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Julella multiloculata sp.nov. (Figs. 5,6) 


Stromata innate, exposed through bark, elongated, black multi- 
locular, 600-900 x 300-400 pm. Locules many, embedded in stroma, 
arranged in a row, globose; each locule measuring 200-320 x 200- 
360 pm; wall composed of flattened cells on inner side and irregular 
to globose cells on outer side. Asci many, in basal layers, bituni- 
cate, cylindrical to club shaped, stalked to subsessile, octosporous, 
paraphysoids present, asci 120-180 x 17-25 pom. Ascospores 8, muriform, 
golden yellow to light brown 30-45 x 14-16 pm. 


Growing on twigs of Punica granatum L. at Pune (Maharashtra) 
dt. 3.X.1981. AMH 7150 (Holotype). 


Stromata immersa in corticem, multiloculata, nigra, erumpentia 
per cortice, exposita pro elongata rima, 600-900 x 300-400 pm. loculi 
pluri, innati in stroma, lineati dispositi, globosi, paries compositae 
ex cellulae hyalineae, compressae; loculi magnit. 200-320 x 200-360 
pm. Asci numerosi in basalis strato, bitunicati, cylindrici vel clavati, 
stipitati vel subsessili, octospori, paraphysoidibus, 120-180 x 17- 
25 pm. Ascosporae 8, muriformae, aureae vel pallide brunneae, 30- 
45 x 14-16 pm. 


Ad caulibus mortuis Punica granavume OU. dt.) 13.X.1981...7 Pune 
(Maharashtra). AMH 7150 (Holotypus). 


This fungus is closest to Julella (Barr, Personal communication, 
Barr 1985). However, presence of multiloculate stromatic fruit bodies 
and 8 lightly pigmented ascospores per mature ascus distinguish 
it from other species of Julella which normally have uniloculate asco- 
mata and usually less than 8 hyaline ascospores per ascus at maturity. 
The species is named to indicate its multiloculate nature. 


ACKNOWLEDGEMENTS 


Grateful thanks are offered to Dr.P.G.Patwardhan, The Director 
M.A.C.S. Research Institute, Pune for facilities. 


REFERENCES 
foeeaarr, IM.B. 1985. On Julella, Delacourea and Decaisnella, three 
dictyosporous genera described by J.F.Fabre. Sydowia 38 


glea hee 
2. Barr, M.E. & C.T. Rogerson 1982. Two new species of Loculo- 
ascomycetes. Mycotaxon 17 : 247-252. 


Didymella capparidis sp.nov. (Figs.1-4) 1. stroma (diagramatic), 
2. locule, 3. ascus, 4. ascospores. Julella multiloculata sp.nov. (Figs. 
5,6) 5. stroma, 6. ascospores. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 507-515 January-March 1989 


THE GENUS CHAETOMASTIA (DACAMPIACEAE) IN NORTH AMERICA 


MARGARET E. BARR 


Department of Botany, University of Massachusetts, Amherst, 
MA 01003 


ABSTRACT 


Chaetomastia is reinstated as an older name for some 


species recently assigned to Massariosphaeria. a€ Venn Oth 
American species are described and illustrated, including 
the type species C. hirtula. New combinations are proposed 


Peeeeweclavispora, C.' phaeospora, 'C. sambuci,.C. sambucina, 
C. typhicola, and the new species C. equiseti is described. 


The name Chaetomastia, Pins te propocedurby \ sa eccardo 
(1883) as subgenus III of Melanomma "Perithecia setulosa", 
has scarcely been used since Berlese (1891) raised it to 


generic rank. twowepeciles Mane: Slisted in they checklist “to 
eee eeneAsScomycotina (Cannon et al. 1985) and the genus is 
Poeomaecd | in "“Unitunicate -Ascomycetes, inc. sed." by 0. 
Eriksson and Hawksworth (1987). Saccardo listed Melanomma 
hirtulum thar s-Cjen') Sia CC wh pilosellum Karsten, Ms 
canescens SiC ts Me hispidulum Sac Ors ira ien | Peake rg 
cucurbitarioides Speg. within the subgenus. Berlese (1891) 
transferred these taxa to Chaetomastia, but was unable to 
Study C. cucurbitarioides. He added C. jfuniperina (Karsten) 
Berlese and:'C. aculeata (Mouton) Berlese. He excluded M. 
(GO: ) plejosporum Mouton with muriform ascospores’ and 
Seuseested*+ that) it belonged) in: Capronia. Chaetomastia 


pilosella is Herpotrichiella pilosella (Karsten) Munk (Munk 
Pee. “shispidula and) C. aculeata probably belong! in 
HWerpetrichiella also. The disposition, of “other species 
meoepe tor C. hirtula is uncertain to me’at present. 


meementsy and, Shear (1931!) fdesienated “Ci hirtula fut 
Seaeeten) |) Sacc.)]) as lectotype species of ‘the genus and 
submerged it under Lasiosphaeris Clements (=Lasiosphaeria). 
Meee ypification was‘acceptediin ING (Farr et al. '1979) and 


feeerLomastia is thus based) upon C.° Hirtula, This species 
was described in some detail as Melanomma hirtulum by Holm 
e2as), “who recognized ‘that disposition out of Melanomma 
would be necessary. Leuchtimann —CLO85)> otransterred. “the 
eapecies to the expanded Montagnula, which includes 
phragmosporous species (Crivelli 1983), because of the dark 
reddish brown, asymmetric, thick-walled, verruculose 


ascospores and the presence of abundant hyphae around 


508 


ascomata. Montagnula, according to the type spectesnaa. 
infernalis (Niessl) Berlese has abundant hyphae, sometimes 
massed as stromatic tissues or forming a clypéus jover ene 
sphaeroid, thin-walled ascomata, and dark reddish brown, 
symmetric, thick-walled, VELTEUGCULose dictyospores. it 
appears to belong. in the Phaeosphaeriaceae (Barre aaug, 
Chaetomastia on the ‘other hand has the characteristlegw@ar 
the Dacampiaceae (Barr 1987), i.e., obpyriform vote ouc vor 
ascomata with a wide apical papilla, peridium of small dark 
cells that is widest and three layered in the upper regions. 
These characteristics and the dark asymmetric phragmospores 
are identical with those of the type species of Massario- 
spheaeria,, M. phaeospora (Muller) Crivelli. Thus 7 Sie aus 
necessary. to utilize the earlier name Chaetomastiag eco 
include phragmosporous species placed in Massariosphaeria by 
Grivellt (1983) ‘and Leuchtmann (1985), in adage tore 


hirtula. The enlarged concept of Massariosphaeria set forth 
by. Crivelli .(1983) that ineludes. dictyosporoUsmispecwes 
not accepted -here, not so.,much on account Ofj=eseoespoare 


septation but because the species, notably M. rubicunda and 
allies, differ in ascoma and loculé structure  tomtiewes, em. 
that they are to be removed to Karstenula (Melanommatales, 


Didymosphaeriaceae). At least ..some of the ‘speeieceecran 
Leuchtmann (1985) added to Massariosphaeria are closely 
related to the type of that gents. He noted that one group 


of species, developed -in arctic-alpine tregionsmyoumeatcoc 
substrates and a second at lower altitudes, mostly littoral, 
on monocots. The North American taxa included here have a 
similar diversity of substrate and distribution. 


Chaetomastia .(Sacc.) Berlese, Icon, Pyren.) lege LAS 
Melanomma subg. II1. Chaetomastia Sacc.Syll3 Fung 
Lae 1883. 
Massariosphaeria. (Miiller) Crivelli, Dissent yu neema 


FACHMS Rea PCAN ROUP ere St. 
Leptosphaeria subg. Massariosphaeria Muller, Sydowia 4: 
PAANSY clin’ ben bY 
Ascomata immersed becoming erumpent and appearing 
superficial when epidermis sloughed, separate or gregarious, 
obpyriform or ovoid to globose, at times somewhat elongate 


ellipsoid, medium sized; apex wide and)4.biunce ostiole 
rounded orirslitlike;: )peridium composed) of browne pseudo- 
parenchymatous cells externally, darkened and thickened in 
upper regions, internally of pallid compressed igawaue. 


cells, with sparse or numerous brown hyphae into substrate. 
Asci bitunicate, basal to peripheral, clavate ov eylinareges 


fies pO = t OC0 ee OSS DO Ged. Pseudoparaphyses cellular, in gel 
matrix’, extending into ostiole. Ascospores yellowish brown 
becoming dark brown or reddish brown, obovoid elongate, 


slightly or strongly asymmetric with obtuse upper hemispore 
usually shorter and wider than more tapered and more acute 
lower hemispore, usually inequilateral to slightly curved, 
three to eleven septate, constricted at first formedmeepoam 
cell above this usually enlarged; wall wide, dark, smooth or 
verruculose, surrounded by gel coating; contents with large 
globule in each cell; biseriate or uniseriate in the agoeues 


509 


Anamorphs coelomycetous where known; conidia hyaline or 


brown, one celled or one septate, (Aposphaeria- or Conio- 
thyrium-like). 
mee O Disc’, perhaps hemibiotrophic, Vig  WOOALY or 


herbaceous stalks and branches, alpine, or in monocot, cillms , 
low altitudes. 

Lectotype species Gris hirtula Char sits.) Berlese 
folements and Shear 1931, Farr et al. LOS lye 


Key to North American Species 


1. Ascospores 3-septate, usually less than eVventeped as cus: 
MERELY Gr A ie, LR gto ee ie ge eS ht ol, ob ete aa Se 2 
1. Ascospores 4-11-septate, usually OL SiG p cigar 6 Cl amie ee 3 
Z. Ascospores 17-25.5 x 6.5-9.5 um, four De tera. Chto. eras 
8 ess gy SE Sn SI: fe a an ae tert alsa Set er mA Yee Commi rr 1A 
Eemeascospores 14-25.'% 6-7'.5 ym, six (rarely eight) per 


RE PMI OUEN The be eos Fin Ss Vdd oe 15 APO My OES a nd (ate 6 dal Re, C. sambuci 
Peeearcospores: (20-)22-32 *« .6-7.5 pm, 4-6-(7-8-)septate; 
PNR oo EE al 4 ak cle" e le tke on sc ade ra | Cy. saémbucina 
Jeunscospores wider, 7-13 Mit ee oe ase Ce te Cue eke Ln 4 

4. Ascospores (3-4-)5-septate, 30-38 x 10-13 pm; lower 

MS Bae aon Mas a tho Wn. ct Boas Lass; Co kas he a C.-e@quiseti 

4. Ascospores (3-)7-1l-septate, relatively, narrower. 3.5 
5. Ascomata elongate to 1 mm; ascospores 31-39 x 7.5-10 pm, 
e  iteseptate, conspicuously asymmetric with long tapered 
Seereeeremilepore, lower altitudes>............0.. Cs. clavaispora 
oie Ascomata rounded; ascospores not so conspicuously 
EEE re tok 8 cS 2 ap eae ol e pues ece ro rik «uc. 4 et ohe' ato ote a ee 6 


6. Ascospores 25-39 x 7-10.5 wm; lower altitudes....... 


eS CB ae A her tie Eslue Sg 6 ges, seo Pe eee a anole el. C. phaeospora 


Chaetomastia clavispora (Cooke & Peck )} sbarra 2 COMD is eNOV.. 
Tha ee 2 eh danas Ta 
Hysterium clavisporum Cooke & Peck, New York State 
Museum Rep... 28: 69. LO 7 Gea DU lie wba al oso. 3: 
Bh. Loh, 


Dothidea clavispora (Cooke & Peck) Peck. News..ork State 
Pieseun ep. 12.98. Oo 3 5 Pod Se 

Rhopographus clavisporus (Cooke & Peck) Sacc. Syl 
Png 627648" Lo S3., 

Calospora clavispora (Cooke & Peck) Theissen & H. Sydow, 
See oaveOw er 13.) 428), ak ree 

Phaeosphaeria clavispora (Cooke & Peck) Barr in Barr et 
Bie NEW LorkeState Museum Bull. 45920115. LO S64 

Leptosphaeria clavicarpa Ellis & Everh. J. My Ou er Lae 
43. LoS D. 

Heptameria clavicarpa (Ellis & Everh.) Cooke, Grevillea 
on ie age EOS9 : 

Ascomata immersed, elongate, 500-1000 um or more, 200- 


300 pam wide, apex obtuse, opening by wide rounded or 
somewhat slitlike pore; peridium 26-40 Ditewi denis "soi 88 = 105 
fees 2). pm. Ascospores 31-39 x 7.5-10 pm, yellowish brown 
to dark brown, obovoid, asymmetric, long taper. to base, 


usually slightly curved, 9-1l-septate, sepctationg 2-1-6) or 


510 


3-1-7, constricted at first formed septum; Wabieeemooen 
surrounded by gel coating. 

In dead culms of Phragmites communis Trin., eastern and 
midwestern North America. 

Material examined: NEW YORK: Tyre,’ Sensca Coe oer 
1871,..C...H. Peck, (NYS, two) collections, isotype} eWeceeea 
Schuyler Co., Sep, Peck, Montezuma, Seneca, Coy pp ep meee 
Bergen swamp, Genessee Co., 14 Aug 1916, Ho) )D vee aiouee 
Bonaparte swamp, Lewis Co., 23 Jun 1920, House; Neweomb, 
Es Sex tenG Oy ae thd die Ri Mech URC Ry Ae ee House (ahh NYS NEBRASKA: 


Callaway, Custer\Co., Apr 1902, J.M. Bates, Funpi totes. 
(MASS QW ONYS )e 

The formation of elongate ascomata with a single locule 
or sometimes several. locules in a row, is one féeaturepor cue 
species. Another is the conspicuous asymmetry of the 
ascospores, the upper hemispore short and wide with two or 
three septa, the lower narrower with a long taper and six or 
S.C ViCiiees Ci Dic. 


Chaetomastia equiseti Barr, sp. nov...) Figs 0 oe 

Ascomata immersa discreta vel gregaria 330-440 pm 
diametro ovoidea, apices lati, peridium 25-35 pm tomentosunm. 
ASCL) DLCUNTCaticsoLavatie Lule LS) teal So <2 contin Ascosporae 30- 
38 xs 10-13 pm spadicea sed pallidae ad- apicem@eiusogaeae 
obovoideae 3-4-5-septatae, parietes verruculosi. 

In caulibus emortuis Equiseti hyemale L., “Spring Farm, 
Carrabassett, Franklin CGo., .Maine, 30 Aug L9/ eager 
Batr, 2007 (lLectim( MASS. noLotypus ).. 

Ascomata immersed, separate or /#greganilous ss je ee 
diam, ovoid, apex wide; peridium. 25-35 pm wide (omen cee 
with brown hyphae’. Asci 100-175 x 18-22 wm, | ASt¢oeno ream Oe 
38 x 10-13 ° pm, -bright brown, ends pallid, fusotameoo ve eee 
only jsiightly etasymmetr i cy 3-4-5-septate, constricted sa 
o> bye Z-3f 8 formed wisepta: cell above slightly" widew wall 
verruculose, surrounded by gel coating. 

On old stalks of Equisetum hiemalis, kno0wniieneweeeeoe 
type rcollectton,: 

This species evidently has not been found previously, 
for none of the taxa described in the literaturemiige meee 
OW mice The broad apices of ascomata make it more conspic- 
uous than Phaeosphaeria berlesei (Larsen & Munk) Hedjaroude 
which occurs in the same collection. 


Pigsiuvle Lota wpecves  oteiCnaeromas fT iar. 1-3.) 0G sce 
Ll. (ascoma.. in) vertical section;) 2). .ascus *s (35) a5 0 
4,9.°C., sambuci: 4. habit .of ascomata;,) 5) /ascos ness 2 ha a 
GC.  sambucina’ 6. habit of ascomatas. 7 vascosporvecs By) em, 
equiseris. 8A ‘habievsot escomats >) \\9ie we asceoseno ream LO dae 
Pphaeospora, ascospores. Na 628 Cyphicola, ascospores. 
12,132 ¢. clavispora;:; 12. habit of sasconata;, 13. Same e meee 
Standard-line = 150 pm for fig. 1, 15 pm for ascusangeaeree 
spores: Habit sketches not to scale. 


511 


512 


Chaetomastia hirtula (Karsten) Berlese, Icon. /FUN@ ees 
USO lee rae seal 5 


Sphaeria hirtula Karsten, Fungi Fenn. (xe 
18695 

Melanomma, hirtulum (Karsten) Sace. Syllivy hinge eee 
US S32 


Montagnula hirtula (Karsten) Leuchtmann, Sydowia 37: 
LWSe, GLOBAL Bie. - 
Melanomma glaciale Rehm, Osterr. Bot. |20 > Game 
1906. 
Melanomma suldensis' Rehm, Ann. Mycol, 5: S39 7s0Ge 
Ascomata gregarious, immersed or appearing superficial 
when epidermis sloughed, ovoid, 300-440 pm diam, up to 450 
pm high; peridium 40-45 pm wide. Asci 88-120 x, Diet See 
spored. Ascospores 1/7-25.5 x, 6.5-9..5 | jimmy ida eee 
obovoid, asymmetric, 3-septate, constricted at irstmiormed 
septum; wall verruculose, surrounded by gel coating. 
On woody canes of Rubus spp., herbaceous stalks of 
Cerastium, Veratrum, arctic alpine Europe, North America. 
Material examined: SWEDEN: Uppland, Dalby pare, 


Jerusalem, 2eMay -sLO8 O08 ska -& Gey Ho lim a coe FINLAND: 
Mustiala, May ©1866, Karsten,, Fungi  Fenn;. exsogueo 7 een 
isotype). CANADA: BRITISH COLUMBIA: ridge north of Mimulus 


Lake, ~Garibaldi Prov. , Park; 2) Aug (195255 MS se og 
(MASS). 


The tetrasporous asci and three-septate, dark 
ascospores are characteristic for this species.) (Ho mm 
found ascospores in the range of 22-30 xX )/ = Ogee 
European material from Rubus .and Cerastiunm. Leuchtmann 
remarked on some variation between collections from Rubus 
and Epilobium, with larger ascospores, and fLromeGenreer tum 
with ‘smaller ascospores (L1l- 20, "xy 725-10) pmee The North 
American specimen on Veratrum viride Ait. has ascospores 


that are intermediate in lengths between the Européan 
materials. 


Chaetomastia phaeospora (Mtiller) Barr, comb. nov. Fig. 10 
Leptosphaeria phaeospora Muller, Sydowia 4: 208. 1950. 
Trematosphaeria phaeospora (Muller) Holm, Symb. Bot. 

Up Saleem 3 deal 6:5. 1 Oc5o/ me, 
Massariosphaeria phaeospora (Muller) Crivelli, Dissert. 
BT 7S an Ge ae ore : 
Ascomata immersed, ca. 440 pm diam, 500-550' pm high, 
obpyriform; peridium ca. 15 pm wide below, light brown, dark 
brown and up to 30 pm in upper regions, surrounded by scanty 
brown hyphae. Asci, 150-180 x 20-23 pm, octdsporomter Asco- 
spores 26-36 x 9-12 pm, dark» brown, obovoid,) \asymmeqaues 
iisUadly Ses livnelyeycurved, 7-9-(11-)septate; wall. ther, 
smooth or verruculose, surrounded by gel coating. 
In stalks of Artémisia, Trisetum, alpine Europemseen eee 


douglasii Hook., alpine western North America. 
Material examined: SWITZERLAND: Kt. Wallis, #indedven- 
LO. Sep; D695 9H Wegeline (Zl sholotype) .» GANADA. BRITISH 


COLUMBIA: Ridge north of Mimulus Lake,;, Garibaldi Praveepagee 
Jane Le oe Mee hee Bey re Oe OM ASS) es 

The North American collection matches the holotype 
material quite precisely; the only difference seen is that 


513 


ascospores in the holotype are smooth walled beneath the gel 
coating whereas those of Barr 614 are verruculose. Muller 
(1950) found ascospores up to 41 pm long. Leuchtmann (1985) 
obtained Aposphaeria-like conidiomata in culture from a 
collection on Trisetun. 


Chaetomastia sambuci (Earle) Barr, comb. nov. PME So - Ge 5 

Melanomma sambuci Earle, Bull. New York Bot. Garden Coe 

Zi  LIOG. 

Ascomata gregarious, sides connected at times, immersed 
becoming widely erumpent, globose, 385-550 pitines Cacainve wot 
forming small stromata up to 1 mm diam containings three or 
four locules, apex rounded with minute papilla, surface 
ar togular or) rugose;. peridium (13-)25-50 pm wide, with brown 
hyphae at base into substrate. AS G49 4657-410 014-1 30) ese 81 01-11 3 
pm, mostly 6-spored. Ascospores (14-)18-25 x 6-7.5 gn, 
reddish brown, 3-septate, cons Caicted wate fits ti tormed 
septum, cell above enlarged slightly. “wall. werruculose, 
surrounded by gel coating. 

Associated conidiomata similar in structure and sizes 


PopsoscOmMata; conidia 5-9 +x 4-7,5 pm, brown, globose to 
ellipsoid, one celled or one septate. (Coniothyrium sambuci 
Earle was described from Nevada as Baker 1181 paxctile 

On dead branches of Sambucus spp., alpine western North 
America. 

Material examined: MONTANA: Entry eG ball siGhactiear 
eomeeeck. «27 Jul 1961, W. Bore V..Gan Cooke 32665 (MASS). 
NEVADA: Snow Valley Peak, Ormsby Co., 14 Jun ie? 0. 2p ese Bs 
Peper tl 82 parti (NY; holotype). 

tieewoth collections cited, sthe “ascomata are widely 
erumpent over immersed, empty ascomata and hyphae that had 
matured beneath the periderm. No features remain to show 


whether these ascomata were of the same or another taxon. 
Chaetomastia sambucina has more elongate and asymmetric 
ascospores that develop additional septa. it Ges pos sib.ve 
that these are simply variations within one species. 


Chaetomastia sambucina (Ellis & Evexh jiaibarteutcomb ae novi 

(ROS Es ane 

Leptosphaeria sambucina Ellis & Everh. Proc. Acad. Nat. 

ecise Philadelphia, 46: 4336. S94. 

Ascomata gregarious, occasionally sides connected, 
immersed, 495-550 pm diam, 500-825 pm high; peridium ca. 30 
pm wide, blackened above and up to 60 pm wide, ample brown 
Byeuaesinto substrate.. Asci 80-120 x 12-15 pm, octosporous. 


Ascospores 22-32 x 6-7 pm, dark brown or reddish brown, 
narrowly obovoid, 4-6(7-8-)septate; wall verruculose, 
surrounded by gel coating. 

In branches of Sambucus melanocarpa Gray, alpine 
western North America. 

Material examined: COLORADO: Cameron Pass, Jackson 


Stpeece Jul. 1894, C.,.F..Baker (MASS sa isotyped t 
Conidiomata are intermixed with and similar in, 


appearance to ascomata; conidia are brown, globose, one 
celled, 7-9 mn. 


514 


Chaetomastia typhicola (Karsten) Barr, comb. nov. Fig. 11 
Leptosphaeria typhicola Karsten, Mycol.) Fennec eeu 
S732 


Phaeosphaeria typhicola (Karsten) Hedjaroude, Sydowia 
Lense OO ae 6/8, 
Massariosphaeria typhicola (Karsten) Leuchtmann, 
Sydowia 37: 168. CLS 84) ) 19857 
Leptosphaeria occidentalis Ellis & Everh. Erythea 2: 
A Oona eh n 
Leptosphaeria baldingerae Fautrey & Lambotte, Rev. 
My cO iw Chauvouel oil 93. io ai, 
Phaeosphaeria baldingerae (Fautrey & Lambotte) 
Hedjaroudeswisydowlaw2 2 ive c/a Lo Gee 
Ascomata immersed, separate or gregarious, ovoid or 
globose, at times ellipsoid, 245-430 pm diam; peridium 15-25 
pm wide, surrounded by tomentum of interwoven hyphae. Asci 
COM). 9 OS Sia kee ak Bel, Ascospores (25-39) °xe@ 310 ae 
yellowish brown becoming dark brown, asymmetric, slightly 
curved, 3-7-9-septate; wall verruculose, surrounded by gel 
Coatings 
In old culms of monocots, low altitudes, Europe, North 


America (Echinochloa crusgalli’(L.) Beauv., Panicum virgata 
Loe; Zea mays) LL. in North. America; ) 

Material examined: MASSACHUSETTS: North Amherst, 
Hampshire)Co., 29; Apr 1959, Mo EqmBarr 25499 (Ao ae KANSAS: 
Rockport, Rooks) Co. “Apr 1893, EF. Bartho lome wee oe 


Evexrhs.s NAFioi2. 925 9 CDAOMi a MASS. isotype L.. oceiventatisg.., 
Stockton, Rooks. Co.,22 May, 1906, E. Bartholomew rungim cos: 
26:2:9) (CMAS Sin 

The concept of this species as a polyphagousm@eone 
follows | that ‘of. Leuchtmann (1985) who also \deseritpeneenre 
formation of Aposphaeria-like conidiomata or ascomata of the 


teleomorph from different isolates. Lucas and Webster 
(1967):,also Sobtained the, anamorph Jinvculture, The isotype 
material of  -L., occidentalis is’ ,not” separabDlepe. .anmee 
typhicola. The North American specimens that I studied do 


not show the full range in variability of ascospores noted 
by Leuchtmann which caused him to combine L. typhicola and 


L. baldingerae as one_ species. Kohlmeyer and Kohlmeyer 
(1979) +» provided a description jof L: typhicolawiteewneen 
ascospore , sizes’. are (\34-152,(=62)% x 7-10-13) geome They 


reported the: ‘species’ from ‘decaying, culms) Of (Viicuegeeue 
Spartina in marine sites in North Carolina and Argentina. 


I acknowledge with gratitude the curators of herbaria 
(FH, NY, .NYS', UPS, ZT) whose loans of specimens @mageme am 
study possible’, ,and thank’ Dr. J.) Boise for "Yeviewin eee 
Manuscript. 


LITERATURE CITED 


Bary sy ieetee LOB 7s Prodromus to Class Loculoascomycetes. 
Publi vbhyethe author, Amherst) MA, To Sear 

Berles ee AGgn: 1891. Ieones fungorum, Vol. 1, fasee ieee 
501% 

Gannon, D. F., D.'L. Hawksworth ,and M. A. Sherwood@s ees 
1IgsS./ Then briatishvAsconycotinarce CNL uke we 3025pe 


515 


Seemente. F.,.—E. and G., L. ‘Shear. 12 a The genera of 
ines. H.W. Wilson’ Co.;" New York, 496 pe 
Seavekii, PP; G, 1983. Ueber die heterogene Ascomyceten- 


gattung Pleospora Rabh.; Vorschlag fur eine Autre dln. 
eee eee 2rd Ch 1 Si 1 HF 
Eriksson, O. and D. L. Hawksworth. 1987. Outline of the 
Peeomyceteish = 198i! Syst Ascomye. 6: 259-337, 
S@rceeewn. ; J. A. Leussink and KF. A. Stafleu. LO Sos Index 


Nominum Genericorum (Plantarum). Regnum Veget. 100: 1- 
De.3.6%, 
Hmoim, L.. te is Etudes taxonomiques sur les Pleosporacées. 


pamper DOCS rUpsal. 4.03) 11-188), 
Kohlmeyer, J. and E. Kohlmeyer. 1979. Marine mycology The 
higher fungi. Academic Press, New York. 690 De 
Ceuchtmann, A. LODO Cs S ay Uber Phaeosphaeria Miyake und 
andere bitunicate Ascomyceten mit mehrfach querseptier- 
PenvAscosporen. Sydowia 37: 75+194. 


Pieces, Le and J. Webster. POG Conidial states of 
British species of Leptosphaeria. PDaANS ee Die ee niey CO: . 
peewee 2) 85-121 | 

Biller! oF, E9505 Die schweizerischen Arten der Gattung 
Leptosphaeria und ihrer Verwandten. Sydowia 4: 185-319 

munky A. SNS Lh Danish Pyrenomycetes. Danskiebot we Ark. 


LIA Cy a er: i 
pacearoo PP. A. 13.83% SyVuLoge.Funvorum, Volo 1. Patavia. 
pee ere Lx Exp 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 517-526 January-March 1989 


THE GENUS DOTHIDOTTHIA (BOTRYOSPHAERIACEAE) IN NORTH AMERICA 


MARGARET E. «BARR 


Department of Botany, University of Massachusetts, Amherst, 
MA 01003 


ABSTRACT 


Dothidotthia is accepted for a group of Botryosphaeria- 
like species whose ascospores are mostly one septate and 
brown. inesscype species (is .redescribed as D:s aspera, .D’. 
ramulicola is accepted, and other North American species: D. 
Peaiaudeoe.  trutcicola,: D.. lasioderma, D. quercicola, are 
proposed, in new combinations. Dothidotthia diapensiae is 
described as new. 


The genus Dothidotthia is recognized as a member of the 
Botryosphaeriaceae (Barr 1987). As. in, species of Botryo- 
sphaeria, the erumpent ascomata, whether separate or 
grouped, have a peridium composed of large, brown, pseudo- 
Patcemcuymatous cells and produce , more or less abundant 
coarse, brown hyphae. Tire was CissaremopLong eclavates with 
Pnictewendotunica, and are basal among relatively wide, 
cellular pseudoparaphyses. The ascospores separate species 
of Dothidotthia from species of Botryosphaeria:, they are 
Puecesm OL brown and one to three . septate. Associated 
coelomycetous anamorphic states are known for a few species 
and are Diplodia-like. 

oma idiot th ia was described oO PSCUGCOt Eni a 
symphoricarpi Rehm, as a member of Eu-Montagnelleae in the 
Montagnellaceae of the Dothideales (von Hohnel, 1918, 1919). 
Petia cl 24.) recognized that, 1t. was separate from Otthia (a 
genus of the Cucurbitariaceae) and compared Dothidotthia 
with Antennularia and Pseudotthia. Later 3(Petrvak. 1927 ))-he 
decided that it was not separable from Dibotryon. Von Arx 
(1954) removed D. symphoricarpi to Gibbera, and Muller and 
Women rx 1962) included both Dothidotthia and Pseudotthia as 
synonyms of Gibbera. Shoemaker (1963) concurred, as do O. 
Eriksson and Hawksworth (1987). But Gibbera, as well as 
Antennularia (= Protoventuria) and Dibotryon, belong in the 
Venturiaceae. Their ascomata have a peridium of small 
Peeudoparenchymatous cells, hyphae are narrow, asci are 
oblong or cylindric and have a thin endotunica, pseudopara- 
physes ave em narrow dand © often. “deliquescent., and known 
anamorphic states are hyphomycetous. CELA. AlLSsos- OULLers 
from Dothidotthia in having a three-layered, smaller-celled 
Petacium; although the .anamorphic state of O. spiraeae 
(Fuckel) Fuckel does not seem to be generically different 
from that of Botryosphaeria obtusa (Schwein.) Shoemaker or 


518 


Dothidotthia ramulicola. 

The examination of numerous collections described or 
disposed under a variety of generic names (chiefly Otthia, 
Didymosphaeria, Parodiella, but also Amphisphaeria), permits 
compilation of the species of Dothidotthia in North America. 
Two series of species’ are evident in the genus) Jone wien 
light clear brown or light reddish brown ascospores (eone lo. 
three septate, the other with dark reddish brown ascospores, 
mostly one septate. 


Dothidotthia von. Hohnel, Ber. Deutsch. .Bot.) Gee oer 

Eos 

Ascomata immersed becoming erumpent, finally appearing 
superficial, usually gregarious, ,often in rows Jor ss Dieome 
rounded groups, at times ‘connected at’ sidés) ) etanboce: 
sphaeroid or ovoid, medium Sized. rarely small; apex 
rounded, with short or well developed papilla, often opening 
widely by ‘rounded ostiole, stuffed with hyalbinewecebie: 
surface smooth or roughened with protruding cells or bearing 
short to elongate hyphal appendages; peridium wide, composed 
externally of rows of large, brown-walled, “pseudoparven- 
chymatous cells, often blackened over surface, internally of 
more compressed rows of pallid cells, at times wedge-shaped 
groups of cells extending from lower sides, or basal portion 
of peridium thickened and hypostromatic; hyphae dark brown, 
coarse, forming slight or well-developed subiculum beneath 
and. connecting ascomata. Asei bitunicate, basal setavaceson 
oblong, endotunica thickened. Pseudoparaphyses cellular, 
usually wide. Ascospores yellowish to clear or dull brown 
or dark reddish brown, ellipsoid, fusoid;, (obGvoi ean. 
obtuse or somewhat acute, Straight. inequilateral or 
slightly ‘curved, one to three "septate, intréequentivagone 
celled or two'-or more septate, not or slightly veconccriceen 
at septum; contents minutely granular, finally) onemetengae 
DeL A Cele aiiwia ll watch Lc ke smooth or wverruculose sae eee 
occasionally surrounded by narrow gel coating; overlapping 
biseriate in the ascus. 


Anamorphic states: coelomycetous where known; 
conidiomata similar in ‘aspect .and structure) toma eeomeeee 
conidiogenesis holoblastic; conidia brown, Ong ute@ed ego 
Scope dices 

Type species: D. symphoricarpi (Rehm) von Hohnel = 
Plowrightia symphoricarpi Ellis & Everh. = D. “asperaueue 


S&S BPVerh |) Bars 


Key to North American Species 


1. Ascospores light brown (clear brown, yellowish brown, 
reddish brown), often inequilateral or slightly curved, 1-3- 
SSP Ca Cer 6 ce ies od ve wis a's hls in Leen! did fe la Nota elite upscale Sm fee a 2 
1. “Ascospores dark reddish brown, usually straighteeoueeee 
C2 )SEPTEt Ger er cel Sie a. ecw shamed oon Weg bet So tape lens eich grantee 6 
2. Ascomata much depressed; ascospores 14.5-20 -3505= 
6 4)5) Se l-septate, in short wide asci|...D. @iepeneeee 
2. Ascomata not strongly depressed; ascospores larger, 
Iin-oblong, clavate ascii... 7. wu. cel oen eel 3 


3. Ascomata clothed with hyphal appendages; ascospores 3- 
Sép Cate ce Oe aoe 6 Comite o> Le i ieee oe eee D. lasioderma 


DL9 


S. Ascomata not clothed with hyphal appendages, slightly 
lee co spores ® lai 3 sep tates, a. Wien ods ees Gee eae Le 4 
4. Ascomata causing hypertrophy in twigs; otpvOuvercus: 
ascospores 20-24 x 9-10(-11) pm, obovoid..D. quercicola 
4. Ascomata not causing noticeable hypertrophy; asco- 
eee oy imei Latig Gio) TAs. eed ih mob atone ore a eg blll 5 
59. Ascomata globose sphaeroid (diam same or greater than 
merent)e s ascospores (17.5-)20-30¢-32) x CS Or Sele P25 


MMII SE os Ay ig Schock cca «ni alah) foc £5 S din, so ss LPT Teh las Oe a ee a Oe Be D. aspera 
59. Ascomata globose to ovoid (diam mostly less than height); 
Seems, O- 39/) 10-29% - 15) 5) amon ds i dee Div tae noo le 
6. Ascomata much depressed; aSGoS pores BUF <12:7 eres 3 am 
eS EARIEPTGS Tse WR ecdonk AU e ae ID oe GP ge ea CER ce Diwiceloidis 


6. Ascomata not strongly depressed; ascospores (15-)18- 
Poiteo4) x (8-)10-13(-16) um in oblong-clavatey asei... 4% 
MN tts eget ea Perce Gesine eke Foy tale Rte Ua aha taps aleve Ra He rake ck Dey ramulicola 


Dothidotthia aspera (Ellis & Everh i barre comb) “nov: Figs, 

1-3 

Amphisphaeria aspera Ellis & Everh. Bull. TOrLeys (bot 
Ciabe2 7/2) 522. £1900) 

Peewee ntial symphoricarpiwEllis’ -& Everh.. Proc. Acad. 
Saree oc. * Philadel philava2s (249). 1890. 

Peewee yiphoricarpil (Ellis. & Everhs)¢ Ellis (&  Everh. 
Norehe Amer. Pyrenomye: 249, :.° 1892. 

Valsaria symphoricarpi (Ellis & Everh:)) Theissen  “& Ht 
ec owor Annee Mycolsy ls 327019 LS. 

Pseudotthia symphoricarpi Rehm, Ann. MYC OTS We 608, 
RLS. 

Dothidotthia symphoricarpi (Rehm Mivon | Hohnet" Ber. 
pegisch.| Bot) Ges i136: 4312: L9LSe 

Dibotryon symphoricarpi (Rehm) Petrak, Ann. Mycol) 25% 
OU Lem O27 %, 

Gibbera symphoricarpi (Rehm) von Atx  AGtay Dot. Neer ly 
Daou, a) 95.4.) 


Gibbera andersonii Shoemaker, Canad. J. BOCan yoink 
BAe hi G3). 

Otthia fendleraecola Earle in Greene, Pl, Bakerianae. 1: 
Pee el 901 

Otthia distegiae Tracy & Earle in Greene, Eis. 


Bakerianae 1: 29, VS O78 te 

Ascomata erumpent, gregarious in ZrOups OL LOWS )s occas = 
Sionally separate, (180-)300-660 proivdiamy) (180-j 320-495. im 
high; apex rounded, shomte/*papiivater Ott ene Ms tightly 
depressed around papilla; surface dull Hlachiwwrsi 1 eh tly 
roughened; peridium (20-)40-60(-80) pm wide, of brown cells 
up to 13 pm diam, with coarse brown hyphae into substrate. 
mera (65-)80-120(-140) x (10-)16.5-23 pm. Ascospores (17.55 
Pere (i 32)). 1 x CO ose ieta) lets Get kee yy ute umn UVehteteceara eb rown, 


Merepsoids ends tapered, obtuse..or” semewhat acute, often 
inequilateral, 1-(3-)septate, slightly constricted: wall 
smooth. 

In woody branches (Fendlera, Lonicera, POpmliys % 


Symphoricarpos, Tetradymia), mid and western North America. 
Material examined: BRITISH COLUMBIA: ‘Soda Creek, 3 Jun 

See). Bancroft’ (UBC 23m 6k NORTHPDAROTAS)) Kime eh iD 

Peenckle, Rehm Ascom. 2040 (BP IQIINYS Misotypel Peeudorehy a4 


520 


symphoricarpi); 7 Mar 1909, N.D. Fungi 260, Fungi Dakotenses 
98 (BPL, NY,.as.Otthia symphoricarpi);.10 May 196 ye ee 
exot.: @xs. -391 (BPI, NY, ‘as -Pseudotthia’ ‘sympnhcreceu en 
MONTANA“* Sand Goulee; 24° Dec 1888, F. W. “Andersqnmagena 
holotype Plowrightia symphoricarptl) ; ‘Helena, I@Aprevouueme. 
D. Kelsey (NY); Great Falls, F. W. Anderson, NAF 2374 (MASS, 
Nise COLORADO: Montrose, E. ‘Bethel ‘S17 “CNY Rhos pe 
Amphisphaeria aspera); Hermosa, 4 Apr 1899, (C7 OF) (Bateray> 
and Pilve Southern Colorado 432 CNY., holotype QLCITIA 
fendleraecola); Berkeley Bluffs, 12 May 1906, Fo See Seavera. 
E.) Bethel (NY as Otthia symphoricarpi); Mancos>=2om uae oee 
Pl. Southern ‘Colorado 1090 (NY; holotype” Otthtiameveee zeae. 
two packets);. Durango, 20 Mar 1899, C.. F. Bakepe iano 
Southern Colorado 43 :(NY);' Durango,* 2) Jul LOO7SR er eee 
SaiGlementspwicryp te Form.) Coorad. e439 sun ye OREGON: 48 km 
N Madras, | Wasco Co., 18 Aug.1977,;' An, Y. RossmanmiGiagae 

Some variability in aspect, shapes and sizes is evidence 
among these collections, but the overlap among them makes it 
impossible to separate more than one species. Although the 
specifiie epithet :.of . Plowrightia. symphorrearp is the 
earliest, combination in Dothidotthia is precludedeaayae. 
symphoricarpi (Rehm) von H@dhnel. Shoemaker (1963) provided 
Gibbera andersonii as replacement. He diseussedseand 
clarified confusion with this species and Griphosphaerioma 
kansensis (Rehm) Shoemaker, which is also present on several 
Oofvstherscolklect ions: Amphisphaeria aspera provides’ the 
earliest available name now. The ascospores are typically 
one septate and ascomata lack the vestiture found in D. 
lasioderma. 

Thegmtyoeve specimentiot raeA: aspera bears conidiomata 
intermixed with ascomata and similar in appearance. Conidia 
are (10-)12-15 -x 6-7 pm, -darki#brown, (l-)3=séptave swarn 
occasionally a-longitudinal septum in, onevecel ia 


Dothidotthia celtidis (Ellis & Everh:.)) Bawrge compe 
Fle Semel Joes 
Didymosphaeria celtidis Ellis & Everh.) Proce Acadmaaes 
scivs Philadelphia’ 47 2842, 1e2 aloo 5e 
Microthelia celtidis (Ellis & Everh.:) 0. ¢8 ite 
Gen AES pice? oy Leos: 
Ascomata immersed, apices. only emergent, scattered to 
gregarious beneath and raising periderm, sphaeroid, 220-440 
ym diam, 130-275 pm high, apex blunt; peridiume 25-308 pmeeoe 


large brown pseudoparenchymatous cells. Asci 60370" x 325-23 
pm. Ascospores 19-27 x 9-12.5 «pm, “dark ‘browne omaue 
ellipsoidal (Oc) l-septate, -not constricted; Gwar ee 


smooth, .at times surrounded by gel coating. 


Figs. 1-15. Species of Dothidotthia. 1-3. D )aeie. eee 


ascoma: in vertical-secition.»2i,ascus,) 3)%ascospomece i ee he 
DetdLapensd 4e2g4aascuseio, kascogpores. 6, 7/7. Di *Erap tears 
6. habit-wot ascomats,. 4/4. ascospores. 8, 9. D. Testoverue 
8. habit of ascoma. 9. ascospores. 10, ll. D. queretegee eee 
habLt. off ascomata., We ees.cosnones. 12, 13. De Seahorse 
12. ascomasingwvertical sectian. 1b3% aseospoces:, 14) ee 


ramulicola: 14. habit of ascomata., 15. ascospores:) Stanamsed 
line,= 150 mpm for ascomata; 15 mm for asci and aseésponea™ 


521 


22 


In) *branches ‘of Celtis occidentalis’ L.j)@ \@aeGe0neeeo 
midwestern North America. 

Material examined: NEW YORK: Orient, Long Island, 14 
Fep “1b923..) Roy La thanieicny9+. KANSAS: Rooks Co., 23 May, Lega, 
E. Bartholomew 1467 (NY, holotype, 2 packets). OKLAHOMA: 


Ripley Bluff, N. of Ripley, Payne Co:, ll Aitg 7a 
Barr 6586 (MASS). 

The: protologue gives May 1895, but the tjcotteggion 
marked as type is dated 23 May 1894. This taxon is retained 
separately from D. ramulicola on the bases of much depressed 
ascomata and shorter asci. The ascospores are quite similar 
in both taxa. An anamorphic state is present on some of the 
twigs; Sphaeropsis celtidis Ellis & Everh. (Amer 7 Natwe 7a. 
1897) was described for such a species, having, brown, yone- 
celled: conidia llS=21)'%)\8>10\nm! 


Dothidotthia diapensiae Barr, sp. nov. Figs. 4, 5 
Ascomata immersa subcuticularia sphaeroidea 105-135 pm 
diametro,.7 75-90 “sm ‘alito, ) peridial ad “255 umperaees Asci 


bi tunieatci oblong? 6645 iwi ote e ome Ascosporae 14.5-20 x 
5.5-6.5 pm flavidae brunneae obovoideae uniseptatae. 

Lark fvo/ lias Diapensiae lapponicae, "Northwest 
Territories, Baffin» Island, head of + Clyde “Intetweaeea. 
1950," a P. Dansereau '5007062299a Lecti' (MASS, hototyowere 

Ascomata immersed, SUDeWE Lewis re visible through 
thickened cuticle, much depressed to almost lens shaped in 
section, 105-135 pm diam, ./5-90 (pm) high, apewemeneeen. 
papillate; peridium ca. 25 pm wide at sides? )LOMiimietowaca 
apex and base, composed of large, brown, pseudoparen- 
chymatous cells, brown hyphae penetrating substrate between 
epidermal cells. » Asci 36-57 x, 25-28 pm. \Ascospot eames 
x 5.5-6.5(mam, yellowish to light dull brown; 6Qevald gas 
obtuse, tapered to base, one septate; wall smooth. 

In leaves of Diapensia lapponica L., “knewn@omivarccom 
theitypevicol lection. 

This small species’ is unustial _in habit for @eteueenaoe 
but the large-celled peridium and ascospore shape _ and 
pigmentatlon permitrmrofyine ‘others ‘disposition: It must be 
rare, for I have seen only one collection, and iBeiie amen 
(1974) doesimoty reportvany isuchyspecies:: 


Dothidotthia fruticola (Ellis & Everh.) Barr, comb. nov. 

Rigs Mi Gruy 

Parodiella fruticola "Ellis & Everh.. J. MyocolSeeigeee 
1888. 

Plowrightia fruticola Ellis & Everh. in FL WeeAmoeewo 
Jiu IMac Odl etait 8. 4h OLB 8.9. 

Otthia fruticola (Ellis & Everh.) Ellis & Everpiweenowen 
Aner. Py renomyc.) 207 HS O2e 


Otthia clematidis Earle in’ Greene, Pl. Bakerianeauwee 
Dd ee Oe) #: 
Ascomata immersed erumpent in long rows, globose or 


ovoid, 275-550 pm diam, 550-660 pm high,  apéexroumeee 
papilla inconspicuous; surface dull, rough with short hyphae 
and protruding cells; peridium ca. 50 pm wide, up )tomheomum 
at base, composed of brown, pseudoparenchymatous cells, with 
ample brown hyphal subiculum surrounding base and lower 


025 


Seoees.  Asctl 115-160 x,20-22 pm.) Ascospores 26-39 x 10-12(- 
15.5) pm, yellowish brown to light brown, ellipsoid, tapered 
to acute ends, 1-(2-)septate, not constricted at septum. 

In stems of Clematis ligusticifolius Nutt., also known 
on Sambucus sp., western North America. 

Material examined: BRITISH COLUMBIA: Soda Creek, 9 Jun 
Pet, sts Bancroft (UBC 2335). MONTANA: Sand Coulee, 11 Feb 
oeec, oF. W. Anderson 124 (NY, holotype of iP. eirurtecala, 2 
packets), also as NAF 2129 (MASS, NY). COLORADO: Gulch 
between Sheep and Humboldt Peak, Sangre de Christo Range, 9 
Sieeeoon, GC. H. .Demetrio’ 212 “(NY as Dothidea DTSECULD La)”: 
Peeemeollins, Apr )1894, C.F. Baker (NY). Jun USO Ge ea O Le 
Meme, 21 Maer 1896; 'C...F.. Baker..(NY as 0: clematidis); 
Durango, DomeM 27 A 8.9:9) 4" 1Cy, Feembaker 6a andy. Piso aSouthern 
Colorado 41 (NY, holotype of O. Clematidig ne For tiscarlands 
eee ane 907 oF. EB. -& E.S,; Clements, Cay pDteurrOormestCo.orad) 
432 (NY as O. clematidis). 

Dothidotthia fruticola has larger ascospores than D. 


aspera and more ovoid ascomata. Most ‘of the collections 
cited above bear also Dothidea insculpta Wallr. = Scirrhia 
Peecuupte. (Wallr.) Barr’ (Barr, 1972). This species forms 
shining, multiloculate ascomata containing numerous. small 
locules. The European and Asian Otthia lisae cde Not. ) 
Sacce. is similar in ascospore shape and sizes and in the 
Bterile base of ovoid ascomata. That species probably 


belongs in Dothidotthia also, but the ascomata have a three- 
layered peridium (Muller and von Arx, 1962). 


Dothidotthia lasioderma (Ellis & Everh.) Barr, comb. nov. 
Paresh Bie. 9 
Leptosphaeria lasioderma Ellis & Everh. Proc. Acad. 
NAtawocil Philadel phiai45 :.135°. LOO 37 

Gibberidea symphoricarpi Tracy & Earle in Greene, Pl. 
BeRe@rd ana el 242 8a) 190 1 

Herpotrichia symphoricarpi (Tracy & Earle) Barr in L. 
Holm evens k Bote Ti ds kins 96 2323.0) 1968. 

Ascomata erumpent separately or in small groups, 
globose or nearly so, 450-550 pm diam, 400-500 ym high, apex 
rounded with short papilla; surface clothed with short to 
elongate, brown, septate hyphae, mostly recumbent, forming 
limited fringe of subiculum around base; peridium 28-33 jum 
wide, of brown pseudoparenchymatous cells 5-8 pm wide. Asci 
meorieoy x 1655-20 pam. AScospores#)2 4:39 O-3'8 i006 2/7 5 lik 
pm, light reddish brown, oblong ellipsoid, tapered to less 
pigmented obtuse ends, usually slightly curved, 3-septate, 
slightly constricted; wall smooth or finely verruculose. 

In woody branches of Artemisia, Symphoricarpos, western 
North America. 


Material examined: GOLORADOi) Bobe Creekw. W) of «Mtl. 
meeperus,, 27 Jun 1898, C. F: Baker; Pl.. Southerm: Colorado 
173 (NY, holotype G. symphoricarpi, 2 packets). NEVADA: 


weeucemont, Oct 1892, ..M. Ei aeOn ese 2) eGNYerh oloty peel 
lasioderma). 

peoam indebted. to .Dr. J.) Boise, fon the suggestion that 
Leptosphaeria lasioderma could provide the earlier name. 
Despite the presence of hyphal appendages on the ascomata, 
Serer. aspects, i.e:, the large pseudoparenchymatous cells 


524 


composing the peridium and ascospore shape, are congruent 
with species of Dothidotthia. Both L. Holm (1968) sande Sane 
(1984) expressed reservations about placing G. symphoricarpi 
in Herpotrichia. The redisposition of this species removes 
a disparate element from Herpotrichia sensu str. 


Dothidotthia quercicola (Ellis & Everh.) Barr, comb. nov. 

Picg Sieg tO aaa 

Otthia quercicola Ellis & Everh. North Amer Pyvencmyae 

2 ORE IC GS ae i 

Ascomata erumpent, densely gregarious on _- stout 
stromatic base, dull black, 220-330 pm diam, apex rounded; 
manittie dv papi lL tat er slightly depressed around papilla; 
peridium ca. 30-40 pm wide at sides and over apex, up to 120 
pam at base, composed of reddish brown, pseudoparenchymatous 
cells, 8-10 pm wide, with coarse reddish brown hyphae’ ints 
substrate. Ascivs/ 0-80 “xe l8 = 2.0) (ome Ascospores 20-24 x 9= 
LO(-11) pm, .light: clear brown, obovoid, apex obtuse yer apeued 
to obtuse base, l-septate, not .or slightly constricted aware 
smooth. 

Causing some hypertrophy in small twigs of Quercus alba 
L.wiNew Jersey, known only, from type collections 

Material examined: NEW JERSEY: Newfield, 30 Jane 1ss07 
Jemibs Bids 263-05 (NYY ahoLotype)= 

This species is close in sizes to the midwestern and 
western collections of D. aspera. The more obovoidmanamiee. 
symmetric ascospores and some hypertrophy of the host serve 
Gos dis tinzsurshivDe -quercicolay 


Dothidotthia ramulicola (Peck) Barr, Mycotaxonm@izeeemoo 

169 Seo re a cen trad 5 

Sphaerta ramulicola Peck, Ann. Rep. New YorkeStatesMuce 
Zoran LOARRaGC OT | Stik )weUolere 

Leptosphaeria ramulicola .(Peck) Sace?) \S¥iiietunceeee 
Paley es” Awei 0) Sh 

Neodeightonia ramulicola (Peck) “Barragin = Bape cee 
Bull. New York State’ Mus: 4595981395) S1ogee 

Didymosphaeria accedens Sac.icr, in Faitman; PEOCG 
RochestermAcadyradys 1890. 

Microthelia accedens- (Sacc.) 0O. Kuntze, “RévoeG@ernmeres 
3 eS Site” A LBSSs. 

Didymosphaeria vagans Ellis & Everh. Proc. Acad. Nat® 
Sei .\Philvadelphial45eC446. S933 

Microthelia vagans (Ellis & Everh.) ‘0. Kunezeneerens 
Coney P Ay, 3 Geo e) U89 Be 

Valsaria magnoliae Ellis & Everh. J. Mycol iO@eeigoe 


1904. 
Ascomatas greparious to crowded with sides ‘grown 
together at times, immersed beneath periderm, becoming 


erumpent in rows or groups, at times separate, globose or 
sphaeroid, (200-)300-450(-550) pm diam, 220-330(2385)0u am 
high; apex rounded or short papillate; peridium 35=50e 
pm wide, surrounded by ample coarse, dark brown hyphae. 
Ascii 25 42 0 G15 5:) XE 43:09" aims at times only 4-spored. 
Ascospores (15-)18-25(-34) x (8-)10-13(-16) pm) dark brown 
ellipsoid,» oblong or subglobose, ends rounded, (O= yas 
septate, noty VOrehecatcely #kconstricted: wall smooth or 


525 


verruculose, surrounded by gel coating at times. 

Anamorph: Diplodia-like: Conidiomata similar to and 
Zia mixedrwith ascomata; -conidia’ @L0-)15-23x% (€5-)7.52410:5 
pm, brown, broadly ellipsoid, one celled or one septate. 

In twigs and branches of various woody angiosperms 
(Carya, Ceanothus, Fraxinus, Magnolia, Menispermum, Ostrya, 


Uilmus), eastern North America with occasional collections 
from midwestern or western states. 
Material examined: ONTARIO: Londoners. Mave els93 \. 1 J); 


Dearness (NY with Fenestella princeps); London, 30 May 1893, 
Semveernessy72110, 211358 and 31 May, 2113 (NY, syntypes of D. 
edie) sid ot o93) (NY as’ DD. vagans): Nov 1903,7°F. Col. 2100 
(MASS, as Valsaria magnoliae). MASSACHUSETTS: Conway, 8 Feb 
meee bart oLSl (MASS); Baptist Hill ;}* Conway; 30 *Deec 1979, 
meio] 6) TOMAS S ).. NEW YORK: Greenbush’, Mayers La 'G:.eH! 
fee ee Ntoe holotype Spheaeria ramulicola), ‘sine data (NYS); 
Lyndonville, May 1889, Ge E. Fairman ENY? isotype D. 
accedens). IOWA: Ledges “State pParky (Boones Co. ;" 325° Jun 
Poo2eee os Barr’ 6940 (MASS). CALIFORNIA: Claremont, Los 
enewemaC peel? Mar, 1911), CC. Fi) Baker’.& Metz (NY on PL. S. 
PeeeGmtitaws562, Of *Thyridium ‘tuberculatum Rehm and Valsa 
ceanothi Rehm). 

Occasionally the ascospores may develop’ several 
additional septa, both transverse and longitudinal, although 
Ehese ‘septa do not become as dark or conspicuous as the 
original one. The anamorphic state is associated with the 
fereomorph in. the type collection as- well as in several 
Sener collections on Ulmus. [Ge cuLltcuLrerobarraoo lo produced 
slow-growing dark colonies in which conidiomata developed. 
Conidiogenesis was holoblastic. The conidia were brown, 
Meostiy sone septate, 10-12(-18) x’ 5-8 pm. On Magnolia and on 
Menispermum similar conidiomata are also present. 

Dothidotthia ramulicola seems to be a dark and septate- 


Spotea sm counterpart to Botryosphaeria obtusa. Bigg) | PAslekcas 
PEpecress ascomata “Show similar variations in size, -and in 
Poemiauron, of papilla, and ascospores are’ (18-)22-27(-34) x 
7-12{-15) pm, but remain hyaline and one celled. Conidia 


(Sphaeropsis sp., Shoemaker, 1964) are 22-26 x 10-12 pm, one 
celled or one septate and brown. 


I acknowledge with gratitude the curators of herbaria 
(BPI, NY, NYS, UBC) whose loans of specimens made this study 
Poeeetple. Dr. J: Boise reviewed the manuscript, and I thank 
her for her suggestions. 


LITERATURE CITED 


ee an A. VON: LOS, Revision einiger Gattungen der 
mecomyceten.. Acta, Bot. Neerlandica, 3: 83-93. 

Barr. M. —£, Ee iy te Preliminary studies on the Dothideales 
in temperate North America. Contr. Univ. Michigan 
We mom. 9523-638, 

weer --- , 1984. Herpotrichia and its segregates. Myco- 
fe Ore. 2 Ona 1 = 3 Be, 

Sie = = le = ‘ LN AY Pe Prodromus to Class Loculoascomycetes. 


Publ “by the author, Amherst, MA. USMS Hehe 
Eriksson, B. LOTAS On Ascomycetes on Diapensales and 


526 


Ericales in Fennoscandia. 2. Pyrenomycetes. Svensk 
Bot Mids kre eel 2a25 48 

Eviks sony Oe land fDyarh eel awks wom: OS Outline of the 
Ascomycetes - 19872. Syst. Ascomyc:)..62 }259S3 57% 

Hohne Lames von 1918. Dritte verld&ufige Mitteilung mycol- 
ogischer Ergebnisse (Nr. 201-304). Ber. Deutsch; Bot, 
Ges 36 sO ORB. 

--- eee e- : LoL Fragmente zur Mykologie. 1177. Uber 


Otthia Symphoricarpi (Ellis et Everhart). Sitzungsber. 
Kaiserl. Akad: Wiss. Math®-Naturwiss. Kl Abu ae 
592-596 

Holm) Ek LIGSe Taxonomic notes on Ascomycetes VI. On the 


genus Gibberidea Fuck. and some alleged relatives. 
svenskiybo tra Tjds kriG2ie) 7 ice le 

Miller, Ex.) and’) J. A. . von \Arx. (1962.0) Die) Gatttngenmeen 
didymosporen Pyrenomyceten. Beitr. Kryptogamenfl. 
Schweiz SLVc2)) sari 224 

Petrak, F., 1924.’ Mykologische: Notizen.: VII 7)0360)qmupen 
die Gattung, Dothidotthia'v.) Hohn., /Anne = Mycommeeae 
18 Ore Oe 

---------- .» 1927... Mykologiische Notizen. IX7S5760uameae 
Systremma spiraeae Murash. Ann. Mycol 925 e296=330e 

Shoemaker, R. A. 19 63 Generic correlations and concepts: 
Griphosphaerioma and Labridella. Canad... Jip Bo Gis ae 
1419-1423. 

ST ae ale lle ; 1964. Conidial states of some Botryosphaeria 
species,on Vitis and. Quercus, Canad. J). 2BO twee 
L201? 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 527-533 January-March 1989 


MARINE FUNGI FROM SEYCHELLES. VIII. RHIZOPHILA MARINA, A 
NEW ASCOMYCETE FROM MANGROVE PROP ROOTS 


Ree Detitly DE and Ghee...) ONES 


Sehool of Btologteal Setences, Portsmouth Polytechnie, 
king Henry I Street, Portsmouth, PO1 2DY, England. 


The number of marine fungi known from mangrove 
habitats has increased as many new species have 
recently been described (Kohlmeyer, 1984; Kohlmeyer & 
Schatz, 1985; Hyde & Borse, 1986a,b; Hyde & Jones, 1986; 
Kohlmeyer & Vittal, 1986). In this paper a new mangli- 
colous fungus from proproots and branches of Rhtzophora 
mucronata Lam. is described. Rhtzgophtla marina gen. et 
sp. nov. is characterised as having one-celled ascospores 
without appendages, unitunicate asci without an apical 
apparatus and large thick-walled ascocarps which are 
immersed in the substratum. 


RHIZOPHILA gen. nov. 


Ascocarptt grandii, globosi vel subglobosi, in ligno 
immersi, ostiolati, papillati, periphysati, coriacei, 
fusci ad nigri, solitarii vel gregarii. Cum paraphyses 
Aset octospori, clavati, pedunculati, unitunicati, lepto- 
dermi cum apice incrassatulo, sine apparatu apicali, ex 
base et lateribus ascocarporum muri evolventes. Asco- 
sporae 2-3 seriatae, ellipsoideae vel fusiformae, uni- 
cellulares, flavidae vel fulvae. Substratum in ligno 
mortuo mangrovis. Typus Rhtzophtla martna Hyde & Jones 
sp. nov. 


Ascocarps large, globose to subglobose, immersed in the 
substratum, ostiolate, papillate, periphysate, coriaceous, 
dark brown to black, solitary or gregarious. Paraphyses- 
like structures irregular in morphology, septate. Ascv 
8-spored, clavate, pedunculate, unitunicate, thin-walled 
with an indistinct apical thickening, without an apical 
apparatus, developing from ascogenous tissue lining the 
base and sides of the ascocarp wall. Ascospores 2-3 
seriate, ellipsoidal or fusiform, one-celled, yellowish to 


528 


yellowish brown. 
Substrate intertidal 
mangrove roots. Type 
spectes Rhtzophtla 
martna Hyde and Jones 
sp. nov. 

Etymology From the 
Latin Rhtzophtlus, 
meaning growing on 
TOOUS Fy alierelauLonsce 
the habitat ‘ofecne 
genus. 


RHIZOPHILA MARINA 
HYDE et JONES sp. nov. 


Ascocarptt 500-965 yum 
lati, 425-730 um longi, 
globosi vel subglobosi, 
immersi, ostiolati, 
papillati, periphysati, 
coriacei, fusci vel 
nigri, solitarii vel gregarii. Pertdtum 45-112 um 
crassum, 2-stratum, stratum externum, crassum, cum pig- 
mento nigro, cum hyphum irregularibusque; stratum 
internum, leptodernum, hyalinum, ex cellulis leptodermis 
elongatis irregularibusque compositum. Collum ad per 

515 um longum x 113-198 um latum, nigrum. Cum paraphyses. 
Aset 74-101 um longi x 13-24 um lati, octospori, clavati, 
pedunculati, unitunicati, leptodermi, subtruncati, sine 
apparatu apicali, ex base et lateribus ascocarporum muri 
orientes. Ascosporae 20-28 x 7-10 um, 2-3 seriatae, 
ellipsoideae ad fusiformes, unicellulares, hyalinae cum 
immaturae, flavidae ad fulvas cum maturae. Colonae in 
SW/CMA, albae, floccosae, celeriter crescentes, sine 
ascocarpis evolventibus. Substrata radices et rami 
Rhtzophorae mucronatae. 


RHIZOPHILA MARINA HYDE et JONES sp. nov. (Figs 1-17). 


Ascocarps 500-965 um in diameter, 425-730 um high, globose 
to subglobose, immersed, ostiolate, papillate, 
periphysate, coriaceous, dark brown to black, solitary or 
gregarious. Pertditum 45-112 um thick, 2-layered; the 


529 


Sigel).  hhizophila marina. Figs. 1-5. Diagrammatic 
Pentesentation of; 1. Cross section through ascocarp; 2. 
Peridium; 3. Paraphyses-like structures; 4. Ascospores and 
5. Ascus. Figs. 6-15, Light micrographs. 6-8. One-celled 
ascospores. 9-11. Asci at various stages of development. 
Note the truncate ascus tip in 11 and the paraphyses-like 
Senuecures in 9 and 10; Bar lines 1,°2:= 100m: 

3-11 = 10um. 


outer layer, thick, composed of a highly melanised layer 
of irregular hyphae interdispersed at the outside with the 
host cells, the inner layer, thin, hyaline, composed of 
irregular thin-walled elongate cells which fuse with the 
paraphyses-like structures. Necks up to 515um long x 113- 
198um in diameter, black, of similar structure to the asco- 
carp outer wall layer. Paraphyses-like structures, 3-llum 


530 


531 


12. T.S. Ascocarp; 13. Peridium. Note the inner wall layer 
(arrowed); 14, 15. Neck of ascocarp with periphyses. Figs. 
16, 17. S.E.M. of ascospores which have a granular wall. 
The mucilaginous material on the polycarbonate membrane 
surface may be the remains of the paraphyses-like struc- 
tures. Bar lines 12-15 = 100um, 16, 17 = 10 um. 


wide, consisting of irregular shaped cells, septate. Asct 
74-101 pm long x 13-24 um in diameter, 8-spored, 2-3 
serlate, clavate, pedunculate, unitunicate, thin-walled 
with an indistinct apical thickening, slightly truncate, 
without an apical apparatus, developing from ascogenous 
tissue lining the base and sides of the ascocarp wall. 
Ascospores 20-28 x 7-10 um, ellipsoidal to fusiform, one- 
celled, hyaline when immature, yellowish to yellowish- 
brown at maturity. Colontes on SW/CMA white, floccose, 
fast growing, no fruiting structures developing. Substrate 
prop roots and branches of Rhtzophora mucronata collected 
from Brillant mangrove, Seychelles. Dtstrtbutton Indian 
Ocean (Seychelles), South China Sea (Brunei), Straits of 
Malacca (North Sumatra, Thailand), Pacific Ocean (Bali). 


Holotype: Rhtzophila marina January 5th, 1984, from the 
Seychelles. Herb. IMI.325411, slides 1-10. 


Isotype: Dried wood from the Seychelles. 


Etymology: From the Latin marinus = marine, describing the 
habitat of the species. 


Other material examined: Intertidal roots of Rhizophora 
sp., Kpg. Serasa mangrove, Brunei, KDHOO01, KDHOO16, KDH 
0046; Intertidal prop roots of Rhtzophora sp., Kpg. 
Nelayan mangrove (near Belawan), North Sumatra, KDH0227; 
Intertidal prop roots of R. mucronata, Phang Nga Bay, 
Thailand, 14/1/88, KDH0O926; Intertidal prop roots of R. 
apteulata Blume, Kuta Beach mangrove, Bali, 5/2/88, 
KDH1LO69, 1010. 


Rhtzophtla marina cannot be assigned to any family of the 
Ascomycotina with any certainty, although it has affinities 
with the Phyllochoraceae and may well belong here. However, 
unlike members of this family, the ascocarps have thick 
walls and the asci lack a narrow apical annulus. Rhtzo- 
phitla shares many characteristics with the genus Glomerella 
Spauld & V. Schrenk, but differs in that the peridial wall 
in Glomerella is composed of angular cells, as opposed to 


532 


irregular hyphae, is one layered rather than two and the 
ascospores are hyaline as opposed to yellow or brown. 
Species of Glomerella also generally develop in leaf 
tissue and often under a stroma, but the extent of stroma- 
tic development varies (v. Arx & Muller, 1954). 


Rhizophtla may also be included in the Gnomoniaceae; 
however, the fungus has paraphyses-like structures and 
asci lack an apical ring-like structure, characteristics 
inconsistent with members of this family. 


Rhizophtla is characterised by having yellowish-brown one- 
celled ascospores without appendages, and clavate unituni- 
cate asci. Ascocarps are large, thick-walled, coriaceous, 
black, periphysate and with paraphyses-like structures. 
The fungus differs from any previous marine species known 
to us and because of its ubiquity in mangrove material is 
described as a new species. 


R. martna was collected in the Seychelles on 50 occasions 
from mangrove prop roots and branches collected at 
Brillant and Anse Boileau mangroves. It was noted as 
Ascomycete sp. (4) in previous publications (Hyde, 1985, 
1986; Hyde and Jones, 1988) and was regarded as a common 
species in the mangrove ecosystem. It has also been 
collected from Kpg. Serasa mangrove, Brunei, Kpg. Nelayan 
mangrove, North Sumatra, Phang Nga Bay, Thailand and Kuta 
Beach mangrove, Bali, Indonesia. It has been identified 
from Rhizophora mucronata and Rhtzophora aptculata prop 
roots. R&. martna was identified on material that was sub- 
merged during high tide and was associated with marine 
borers, barnacles, seaweeds and other marine fungi (i.e. 
Lulworthta grandispora Meyers, Ketsslertella blepharo- 
spora Kohlmeyer & Kohlmeyer). 


ACKNOWLEDGEMENTS 


Thanks are extended to Mr. C. Derrick of Portsmouth Poly- 
technic for photographic assistance; to Mrs. Monica Heywood 
Kenny for help with the Latin diagnosis and Professor J. 
Kohlmeyer for reviewing the manuscript. 


LITERATURE CITED 


von ARX, J.A. and MULLER, E. 1954. Die Gattungen der 
amerosporen Pyrenomyceten. Beitrage Krypt. Schweiz, 
Bande Hetty pp 7434. 


533 


HYDE, K.D. 1985. Spore settlement and attachment in 
marine fungi. Ph.D. Thesis (CNAA) Portsmouth 
Polytechnic. 


HYDE, K.D. 1986. Frequency of occurrence of lignicolous 
marine fungi in the tropics. In: The Biology of 
Marine Fungi, (Ed. S.T. Moss), pp. 311-322, Cambridge 
University Press, Cambridge. 


HYDE, K.D. and BORSE, B.D. 1986a. Marine fungi from 
Seychelles. V. Btatrtospora marina gen. et sp. nov. 
from mangrove wood. Mycotaxon 26: 263-270. 


HYDE, K.D. and BORSE, B.D. 1986b. Marine fungi from 
Seychelles. VI. Massarina velataspora, a new 
ascomycete from mangrove wood. Mycotaxon 27: 161-167. 


HYDE, K.D. and JONES, E.B.G. 1986. Marine fungi from 
Seychelles. IV. Cucullospora mangrovet from dead 
mangrove wood. Bot. Mar. 26: 491-495, 


HYDE, K.D. and JONES, E.B.G. 1988. Marine mangrove fungi. 
Marine Ecology (PSZNI) 9: 15-33. 


KOHLMEYER, J. 1984. Tropical marine fungi. Marine Ecology 
(PSZNI) 5: 329-378. 


KOHLMEYER, J. and SCHATZ, S. 1985. Atgtalus gen. nov. 
(Ascomycetes) with two new marine species from 
Meameroves. —Trans. Br. mycol. Soc. 85: 699-707. 


KOHLMEYER, J. and VITTAL, B.P.R. 1986. Lophtostoma 
mangrovts, a new marine ascomycete from the Eropicsr 
Mycologia 78: 485-489, 


KOHLMEYER, J. and VOLKMANN-KOHLMEYER, B. 1987. Marine 
fungi from Belize with a description of two genera 
of ascomycetes. Bot. Mar. 30: 195-204. 


! 


f 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 535-540 January-March 1989 


MARINE FUNGI FROM INDIA. III. ACROCORDIOPSIS PATILII 
GEN. ET. SP. NOV. FROM MANGROVE WOOD 


B.D. BORSE! and K.D. HYDE 


lDepartment of Botany, Arts, Commerce and Sctence College, 
Erandol 425109, Indta. 


2school of Btologteal Setences, Portsmouth Polytechnic, 
King Henry I Street, Portsmouth, PO1 2DY, England. 


During investigations into the mangrove inhabiting 
fungi from the coast of Maharashtra (India) and Brunei, an 
ascomycete with large black conical ascocarps was encoun- 
tered. A new genus is described to accommodate this 
fungus which cannot be assigned to any genera of the 
Ascomycotina. 


ACROCORDIOPSIS gen. nov. 


Ascocarpiis concis vel semiglobosis, superficialibus, 
solitariis vel gregariis, nigris, ostiolatis, carbonaceis. 
Pseudoparaphysibus numerosis, filiformibus, persistenti- 
bus, anastomosantibus, septatis. Asci octosporis, bituni- 
catis, cylindricis, pedunculatis. Ascosporis hyalinis, 
uniseptatis constrictus, ad septis, ovoideis vel ellip- 
soideis, guttulatis. Typus generis: Acrocordtopsts 
pattltt Borse et Hyde. 


Ascocarps conical or semiglobose, superficial, solitary or 
gregarious, black, ostiolate, carbonaceous. Pseudopara- 
physes numerous, filiform, persistent, anastomosing, sep- 
tate. Asci eight-spored, bitunicate, cylindrical, 
pedunculate, with ocular chamber. Ascospores hyaline or 
slightly yellow, one-septate, constricted at the septum, 
obovoidal or ellipsoidal, guttulate. Type species: 
Acrocordtopsts patitltt Borse et Hyde. 


Etymology: Acrocordtopsts in reference to the congruity 
with the genus Acrocordta Massal. 


536 


ACROCORDIOPSIS PATILII sp. nov. (Figs 1-10) 


Ascocarpiis 1-3mm latis, 1-2mm longis, conicis vel 
semiglobis, superficialibus, solitariis vel gregariis, 
nigris, ostiolatis, carbonaceis. Peridiis 240-350um 
crassis. Pseudoparaphysibus hyalinis, filiformibus, 
persistentibus, numerosis, septatis, anastomosantibus. 
Ascis 150-250um longis, 12-l6um latis, octosporis, 
bitunicatis, cylindricis, pedunculatis. Ascosporis 
16-25um longis, 10-léyum latis, hyalinis, uniseptatis, 
constrictis ad septis, ovoideis vel ellipsoideis, 
guttulatis. Substratum lignum mortuum Avtcenntae albae 
Bl. et Rhtzophorae mucronatae Lam. Holotypus: I.M.I. 
DIOS, 


Ascocarps 1-3mm in diameter x 1-2mm high, conical or semi- 
globose, superficial, solitary or gregarious, black, 
ostiolate, carbonaceous. Peridium 250-350um thick, two 
layered; outer layer 240-338um, thick, black, pseudo- 
parenchymatous, composed of thick-walled black cells; 

inner layer 9-14yum thick, hyaline-light brown, merging 
with pseudoparaphyses, composed of irregular elongate thin- 
walled cells. A third area of rectangular thick-walled 
cells, brown in colour, fills the angle between the cover 
and basis of the ascoma. Pseudoparaphyses 0.5-1.5um, 
hyaline, filiform, persistent, numerous, anastomosing and 
sparsely septate. Asci 150-250 x 12-l6um, eightwenoreds 
bitunicate, cylindrical, pedunculate, with an apically 
thickened refractive region and an ocular chamber, develop- 
ing from base and corners of the ascocarp. Ascospores 
16-25 x 10-l6um, hyaline, one septate, constricted at the 
septum, ovoidal or ellipsoidal, one large guttule per cell. 
Substratum on intertidal wood of Avtcennta alba Bl. and 
Rhtzophora mucrobata Lam. 


Figs 1-10. Acrocordtopsts patili1t: light micropraphse ais 
2. Conical ascocarps with released spore mass (arrowed). 
3. Section through ascocarp. The wall is black, 
carbonaceous and individual cells are indistinguishable in 
mature specimens. A pallisade-like area of parallel cells 
is present at the rim. 4..Gregarious ascocarps meee 
Squash showing asci, filiform pseudoparaphyses and asco- 
spores. Note the ocular chamber of the ascus and 
refractive apical thickening (5 arrowed). 7-10. Asco- 
spores. Bars: Figs. 1,2,4 = lmm; Fig. 4 = 500um; Figs. 
5-10 = 10um. 


537 


538 


Holotype: collected from Malvan (Maharashtra), Indian 
Ocean (India), 30.10.1981 on intertidal wood of Avicennta 
apa Bl el M orem 97 769% 


Specimens examined: MFM No. 58, 30.4.1982, on intertidal 
wood of Aviecennta alba; MFM No. 59, Bankot, 29.3.1983, on 
intertidal wood of Rhtzophora mucronata Lam; MRM No. 60, 
Malvan, 18.4.1983, on intertidal wood of Avtcennta alba; 
KDH 0071, October 1985, on mangrove wood, Brunel. 


Attempts to locate a suitable genus to accommodate this 
mangrove species were unsuccessful and therefore a new 
genus Acrocordtopsts is formed to accommodate this taxon. 
Acrocordtopsts is characterized by having large conical 
superficial ascocarps (Figs 1-4), with a thick 
carbonaceous wall, narrow anastomosing hamathecial tissue 
(Figs 5,6) and cylindrical asci with an ocular chamber and 
a refractive apical thickening (Figs 5,6). Ascospores are 
hyaline, symmetrical, bi-celled and without appendages 
(Figs 7-10). These characteristics indicate that 
Acrocordtopsts can be placed in the Melanommatales (sensu 
Barr, 1983), Dothideales or Pyrenulales (sensu Hawksworth 
and Eriksson, 1986). 


Acrocordtopsts may belong in the Pyrenulaceae 
(Pyrenulales) where it has affinities with Acrocordta 
Massal and Acrocordtella 0. Eriksson (Requinella Fabre?). 
However, Acrocordta is a lichen forming genus and the asci 
are capped by a meniscus staining in Congo Red (Eriksson, 
1981). The meniscus does not occur in Acrocordtopsts, 
although the tip of the ascus is thickened and refractive. 
In Acrocordtella (bark saprophyte), the ascospores are 
multiseptate with lenticular locules, the asci also have a 
meniscus (manubrium), while the ascocarps are immersed 
(Eriksson, 1982). In Acrocordtopsts, the ascocarp wall 
structure is also different from the above species. 


Acrocordtopsts also has affinities with the genus 
Pertdtothelta Hawksw. (Phaeosphaeriaceae), a non-lichenized 
saprophytic (?) genus (Hawksworth, 1985). However, 
Acrocordtopsts cannot be accommodated here as in 
Pertdothelta, the pseudoparaphyses are cellular and blue in 
iodine, the ascocarps are smaller and the ascospores are 
brown (Hawksworth, 1985). 


Acrocordtopsts is probably best included in the 
Melannommataceae (Melanommatales - sensu Barr, 1983; 


539 


Dothideales - sensu Hawksworth and Eriksson, 1986), since 
the ascocarps are ostiolate and possess a well developed 
hamathecium of the narrow, branched and anastomosing 
sparsely septate trabeculate type. It has affinities with 
the genus Astrosphaertella Sydow (Hawksworth, 1981) 
although ascospores in these genera differ greatly. In 
Aerocordtopsts the ascocarp wall is composed of thick- 
walled black cells which are indistinguishable in mature 
specimens, while at the rim an area of pallisade-like rows 
of parallel cells is present. This type of wall structure 
is also found in Astrosphaertella, while the ascocarps are 
also conical. However, ascospores differ in 
Astrosphaertella as they are elongate-fusiform, usually 
brown or red-brown and surrounded by a gelatinous sheath 
when young (Hawksworth, 1981). Furthermore 
Astrosphaertella species develop on monocotyledon 
petioles. 


Aerocordtopsts pattltt was collected in Brunei and India 
in the upper intertidal region of mangroves. The asco- 
carps are large (1-3mm high x 1-2mm long) compared to most 
fungi from marine or brackish water habitats; other marine 
species with large superficial ascocarps include 
Rosellinta sp. (undescribed ascomycete) and Caryosporella 
rhizophorae Kohlmeyer, and both species are known from 
mangrove habitats. It is also interesting to note that 
many very large spored ascomycetes have been collected 
from the mangrove habitat, which is an unusual feature for 
Marine ascomycetes. Atgtalus grandts Kohlm., A. parvus 
Kohlm., Cuecullospora mangrovet Hyde and Jones and 
Halosarpheta ratnagirtensts Patil and Borse all have asco- 
spores with dimensions above 55um. Lulworthta grandtspora 
is the largest Lulworthia sp. and this is also found in 
mangrove habitats. 


ACKNOWLEDGEMENTS 


B.D. Borse is much indebted to Professor S.D. Patil 
for the use of laboratory facilities and for his 
encouragement. We are thankful to C. Derrick and 
R. Mouzouras for photographic assistance and to Professor 
E.B.G. Jones for his support. We are also grateful to 
Professor D.L. Hawksworth for his general advice with 
respect to the ideas presented in this paper, and for 
reviewing the manuscript. 


540 


LITERATURE CITED 


BARR, M.E. 1983. The Ascomycete connection. Mycologia, 75: 
Bld Shes 


ERIKSSON, O. 1981. The families of bitunicate ascomycetes. 
Opera’ Botanica, ows 1-270), 


ERIKSSON, O. 1982. Notes on ascomycetes and coelomycetes 
from’ N.W. Europe. Mycotaxon, 15%7 189-2028 


HAWKSWORTH, D.L. 1981. Astrosphaertella Sydow, a 
misunderstood genus of Menanommataceous pyrenomycetes. 
BOY Rie elal i O0C . moe emo. 


HAWKSWORTH, D.L. 1985. A redisposition of the species 
referred to the ascomycete genus Microthelta. Bull. 
Brea Mus, nat. Hist.. (BOty) . 04 eo tore 


HAWKSWORTH, D.L. and ERIKSSON, O. 1986. The names of 
accepted orders of ascomycetes. Systema Ascomycetum, 
Dee Log. 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 541-564 January-March 1989 


NEW SPECIES IN THE LICHEN GENUS XANTHOPARMELIA 
(ASCOMYCOTINA: PARMELIACEAE) 


Mason E. Hale 


Department of Botany, Smithsonian Institution 
Washington, DC 20560, U.S.A. 


Abstract: The following new species in the lichen genus Xanthoparmelia from 
North and South America, Australia, Lesotho, South West Africa/Namibia, and 
South Africa are described as new: X. afrolavicola Hale, X. amplexuloides Hale, 
X. ausiana Hale, X. catarinae Hale, X. cirrhomedullosa Hale, X. coneruptens Hale, 
X. conjuncta Hale, X. granulata Hale, X. harrisii Hale, X. idahoensis Hale, X. 
imbricata Hale, X. inflata Hale, X. khomasiana Hale, X. kotisephola Hale, X. 
lipochlorochroa Hale & Elix, X. luderitziana Hale, X. mapholanengensis Hale, X. 
maxima Hale, X. neowyomingensis Hale, X. norcolorata Hale, XxX. 
norlobaronica Hale, X. norwalteri Hale, X. protolusitana Hale, X. protoquintaria Hale, 
X. psornorstictica Hale, X. pustulosorediata Hale, X. salamphixantha Hale, X. 
saleruptens Hale, X. salkiboensis Hale, X. springbokensis Hale, xX. 
subamplexuloides Hale, X. subbullata Hale, X. subconvoluta Hale, X. subluminosa 
Hale, X. submougeotii Hale, X. substenophylloides Hale, and X. tsekensis Hale. 


Xanthoparmelia afrolavicola Hale, sp. nov. (Figs. 1, 2) 

Thallus laxe adnatus, fragilis, 5-7 cm latus, lobis sublinearibus, 1.5—4 mm latis, superne 
continuus, emaculatus, isidiatus, isidiis cylindricis, subtus pallide brunneus, sparse rhizinosus. 

Thallus loosely adnate on rock, rather brittle and easily breaking apart, 5-7 cm broad, darkish 
yellow green; lobes subirregular to sublinear, 1.5-4 mm wide, irregularly dichotomously branched, 
subimbricate; upper surface continuous to faintly white maculate in patches, shiny, moderately to 
densely isidiate, the isidia cylindrical, thin, 0.06-0.08 mm in diameter, 0.1—0.4 mm high, black 
tipped, unbranched or sparingly branched (fig. 2); lower surface plane, light brown, sparsely 
rhizinate, the rhizines pale brown, coarse, simple, 0.2-0.5 mm long. Pycnidia common; conidia 
cylindrical, 0.5 x 5-6 um. Apothecia lacking. 

Chemistry: psoromic, norpsoromic, usnic acids. 

Type collection: Natal Table Mountain near Pietermaritzburg, elev. 650 m, Natal Province, South 
Africa, Grid 2930 DA, Hale 74104 (US, holotype; PRE, isotype). 

Additional specimens examined. S.W.A./Namibia: Waterberg Plateau National Park (E of 
Otjiwarongo), Grid 2017 AC, Hale 80923, 80984, 81048, 81058 (US). . 

This rare species is related to the American psoromic acid-containing X. lavicola (Gyel.) Hale, 
which has globose, partially erumpent isidia and a darker, moderately rhizinate lower surface. 


Xanthoparmelia amplexuloides Hale, sp. nov. (Figs. 3,4) 

Thallus adnatus, saxicola, 5—8 cm latus, lobis subirregularibus, 1.2.5 mm latis, superne continuus, 
-emaculatus, modice isidiatus, isidiis subglobosis, subtus planus, brunneus, modice rhizinosus. 

Thallus adnate, rather soft and fragile, 5-8 cm broad, light yellowish green to yellowish green; 
lobes subirregular, 1-2.5 mm wide, apically subrotund and irregularly branched, imbricate; upper 
surface continuous, shiny, rugose with age, moderately to densely isidiate, the isidia initially 
subglobose, subcylindrical at maturity, 0.08-0.12 mm in diameter, to 0.5 mm high, tips syncorticate 
to weakly epicorticate, pale, very rarely erumpent or breaking off, unbranched (fig. 4); lower surface 


542 


Figures 1-4. Species of Xanthoparmelia: 1, X. afrolavicola (Hale 74104, holotype); 2, SEM 


enlargement of isidia of X. afrolavicola; 3, X. amplexuloides (Hale 78874, holotype); 4, SEM 
enlargement of isidia of X. amplexuloides. Scale in mm. 


543 


plane, pale brown, moderately rhizinate, the rhizines pale brown, simple, 0.2—0.4 mm long. Pycnidia 
lacking. Apothecia substipitate, 1-2 mm in diameter; spores S—6 x 9-11 um. 

Type collection: 38 km W of Bloemfontein on north side of Hwy R64, elev. 1500 m, Orange Free 
State, South Africa, Grid 2926 BA, Hale 78874, 3 Nov. 1986 (US, holotype; PRE, isotype). 

Chemistry: norlobaridone, unknown Rfc 12, and usnic acid (det. J. A. Elix). 

Additional specimens examined. South Africa. Transvaal: Hanglip Forest Reserve, Soutpansberg 
Mountains, Grid 2229 DD, Hale 76373 (US); Breedtsnek, Magaliesberg, Grid 2527 CD, Hale 80871, 
80876 (US); Long Tom Pass, Grid 2530 BA, Hale 76070 (US). Orange Free State: 46.9 km S of 
Wepener, Grid 3027 AA, Hale 73483, 73484 (US); 38.5 km S of Standerton, Hale 73481 (US). Cape 
Province: 33.8 km W of Dordrecht, Grid 3126 BC, Hale 73480, 73482 (US); Penhoek Pass on Hwy 
R30, Grid 3126 BC, Hale 80558 (US); 20 km WSW of Elliott, Grid 3127 BC, Hale 72531 (US); 27.6 
km WNW of Elliott, Grid 3127 BA, Hale 77087 (US); 17.9 km NE of Maclear, Grid 3028 CD, Hale 
76891 (US); 37.4 km NW of Queenstown, Grid 3126 AC, Hale 72530 (US); Karoo Nature Reserve, 
Grid 3224 BC, Hale 73479 (US). Natal: Mzintlava River crossing on Hwy R394, 22km N of Kokstad, 
Grid 3029 AD, Hale 81316 (US); Giants Castle Game Reserve, Grid 2929 BC, Hale 80811 (US). 
Lesotho: 6 km S of Sehlabathebe National Park, Grid 2929 CC, Hale 81347, 81349, 81354 (US); 9 km 
W of Ramatsiliso Nek, Grid 3028 BB, Hale 81354 (US). 

The Australian X. amplexula is a more robust, leathery lichen with branched, tall, black-tipped 
cylindrical isidia and usually contains loxodin along with norlobaridone and rarely the scabrosin 
complex. Xanthoparmelia scabrosa (Tayl.) Hale, common in Australasia and Japan, has distinctly 
pustulate isidia. Another isidiate, norlobaridone-containing species in $.W.A./Namibia, X. 
subamplexuloides Hale (below), is tightly adnate with narrow lobes, 0.4-1.3 mm wide. 


Xanthoparmelia ausiana Hale, sp. nov. (Fig. 5) 

Thallus arcte adnatus, saxicola, 4-8 cm latus, lobis subirregularibus, 1-2 mm latis, superne 
continuus, emaculatus, isidiis sorediisque destitutus, subtus pallide brunneus, modice rhizinosus. 

Thallus tightly adnate to adnate on rock, somewhat dispersed, 4-8 cm broad, light yellow green; 
lobes subirregular, 1-2 mm wide, short and irregularly branched, separate to subimbricate; upper 
surface continuous to faintly maculate in patches, shiny, rugulose and transversely cracked with age, 
isidia and soredia lacking; lower surface plane, pale brown, moderately rhizinate, the rhizines pale 
brown, simple, 0.3-0.5 mm long. Pycnidia commonly developed; conidia bifusiform, 0.5 x 5-6 um. 
Apothecia lacking. 

Chemistry: norstictic, salazinic and usnic acids. 

Type: On sheltered granite ledge, 5 km NE of Aus on hwy 35 (road to Helmeringshausen), 4.7 km 
N of Hwy B4, elevation 1300 m, South West Africa/Namibia, Grid 2616 CB, Hale 81143, 27 April 
1988 (US, holotype; PRE, isotype). 

The thallus color and lobe configuration remind one of X. colorata (Mill. Arg.) Hale, a much 
larger lichen with the same chemistry but a black lower surface. 


Xanthoparmelia catarinae Hale, sp. nov. (Figs. 6, 7) 

Thallus laxe adnatus, saxicola, 5—8 cm latus, lobis sublinearibus, 0.8-2 mm latis, superne modice 
isidiatus, isidiis subglobosis vel irregulariter cylindricis, subtus niger, sparse rhizinosus. 

Thallus loosely adnate on rock, brittle, 5—8 cm broad, yellowish green; lobes sublinear, 0.8-2 mm 
wide, dichotomously branched, imbricate; upper surface continuous, shiny, sparsely to moderately 
isidiate, the isidia initially subglobose, irregularly cylindrical at maturity, 0.14-0.18 mm in diameter, 
0.2-0.8 mm high, mostly simple to sparsely coralloid branched (fig. 7); lower surface plane to 
weakly convoluted at the tips, black with a barely raised dark to yellowish rim, shiny, sparsely 
rhizinate, the rhizines black, coarse, unbranched, 0.2-0.5 mm long. Pycnidia common; conidia 
bifusiform, 0.5 x 5-6 um. Apothecia lacking. 

Chemistry: stictic, constictic, norstictic, cryptostictic (tr.) and usnic acids. 

Type collection: Florianopolis-Estreito, Santa Catarina, Brazil, Poeschmann-Hajkova in Vezda, 
_Lichenes Selecti Exsiccati 761 (US, holotype). 

Additional specimen examined. South America. Brazil. Sao Sepe, Passo dos Freire, Rio Grande 
do Sul, Fleig ICN 55973 (US). 

This is an isidiate species related to X. hypopsila (Mill. Arg.) Hale, a common nonisidiate 
lichen in Argentina. It is differentiated from North American X. isidiascens by the subconvolute 
lobe tips, narrower lobes, and sparse rhizines. 


544 


BHO4h Laky 


Figures 5-8. Species of Xanthoparmelia: 5, X. ausiana (Hale 81143, holotype); 6, X. catarinae 


(Poeschmann-Hajkova, holotype); 7, SEM enlargement of isidia of X. catarinae; 8, X. 
cirrhomedullosa (Hale 80605, holotype). Scale in mm. 


—— 
Bal 
- 
Saad 
— 
— 
-= 
~ 
~ 
sell 


545 


Xanthoparmelia cirrhomedullosa Hale, sp. nov. (Fig. 8) 

Thallus adnatus, saxicola, 5-7 cm latus, lobis subirregularibus, 1-4 mm latis, superne albo- 
maculatus, rugosus, isidiis sorediisque destitutus, medulla pro parte ochracea, subtus brunneus, 
modice rhizinosus. 

Thallus adnate on sheltered rock, leathery, 5—7 cm broad, darkish yellow green; lobes 
subirregular with rotund tips, 1-4 mm wide, short, contiguous at the margin, crowded at the center; 
upper surface uniformly white maculate, shiny, strongly rugose with age; isidia and soredia lacking; 
lower surface plane, brown or faintly orange, smooth to rugulose, sparsely to moderately rhizinate, 
the rhizines brown, 0.5-1 mm long. Pycnidia common; conidia bifusiform, 0.5 x 5-6 wm. Apothecia 
substipitate, 2-4 mm in diameter. 

Chemistry: salazinic and usnic acids, skyrin and the chalybaeizans unknown. 

Type collection: Large S-facing Cave sandstone ledges, Greyling Pass on W side of Hwy R396, 9 
km N of Rossouw, elev. 2100 m, Cape Province, South Africa, Grid 3127 AB, Hale 80605, 6 April 
1988 (US, holotype; PRE, isotype). 

The white maculate surface and medullary pigment skyrin are found only in this species and in X. 
subcolorata Hale, a large, sublinear-lobed, loosely adnate lichen with an eroding lower surface. It is 
known only from the type collection. 


Xanthoparmelia coneruptens Hale, sp. nov. (Fig. 9) 

Thallus arcte adnatus, saxicola, 3-5 cm latus, lobis subirregularibus, 0.5—-1 mm latis, superne 
continuus vel vix reticulato-maculatus, pustulatus, pustulis eruptentibus, crasse sorediatis, subtus 
planus, pallide brunneus, modice rhizinosus. 

Thallus tightly adnate on rock, light yellowish green, 3-5 cm broad; lobes subirregular, 0.4-1 mm 
wide, short and crowded, imbricate; upper surface continuous to faintly reticulate-maculate, shiny to 
dull white pruinose with age, transversely cracked, pustulate-isidiate, the pustules entire or erupting, 
breaking open and become coarsely sorediate; lower surface plane, pale brown, moderately 
rhizinate, the rhizines pale brown, simple, ca 2 mm long. Pycnidia and apothecia lacking. 

Chemistry: lecanoric and usnic acids. 

Type collection: On overhanging sandstone ledge, on Hwy R364, 32.4 km S of jet with R27, elev. 
600 m, Cape Province, South Africa, Grid 3119 CD, Hale 79859, 24 March 1988 (US, holotype; PRE, 
isotype). 

Additional specimens examined. South Africa. Cape Province: same locality as the holotype, Hale 
79854, 79855, 79859 (US); 7 km W of Oliewenboskraal (W of Hwy N7), Grid 3218 BD, Hale 79906 
(US); 18 km E of Aggeneys on Hwy R64, Grid 2918 BB, Hale 79474, 79475 (US). 

This is a chemotype of X. eruptens Hale, in fact the commonest member of this pustulate-isidiate 
group, typically found on the underside of large overhanging sandstone ledges. 


Xanthoparmelia conjuncta Hale, sp. nov. (Fig. 10) 

Thallus arcte adnatus, saxicola, coriaceus, 4-6 cm latus, lobis subirregularibus, 0.7-1.5 mm latis, 
superne continuus, emaculatus, isidiis sorediisque destitutus, subtus planus, brunneus, modice 
rhizinosus. 

Thallus tightly adnate to adnate on rock, leathery, 4-6 cm broad, dark yellowish green; lobes 
subirregular, 0.7-1.5 mm wide, irregularly branched, contiguous to subimbricate; upper surface 
continuous, transversely cracked with age, isidia and soredia lacking; lower surface plane, brown, 
moderately rhizinate, the rhizines brown, simple, 0.2—0.4 mm long. Apothecia adnate, 1-3 mm in 
diameter. Pycnidia common; conidia bifusiform, 0.5 x 4-6 um. 

Chemistry: diffractaic (major), 4-O-demethyldiffractaic, barbatic (tr.), 4-O-demethylbarbatic (tr.), 
and usnic acids. 

Type collection: On small dolerite boulders in meadow, 2.6 km NW of Sani Pass, elev. 2900 m, 
Lesotho, Grid 2929 DB, Hale 81327, 6 May 1988 (US, holotype; PRE, isotype). 

This rare high elevation species is externally similar to X. lesothoensis Hale, which contains both 
diffractaic and salazinic acids but lacks 4-O-demethyldiffractaic acid. 


Xanthoparmelia granulata Hale, sp. nov. (Fig. 11) 

Thallus adnatus, saxicola, fragilis, 4-6 cm latus, lobis subirregularibus, 0.8-1.3 mm latus, superne 
continuus, emaculatus, valde rugosus, pustulatus, pustulis eruptentibus, crasse sorediatis, subtus 
planus, pallide brunneus, sparse rhizinosus. 

Thallus adnate on rock (removed free when wetted), fragile and breaking into pieces, dull 


546 


“eescin 
weete 
sSeaaee 
— 
ses 
ome 
sama 
a 
bead 


Figures 9-12. Species of Xanthoparmelia: 9, 


X. conjuncta (Hale 81327, holotype); 11, X. 
(Hale 81203, holotype). Scale in mm. 


X. coneruptens (Hale 79859, holotype (x10)); 10, 
granulata (Hale 81540, holotype); 12, X. harrisii 


547 


yellowish green, 4—6 cm broad; lobes subirregular, 0.8- 1.3 mm wide, black rimmed, subimbricate; 
upper surface continuous, dull to faintly pruinose, soon strongly rugose, the wrinkles becoming 
irregularly pustulate, erupting and forming coarse soredia; lower surface plane, pale brown, sparsely 
to moderately rhizinate, the rhizines simple, 0.3-0.5 mm long. Pycnidia and apothecia lacking. 

Chemistry: fumarprotocetraric, succinprotocetraric, and usnic acids. 

Type collection: On sheltered dolerite ledge, 56.1 km N of Mapholaneng on Sani Pass-Moteng 
Pass road, elev. ca 3000 m, Lesotho, Grid 2828 DD, Hale 81540, 7 May 1988 (US, holotype; PRE, 
isotype). 

The granular pustules are vaguely similar to those of Flavoparmelia caperata (L.) Hale, but the 
narrow lobes and pale lower surface are typical of Xanthoparmelia. It is known only from the type 
collection at high elevation in Lesotho. 


Xanthoparmelia harrisii Hale, sp. nov. (Fig. 12) 

Thallus arcte adnatus, saxicola, 1-2 cm latus, lobis sublinearibus, 0.5—1 mm latis, superne sparse 
isidiatus, isidiis cylindricis, subtus pallide brunneus, sparse rhizinosus 

Thallus tightly adnate on quartz pebbles, light yellow green at the tips but darkening to blackish 
brown at the center, 1-2 cm broad; lobes sublinear, 0.5—1 mm wide, convex, little branched, 
separate to contiguous; upper surface continuous, dull and light pruinose, sparsely to densely 
isidiate, the isidia coarse, cylindrical, 0.1—0.2 mm in diameter, to 2 mm high, the tips syncorticate, 
becoming richly branched; lower surface plane, pale brown, sparsely rhizinate, the rhizines short, 
coarse, 0.1-0.2 mm long, unbranched. Pycnidia and apothecia lacking. 

Chemistry: hypostictic, hyposalazinic, hypoconstictic, caperatic (+) and usnic acids, associated 
undetermined spots. 

Type collection: Gravel flats, 18.2 km NE of Cape Cross on E side of Hwy D2301, elev. 100 m, 
S.W.A./Namibia, Grid 2113 DB, Hale 81203, 22 April 1988 (US, holotype; LD, PRE, isotypes). 

Additional specimens examined. $.W.A./Namibia: East of Wlotzska’s Baken off Hwy D2301, 
Swakopmund District, Grid 2214 AD, Hale 80946 (US), R. Harris 18513 (NY); 33.1 km NE Cape 
Cross at "Pink Mountain," Grid 2114 CA, Hale 81023 (US); 

Comments: This widespread, easily overlooked Namib desert lichen grows best on the protected 
lower edges of quartzite pebbles. The center is almost black but the marginal lobes a bright yellow. 
It is unique among the species with the hypostictic acid series and confined to the coastal fog zone in 
Namibia. It is named in honor of Dr. Richard Harris, who first collected it. 


Xanthoparmelia idahoensis Hale, sp. nov. (Fig. 13) 

Thallus vagans, terricola, coriaceus, 2-4 cm latus, lobis sublinearibus, 1.5—4 mm latis, contortis, 
subterete laciniatus, superne valde albo-maculatus, sorediis isidiisque destitutus, subtus 
canaliculatus, sparsissime rhizinosus. 

Thallus vagrant, free growing on soil, firm, breaking apart when collected, 2-4 cm broad, light 
yellowish green; lobes sublinear, 1.5-4 mm wide, contorted and twisted, the tips irregularly dilated 
and divided into subterete black-tipped laciniae ca 0.5 mm wide, separate; upper surface strongly 
white maculate, shiny, soredia and isidia lacking; lower surface irregularly canaliculate with a raised 
rim, pale yellowish brown or turning brown toward the tips, foveolate-rugose, very sparsely rhizinate, 
the rhizines brown, coarse, 0.2—0.3 mm long, simple. Pycnidia and apothecia lacking. 

Chemistry: salazinic, consalazinic, protocetraric (tr.), and usnic acids. 

Type collection: On calcareous lacustrine ash soil, SE of Salmon, Lemhi County, Idaho, T 21 N, R 
22 E, sect. 28, R. Rosentreter 3828, 4 Jan. 1986 (US, holotype; Rosentreter herbarium, isotype). 

This rare soil lichen with contorted lobes is unique in the genus. The waxy-appearing upper 
surface is strongly maculate in patches. It almost seems like an Evernia, but with the scanning 
electron microscope I was able to find epicorticate areas. 


Xanthoparmelia imbricata Hale, sp. nov. (Fig. 14) 

Thallus laxe adnatus, saxicola, subpulvinatus, fragilis, 8-10 cm latus, lobis subirregularibus, 1-2 
mm latis, apice digitato-laciniatis, superne continuus, emaculatus, isidiis sorediisque destitutus, 
subtus planus, nigricans, sparsissime rhizinosus. 

Thallus loosely adnate on rock, subpulvinate, rather fragile and breaking apart, 8-10 cm broad, 
light yellowish green; lobes subirregular, 1-2 mm wide, short with digitate-laciniate branched tips, 
the tips black rimmed and often suberect and weakly subterete, strongly imbricated; upper surface 
continuous, shiny, isidia and soredia lacking; lower surface plane, dark brown at the tips with a 


548 


Figures 13-16. Species of Xanthoparmelia: 13, X. idahoensis (Rosentreter 3828, holotype); 14 


X. imbricata (Hale 81541, holotype); 15, X. inflata (Hale 81531, holotype); 16, X. khomasiana 
(Hale 80955, holotype (x10)). Scale in mm. 


’ 


549 


yellowish rim, blackening at the center, very sparsely rhizinate, the rhizines black, simple, 0.4-0.8 
mm long. Pycnidia numerous; conidia subbifusiform, 0.5 x 5-6 um. Apothecia lacking. 

Chemistry: unknown fatty acids Rfc 33 and 37 (in X. subdecipiens), constipatic (tr.), 
protoconstipatic (tr.), and usnic acids. 

Type collection: On sheltered dolerite ledges on hillside, 56.1 km N of Mapholaneng on Sani Pass- 
Moteng Pass road, elev. ca 3000 m, Lesotho, Grid 2828 DD, Hale 81541, 7 May 1988 (US, holotype; 
PRE, isotype). 

Additional specimen examined. Lesotho: 7.5 km NW of Sani Pass, Grid 2929 CB, Hale 81529 
(US). 

The digitate-laciniate lobes and chemistry set this species apart from other fatty acid-containing 
species. 


Xanthoparmelia inflata Hale, sp. nov. (Fig. 15) 

Thallus adnatus, saxicola, coriaceus, 5-10 cm latus, lobis late sublinearibus, 2-3 mm latis, 
convexis, superne continuus, emaculatus, isidiis sorediisque destitutus, subtus planus, dense 
rhizinosus. 

Thallus adnate to loosely adnate on rock, rather leathery and appearing inflated, 5-10 cm broad, 
light yellowish green; lobes broadly sublinear, 2~3 mm wide, little branched, separate to contiguous; 
upper surface continuous, dull, rugose at the center, isidia and soredia lacking; lower surface plane, 
pale brown, densely rhizinate, the rhizines pale brown, simple to sparsely furcately branched, 0.5-1 
mm long. Pycnidia numerous; conidia bifusiform, 0.5 x 6-7 um. Apothecia well developed, 
substipitate, 3-7 mm in diameter; spores 5—6 x 9-10 um. 

Chemistry: Hypoprotocetraric, 4-O-demethylnotatic, and usnic acids. 

Type collection: On small dolerite boulders on S-facing slope, east side of Kotisephola Pass, 11.3 
km NW of Sani Pass, elev. 3200 m, Lesotho, Grid 2929 CB, Hale 81531, 6 May 1988 (US, holotype; 
PRE, isotype). 

Additional specimens examined. Lesotho: Same locality as the holotype, Hale 81398 (US); 
Kotisephola Pass summit, Grid 2929 CA, Hale 81394 (US); E slope of Mahlasela Pass (S of Oxbow), 
Grid 2828 DC, Hale 81444 (US). 

Xanthoparmelia inflata represents another member of the hypoprotocetraric acid-containing X. 
prodomokosii Hale & Elix group. The diagnostic features are the high elevation habitat and the 
puffy, little branched lobes. During the long winter season the species seems to be shaded and 
covered with hoarfrost or snow most of the time. 


Xanthoparmelia khomasiana Hale, sp. nov. (Fig. 16) 

Thallus arcte adnatus, saxicola, lobis subirregularibus, 0.3-0.8 mm latis, superne continuus, 
emaculatus, modice isidiatus, isidiis crassis, pustulatis, subtus planus, pallide brunneus, modice 
rhizinosus. 

Thallus very tightly adnate on rock, 3-4 cm broad, dull yellowish green; lobes subirregular, 
0.3—0.7 mm wide, short and irregularly branched, contiguous to subimbricate; upper surface 
continuous, shiny, sparsely isidiate, the isidia subglobose, 0.2 mm in diameter, to 0.2 mm high, the 
tips epicorticate, pale, hollow, pustular and erupting but without forming soredia; lower surface 
plane, pale brown, moderately rhizinate, the rhizines pale brown, fragile, unbranched, 0.1-0.3 mm 
long. Pycnidia and apothecia lacking. 

Chemistry: hypoprotocetraric, 4-O-demethylnotatic, and usnic acids. 

Type collection: Khomas Highlands, 47 km W of Windhoek on Hwy R56, 3.4 km W of Neu 
Heusis, $S.W.A./Namibia, Grid 2216 DA, Hale 80955, 25 April 1988 (US, holotype; PRE, isotype). 

This rare, easily overlooked species is related to the X. weberi (Hale) Hale group, but the isidia 
are strongly globose and the tightly adnate lobes much narrower. 


Xanthoparmelia kotisephola Hale, sp. nov. (Fig. 17) 

Thallus laxe adnatus, terricola, coriaceus, lobis sublinearibus, 1-3 mm latis, superne continuus, 
emaculatus, sorediis isidiisque destitutus, subtus convolutus, brunneus, dense rhizinosus. 

Thallus loosely adnate to nearly free growing on soil, remaining mostly intact to scattered, firm 
and leathery, somewhat pulvinate, 4-12 cm broad, yellowish green or darkening; lobes sublinear, 
1-3 mm wide, irregularly branched, separate to contiguous; upper surface continuous, shiny, 
irregularly cracked with age, isidia and soredia lacking; lower surface weakly to distinctly convoluted, 
pale brown to brown but darkening toward the tips, densely rhizinate, the rhizines brown or 


550 


Figures 17-20. Species of Xanthoparmelia: 17, X. kotisephola (Hale 79519, holotype); 18, X. 
lipochlorochroa (Buckingham §22115, holotype in HO); 19, X. luderitziana (Hale 80123, 
holotype); 20, X. mapholanengensis (Hale 81487, holotype). Scale in mm. 


551 


darkening, rather coarse and often projecting as a dense mat from below, simple, 0.5-1 mm long. 
Pycnidia poorly developed; conidia bifusiform, 0.5 x 5-6 um. Apothecia lacking. 

Chemistry: Salazinic, consalazinic, norstictic (tr.), protocetraric (tr.), and usnic acids. 

Type collection: On grassy humus in pasture, summit of Kotisephola Pass, 13.8 km NW of 
Sani Pass, elev. 3240 m, Lesotho, Grid 2929 CA, Hale 79519, 6 May 1988 (US, holotype; ANUC, 
ASU, LD, PRE isotypes). 

Additional specimens examined. Lesotho: Mahlasela Hill, elev. 3100 m, Lesotho, Grid 2828 DC, 
Deal & Killick 83, Hale 81495 (PRE, US); Black Mountain, Qachas Nek, Kofler 2—2-15a (LD, US). 

This convoluted species occurs in the dolerite highlands of Lesotho, widely scattered among mats 
of short grasses. It is distinguished from the American X. chlorochroa (Tuck.) Hale by the dense 
mat of dark rhizines below and the open branching pattern. 


Xanthoparmelia lipochlorochroa Hale & Elix, sp. nov. (Fig. 18) 

Thallus vagans, terricola, coriaceus, 3-5 cm latus, lobis sublinearibus, separatis, superne 
continuus, emaculatus, sorediis isidiisque destitutus, subtus valde convolutus, sparse rhizinosus. 

Thallus vagrant, free growing on soil, leathery, breaking apart into smaller colonies 3-5 cm 
broad, light yellowish green; lobes sublinear, 2-5 mm wide (to 10 mm when unrolled), sparsely 
branched, separate; upper surface continuous, shiny to mostly dull, transversely cracked and rugose 
with age, soredia and isidia lacking; lower surface strongly convoluted with little of the lower surface 
visible, light brown or darkening, smooth to foveolate-ridged, sparsely rhizinate, the rhizines brown, 
delicate, 0.3-0.6 mm long, simple. Pycnidia and apothecia lacking. 

Chemistry: traces of unidentified fatty acids, usnic acid. 

Type collection: On sandy soil, 15 mi S of Kammerer, Lincoln County, Wyoming, B. J. 
Buckingham $22115, in Weber, Lichenes Exsiccati 29 (HO, holotype; MEL, isotype). 

This is a rare fatty acid chemotype of X. chlorochroa, a common western lichen with salazinic acid. 
I wish to thank Dr. J. A. Elix for bringing the species to my attention and determining the chemistry. 
The number of Weber’s exsiccate in US contains salazinic acid and represents X. chlorochroa 
(Tuck.) Hale; the various duplicates in this widely distributed exsiccate must be retested for 
chemistry. 


Xanthoparmelia luderitziana Hale, sp. nov. (Fig. 19) 

Thallus laxe adnatus, saxicola, fragilis, 1-3 cm latus, lobis sublinearibus, 1-2 mm latis, apice 
adscendentibus, superne albo-maculatus, isidiis sorediisque destitutus, subtus planus, niger, pro 
parte flavo— marginatus, sparsissime rhizinosus. 

Thallus loosely adnate on pebbles, rather brittle, 1-3 cm broad, dull yellowish green; lobes 
sublinear, 1-2 mm wide, separate to subimbricate with ascending tips; upper surface white 
maculate, shiny, transversely cracked with age, isidia and soredia lacking; lower surface plane, 
smooth to rugulose, black, shiny, in part yellow rimmed, very sparsely rhizinate, the rhizines black, 
coarse, 0.2—0.3 mm long. Pycnidia and apothecia lacking. 

Chemistry: diffractaic (major), barbatic (tr.), squamatic (tr.), and usnic acids. 

Type collection: On pebbles in loose granite-schist desert pavement, 9 km S of Liideritz, elev. 20 
m, South West Africa/Namibia, Grid 2615 CA, Hale 80123, 28 April 1988 (US, holotype; PRE, 
isotype). 

Additional specimens examined. $.W.A./Namibia: Same locality as the holotype, Hale 80019 
(US). 

This rare species is closely related to X. walteri Knox, although the lobes are shorter and the 
thallus quite brittle. It occurs with X. norwalteri Hale (see below) on desert pavement well within the 
coastal fog zone. 


Xanthoparmelia mapholanengensis Hale, sp. nov. (Fig. 20) 

Thallus adnatus vel laxe adnatus, saxicola, coriaceus, 8-15 cm latus, lobis sublinearibus, 0.8 -2 
mm latis, convexis, sublaciniatis, apice subteretibus, superne continuus, emaculatus, isidiis 
sorediisque destitutus. 

Thallus adnate to loosely adnate on rock, rarely on soil, rather firm, often forming a centrifugate 
pattern, 8-15 cm broad, light yellowish green; lobes sublinear, 0.8-2 mm wide, convex and in part 
appearing inflated, sublaciniate, the tips blunt, subterete, darkening, separate to subimbricate; upper 
surface continuous, shiny, transversely cracked with age, isidia and soredia lacking; lower surface 
plane, shiny, brown at the tips but blackening at the center, sparsely rhizinate, the rhizines brown to 


poe 


black, 0.2—0.4 mm long. Pycnidia common; conidia bifusiform, 0.5 x 5-6 um. Apothecia lacking. 

Chemistry: salazinic (major), protocetraric (major), norstictic (minor), consalazinic, and 
usnic acids. 

Type collection: On vertical dolerite ledge face, 23.4 km N of Mapholaneng on the Sani 
Pass — Moteng Pass road, elev. 3100 m, Lesotho, Grid 2928 BB, Hale 81487, 7 May 1988 (US, 
holotype; LD, PRE, isotypes). 

Additional specimens examined. Lesotho: Same locality as the holotype, Hale 81477, 81478 (US); 
11.3 km NW of Sani Pass on the east slope of Kotisephola Pass, Grid 2929 CB, Hale 81533, 81535 
(US); Kotisephola Pass, 13.4 km NW of Sani Pass, Grid 2929 CA, Hale 81395, 81427, 81428 (US);; 
30.5 km N of Mapholaneng (1.5 km S of Letsengla), Grid 2928 BB, Hale 81537, 81539 (US); 
Mahlasela Pass (S of Oxbow), Grid 2828 DC, Hale 81497 (US); Moteng Pass, Grid 2828 DA, Hale 
81555, 81557 (US). 

This common high elevation species has a unique combination of acids, salazinic and protocetraric 
acids both forming strong spots on TLC plates, inflated, subterete lobes, and often a centrifugate 
growth pattern. 


Xanthoparmelia maxima Hale, sp. nov. (Fig. 21) 

Thallus laxe adnatus, saxicola, firmus, 6-9 cm latus, lobis subirregularibus, late apice rotundatus, 
3-9 mm latis, superne continuus, emaculatus, pro parte pruinosus, isidiis sorediisque destitutus, 
subtus planus, pallide brunneus, modice rhizinosus. 

Thallus loosely adnate on rock, leathery but breaking apart when collected, 6—9 cm broad, light 
yellowish green; lobes subirregular and broadly rotund, 3-9 mm wide, subimbricate; upper surface 
continuous, dull and in part white pruinose, rugose and transversely cracked with age, isidia and 
soredia lacking; lower surface plane, light brown, moderately to densely rhizinate, the rhizines pale 
brown, robust, simple to splayed or furcate, 0.5—-1 mm long. Pycnidia numerous; conidia bifusiform, 
0.5 x 5-6 um. Apothecia well developed, substipitate, 2-5 mm in diameter, the disc becoming 
pruinose, rarely perforate, the rim usually inrolled. 

Chemistry: norstictic, salazinic, consalazinic, and usnic acids. 

Type collection: Mudstone outcrops along hwy R388, 32 km N of junction with R63 (S of 
Richmond), elev. 1400 m, Cape Province, South Africa, Grid 3124 CA, Hale 80651, 4 April 1988 
(US, holotype; PRE, isotype). 

Additional specimen examined. South Africa. Cape Province: Same locality as the holotype, Hale 
80560 (US); on Hwy R388, 13 km N jet with Hwy R63 (S of Richmond), Grid 3124 CA, Hale 80617 
(US). 

The unusually broad lobes resemble those of chemically similar X. colorata (Gyel.) Hale, which is 
black below, as well as salazinic acid-containing X. crassilobata Hale, which grows on soil and is 
more convoluted. 


Xanthoparmelia neowyomingica Hale, sp. nov. (Fig. 22) 

Thallus laxe adnatus, terricola, 3-4 cm latus, lobis sublinearibus, 0.8-2 mm latis, margine tereti- 
laciniatis, superne continuus, emaculatus, sorediis isidiisque destitutus, subtus pallide brunneus, 
dense rhizinosus. 

Thallus loosely adnate to nearly free growing on soil, forming orbicular colonies 3-4 cm in 
diameter, firm, light yellowish green; lobes sublinear, 0.8-2 mm wide, dichotomously branched at 
the tips, subascending, soon marginally laciniate to form dense masses of weakly terete laciniae 
0.3-0.6 mm wide; upper surface continuous, shiny, isidia and soredia lacking; lower surface 
convoluted with a more or less prominent raised yellowish rim, pale brown to brown, moderately to 
densely rhizinate, the rhizines pale brown or darkening, rather coarse with splayed tips, simple to 
branched and tufted, 0.5-1 mm long. Pycnidia common; conidia bifusiform, 0.5 x 4-5 um. 
Apothecia rare, substipitate, 2-4 mm in diameter; spores not developed. 

Chemistry: stictic, constictic, norstictic, and usnic acids. 

Type collection: 3 mi S of Ward, vicinity of Glacier Lake, Boulder County, Colorado, U.S.A., 
Wirth s.n., 1 June 1962 (US, holotype; COLO, isotype). 

Except for the more conspicuous development of terete laciniae and the chemistry, this species is 
very close to another more common, salazinic acid-containing, high elevation soil lichen X. 
wyomingica (Gyel.) Hale. 


553 


Figures 21-24. Species of Xanthoparmelia: 21, X. maxima (Hale 80651, holotype); 22, X. 
neowyomingica (Wirth s.n., holotype in US); 23, X. norcolorata (Hale 81260, holotype); 24, X. 
norlobaronica (Hale 72523, holotype). Scale in mm. 


554 


Xanthoparmelia norcolorata Hale, sp. nov. (Fig. 23) 

Thallus adnatus, saxicola, 6-8 cm latus, lobis subirregularibus, sublobulatis, 1-2 mm latis, 
superne continuus, emaculatus, isidiis sorediisque destitutus, subtus planus, niger, sparse rhizinosus. 

Thallus adnate on rock, 6-8 cm broad, light yellowish green; lobes subirregular, 1-2 mm wide, 
rather short and crowded, subimbricate, sublobulate, becoming suberect and revolute at the center; 
upper surface continuous, dull, rugulose with age, isidia and soredia lacking; lower surface plane, 
black, sparsely rhizinate, the rhizines black, simple to sparsely branched, 0.3—0.6 mm long. Pycnidia 
common; conidia bifusiform, 0.5 x 6-8 um. Apothecia substipitate, 1-2 mm in diameter; spores 
5-6 x 9-10 um. 

Chemistry: norstictic, connorstictic, and usnic acids. 

Type collection: On sheltered coarse granite in large koppie, 13.6 km N of Rehoboth on east side 
of Hwy B2, elev. 1700 m, South West Africa/Namibia, Grid 2317 AA, Hale 81260, 30 April 1988 
(US, holotype; PRE, isotype). 

This rare species seems to fall near the X. colorata (Gyel.) Hale (norstictic and salazinic acids 
present) group, but it is a smaller, narrow-lobed lichen without the additional salazinic acid. 


Xanthoparmelia norlobaronica Hale, sp. nov. (Fig. 24) 

Thallus adnatus vel laxe adnatus, saxicola, fragilis, S—7 cm latus, lobis subirregularibus, 1-3 mm 
latis, superne continuus, emaculatus, sorediis isidiisque destitutus, subtus pallide brunneus, modice 
rhizinosus. 

Thallus adnate to loosely adnate, fragmenting easily, 5-7 cm broad, light yellowish green; lobes 
subirregular, 1-3 mm wide, apically rotund, short and irregularly branched, imbricate; upper surface 
continuous, shiny, rugulose with age, soredia and isidia lacking; lower surface plane, pale brown, 
moderately rhizinate, the rhizines pale brown, simple, 0.2—0.5 mm long. Pycnidia well developed; 
conidia bifusiform, 0.5 x 4-6 um. Apothecia lacking. 

Chemistry: norlobaridone, unknown Rfc 12, usnic acid, several undetermined fatty acids (det. J. A. 
Elix). 

Type collection: 3.8 km SW of Montagu in pass area on west side of road. Elev. 200 m. Cape 
Province, South Africa, Grid 3320 CC, Hale 72523, 3 Feb. 1986 (US, holotype; PRE, isotype). 

Additional specimens examined. South Africa. Cape Province: Same locality as the holotype, Hale 
72522 (US); 21.6 km S of Tarkastad on W side of Hwy R344, Grid 3226 AB, Hale 77367 (US). 

This species is comparable to the Australasian X. filarszkyana (Gyel.) Hale, which is more closely 
adnate and normally contains loxodin in addition to norlobaridone. 


Xanthoparmelia norwalteri Hale, sp. nov. (Fig. 25) 

Thallus laxe adnatus, saxicola, fragilis, 1-3 cm latus, lobis sublinearibus, 1-2 mm latis, apice 
adscendentibus, superne albo-maculatus, transversim fissuratus, isidiis sorediisque destitutus, subtus 
planus, niger, flavo-marginatus, sparsissime rhizinosus. 

Thallus loosely adnate on pebbles, rather brittle, 1-3 cm broad, dull yellowish green; lobes 
sublinear, 1-2 mm wide, separate to subimbricate with ascending tips; upper surface white 
maculate, shiny, transversely cracked with age, isidia and soredia lacking; lower surface plane, 
smooth to rugulose, black, shiny, in part yellow rimmed, very sparsely rhizinate, the rhizines black, 
coarse, 0.2—0.3 mm long. Pycnidia and apothecia lacking. 

Chemistry: norstictic, connorstictic, and usnic acids. 

Type collection: 9 km S of Liideritz, South West Africa/Namibia, Grid 2615 CA, Hale 81205, 28 
April 1988 (US, holotype; PRE, isotype). 

This unusual relative of X. walteri Knox occurs at the same locality as diffractaic acid-containing X. 
luderitziana Hale (see above). It is also smaller and more brittle than typical X. walteri. 


Xanthoparmelia protolusitana Hale, sp. nov. (Fig. 26) 

Thallus adnatus vel laxe adnatus, saxicola, firmus, 4-8 cm latus, lobis sublinearibus, 0.7-2 mm 
latis, laciniatis, superne continuus, emaculatus, isidiis sorediisque destitutus, subtus planus, niger, 
modice rhizinosus. 

Thallus adnate to loosely adnate on rock, rather stiff, 4-8 cm broad, darkish yellow green; lobes 
subirregular to sublinear, 0.7—-2 mm wide, imbricate, becoming laciniate, the laciniae sublinear, 
0.2—0.4 mm wide, black rimmed; upper surface continuous, shiny, rugulose and sparsely transversely 
cracked with age, isidia and soredia lacking; lower surface plane, black and shiny, sparsely to 


DOO 


Figures 25-28. Species of Xanthoparmelia: 25, X. norwalteri (Hale 81205, holotype); 26, X. 
protolusitana (Hale 81205, holotype); 27, X. protoquintaria (Hale 78171, holotype); 28, X. 
psornorstictica (Hale 79495, holotype). Scale in mm. 


556 


moderately rhizinate, the rhizines black, simple, 0.3-0.5 mm long. Pycnidia lacking. Apothecia 
numerous, substipitate, 1-4 mm in diameter; spores not found. 

Chemistry: stictic, constictic, and usnic acids, lusitana unknown. 

Type collection: On low sandstone ledges along small stream, 9 km W of Ramatsiliso Nek, elev. 
2400 m, Lesotho, Grid 3028 BB, Hale 81205, 28 April 1988 (US, holotype; PRE, isotype). 

Additional specimens examined. Lesotho: Same locality as the holotype. Hale 81333 (US); 7.5 km 
NW of Sani Pass on road to Mokhotlong, Grid 2929 CB, Hale 81518 (US). 

Although more or less related to X. angustiphylla (Gyel.) Hale, the thallus is rather stiff. The lack 
of norstictic acid and presence of the "lusitana" unknown (a faintly reacting yellowish spot below 
norstictic acid) suggest a closer relationship with isidiate X. /usitana (Nyl.) Krog & Swinscow. 


Xanthoparmelia protoquintaria Hale, sp. nov. (Fig. 27) 

Thallus arcte adnatus, saxicola, 2-4 cm latus, lobis sublinearibus, 0.4-0.9 mm latis, superne 
continuus, emaculatus, isidiis sorediisque destitutus, subtus planus, niger, modice rhizinosus. 

Thallus tightly adnate on rock, 2-4 cm broad, darkish yellow green; lobes subirregular to 
sublinear, 0.4-—0.9 mm wide, short, dichotomously branched, contiguous to subimbricate; upper 
cortex continuous, shiny, transversely cracked at the center, isidia and soredia lacking; lower surface 
plane, black, shiny, moderately rhizinate, the rhizines black, simple to furcate, 0.1-0.2 mm long. 
Pycnidia and apothecia lacking. 

Chemistry: hypostictic, hyposalazinic, hypoconstictic, and usnic acids. 

Type collection: On sandstone ledges in Fynbos, Fernkloof Nature Reserve at Hermanus, Cape 
Province, South Africa, Grid 3419 AD, Hale 78171, 25 Oct. 1986 (US, holotype; PRE, isotype). 

The only other hypostictic acid-containing Xanthoparmelia with a black lower surface is the 
Australian X. multipartita (R. Br.) Elix, a loosely adnate, linear-lobed species. 


Xanthoparmelia psornorstictica Hale, sp. nov. (Fig. 28) 

Thallus laxe adnatus, saxicola, coriaceus, 5-7 cm latus, lobis subirregularibus, rotundatis, 1.5-3 
mm latis, superne continuus, emaculatus, isidiis sorediisque destitutus, subtus planus, nigricans, 
modice rhizinosus. 

Thallus adnate to loosely adnate on rock, firm and leathery, 5-7 cm broad, light yellowish green; 
lobes subirregular with rotund tips, 1.5-3 mm wide, relatively short and crowded; upper surface 
continuous, dull, rugulose with age, isidia and soredia lacking; lower surface plane, brown at the tips 
but black toward the center, sparsely to moderately rhizinate, the rhizines black, simple, 0.5-1 mm 
long. Pycnidia common; conidia bifusiform, 0.5 x 6-8 um. Apothecia substipitate, 1-3 mm in 
diameter; spores 5-6 x 9-10 um. 

Chemistry: psoromic, norstictic, and usnic acids. 

Type collection: On coarse granite outcrops in pasture, 18 km NE of Keimoes on S side of Hwy 
R27, elev. 900 m, Cape Province, South Africa, Grid 2821 CA, Hale 79495, 17 March 1988 (US, 
holotype; PRE, isotype). 

Additional specimens examined. South Africa. Cape Province: Same locality as the holotype, Hale 
79493, 79494 (US). 

A narrow-lobed member of the X. colorata (Gyel.) Hale group, this species has a unique 
combination of acids. It was abundant at the type locality, an arid desert site, but has not been found 
elsewhere. 


Xanthoparmelia pustulosorediata Hale, sp. nov. (Fig. 29) 

Thallus adnatus, saxicola, fragilis, 5-8 cm latus, lobis subirregularibus, 1-3 mm latis, congestis, 
superne continuus vel albo-maculatus, rugosus, pustulatus, pustulis eruptentibus, sorediatis, subtus 
planus, modice rhizinosus. 

Thallus adnate on rock, brittle and breaking apart when collected, 5-8 cm broad, dull yellowish 
green; lobes subirregular, 1-3 mm wide, short and crowded; upper surface continuous to faintly 
maculate in part, shiny to dull pruinose, soon strongly rugose and developing coarsely sorediate, 
erupting pustules covering much of the thallus; lower surface plane, pale brown or darkening, 
moderately rhizinate, the rhizines brown, simple, 0.2-0.8 mm long. Pycnidia and apothecia lacking. 

Chemistry: stictic, constictic, and usnic acids, lusitana unknown. 

Type collection: On crumbling sheltered granite in large koppie, 5 km NE of Aus on hwy 35 (road 
to Helmeringshausen), 4.7 km N of Hwy B4, elevation 1300 m, South West Africa/Namibia, Grid 
2616 CB, Hale 81134, 27 April 1988 (US, holotype; LD, PRE, isotypes). 


D7, 


Figures 29-32. Species of Xanthoparmelia: 29, X. pustulosorediata (Hale 81134, 
holotype); 30, X. salamphixantha (Hale 79962, holotype); 31, X. saleruptens (Hale 79905, 
holotype); 32, X. salkiboensis (Santesson 21163, holotype). Scale in mm. 


558 


Additional specimens examined. South Africa. Cape Province: 3 km SW of Rooifontein on road to 
Gamoep, Grid 3018 AB, Hale 79710 (US). S.W.A./Namibia: Same locality as the holotype, Hale 
81141 (US). 

Dense pustulate-sorediate outgrowths characterize this unusual species, collected in extremely 
arid sites at the base of granite ledges where water accumulates during infrequent rains. 


Xanthoparmelia salamphixantha Hale, sp. nov. (Fig. 30) 

Thallus laxe adnatus, terricola, 4-8 cm latis, lobis sublinearibus, 2-5 mm latis, separatis, superne 
albo-maculatus, isidiis sorediisque destitutus, subtus canaliculatus, flavo-marginatus, nigricans, 
sparse rhizinosus. i 

Thallus loosely adnate to nearly free growing on soil and mosses, easily breaking apart, 4-8 cm 
broad, light yellowish green; lobes sublinear, 2-5 mm wide, dichotomously branched, more or less 
separate; upper surface uniformly white maculate, shiny, rugose and irregularly cracked with age, 
isidia and soredia lacking; lower surface weakly canaliculate with a raised yellow rim, pale brown or 
darkening and in part jet black, shiny, sparsely rhizinate, the rhizines dark brown, simple, 0.5-2 mm 
long. Pycnidia common; conidia 0.5 x 4-6 wm long. Apothecia substipitate, 3-5 mm in diameter; 
spores 5-6 x 9-10 um. 

Chemistry: salazinic (major), consalazinic, norstictic, and usnic acids, chalybaeizans unknown. 

Type collection: On sterile soil in karoo pasture, on Hwy R364, 29 km S of junction with Hwy 
R27, elev. 600 m, Cape Province, South Africa, Grid 3119 CD, Hale 79962, 24 March 1988 (US, 
holotype; LD, PRE, isotypes). 

Additional specimens examined. South Africa. Cape Province: Same locality as the holotype, Hale 
79953 (US); near Boesmanskloof River (W of Piketberg), Grid 3218 DC, Leighton 149 (BOL, US). 

While superficially similar to X. amphixanthoides (Stein. & Zahlbr.) Hale, X. salamphixantha has 
broad lobes with a heavily maculate surface, lacks terete laciniae, and darkens below. 


Xanthoparmelia saleruptens Hale, sp. nov. (Fig. 31) 

Thallus arcte adnatus, saxicola, fragilis, 1-3 cm latus, lobis subirregularibus, 0.8—1.5 mm latis, 
superne continuus, emaculatus, pruinosus, sparse pustulato-isidiatus, esorediatus, subtus planus, 
pallide brunneus, sparse rhizinosus. 

Thallus tightly adnate on rock, fragile, 1-3 cm broad, light yellowish green; lobes subirregular, 
0.8-1.5 mm wide, contiguous to subimbricate; upper surface continuous, dull white pruinose, 
sparsely pustulate-isidiate, the isidia irregularly developed from ridges, not becoming sorediate; 
lower surface plane, pale brown, sparsely rhizinate, the rhizines pale brown, simple, 0.1—0.3 mm 
long. Pycnidia and apothecia lacking. 

Chemistry: salazinic, connorstictic, and usnic acids, chalybaeizans unknown. 

Type collection: On large overhanging Table Mountain sandstone ledges, 7 km W of 
Oliewenboskraal on road to Paleisheuwel (west of Hwy N7), elev. 900 m, Cape Province, South 
Africa, Grid 3218 BD, Hale 79905, 25 March 1988 (US, holotype; PRE, isotype). 

This is a rare member of the X. eruptens Hale group, also growing on the underside of large 
overhanging ledges. 


Xanthoparmelia salkiboensis Hale, sp. nov. (Fig. 32) 

Thallus laxe adnatus, saxicola vel muscicola, pulvinatus, fragilis, 4-8 cm latus, lobis sublinearibus, 
0.8-2 mm latis, superne continuus, emaculatus, sorediis isidiisque destitutus, subtus planus, niger, 
sparse rhizinosus. 

Thallus loosely adnate on rock or mosses over rocks, pulvinate, rather brittle, 4-8 cm broad, dark 
yellowish green; lobes sublinear, 0.8—2 mm wide, dichotomously branched, black rimmed, imbricate, 
irregularly laciniate with age; upper surface continuous, shiny, rugulose and transversely cracked 
with age, soredia and isidia lacking; lower surface plane, black, shiny, smooth to slightly rugulose, 
sparsely to moderately rhizinate, the rhizines black, rather coarse and unbranched, 0.3-0.5 mm long. 
Pycnidia common; conidia bifusiform, 0.5 x 4-5 um. Apothecia not commonly developed, 
substipitate, 3-5 mm wide, the rim inrolled; spores 5-6 x 9-10 wm. 

Chemistry: salazinic, consalazinic, norstictic (tr.), and usnic acids. 

Type collection: Near Johnsell Point, Shira Plateau, Mt. Kilimanjaro, Tanzania, elev. 3950 m, 
Santesson 21163, 13 Jan. 1970 (UPS, holotype; US, isotype). 

Additional specimens examined: Kenya: Sirimon Track, Mt. Kenya National Park, Kenya, Moberg 
3948 (UPS); Wandare’s Track, Aberdare National Park, Rift Valley/Central Province, Kenya, 


559 


Moberg 4321 (UPS). Tanzania: Same locality as the holotype, Santesson 21177 (UPS); Peters Hut, 
Mt. Kilimanjaro, Hedberg 1395, Santesson 21163 (UPS); Koitobboss, Mt. Elgon, Holm 17:6 (UPS). 

This high-elevation, black-rimmed lichen appears to be common at an elevation of 3600-4000 m 
in Kenya and Tanzania. It differs from X. kiboensis (Dodge) Krog & Swinsc. in the sublinear lobes, 
pulvinate growth habit, shiny lower surface, and chemistry (a dull rugose lower surface and 
norstictic and connorstictic acids in X. kiboensis). 


Xanthoparmelia springbokensis Hale, sp. nov. ( Fig. 33) 

Thallus laxe adnatus, saxicola) 5-8 cm _ latus, lobis subirregularibus, 1.5—3 mm latis, 
superne continuus vel albo-maculatus, aetate valde rugosus, sorediis isidiisque destitutus, subtus 
planus, brunneus, modice rhizinosus. 

Thallus adnate on rock, 5—8 cm broad, dull olive green; lobes subirregular, 1.5—3 mm wide, short 
and irregularly branched, more or less suberect at the tips, imbricate; upper surface continuous to 
distinctly white maculate, shiny, transversely cracked and strongly rugulose with age; medulla often 
pale yellow, the lower part ochraceous in patches; lower surface plane, pale brown, but often 
darkening at the tips, moderately rhizinate, the rhizines pale brown, 0.3-—0.5 mm long, unbranched. 
Pycnidia common; conidia bifusiform, 0.5 X 5-6 um. Apothecia well developed, substipitate, 2-5 
mm in diameter; spores poorly developed, 5 X 9 um. 

Chemistry: salazinic (major), consalazinic, norstictic (minor), protocetraric (trace), and usnic 
acids, chalybaeizans unknown and skyrin (det. J. Elix). 

Type collection: Sloping granite domes in karoo, 1 km E of Springbok, N side of Hwy R64, elev. 
1100 m, Cape Province, South Africa, Grid 2917 DB, Hale 72249, 27 Jan. 1986 (US, holotype; 
PRE, isotype). 

Additional specimens examined. South Africa. Cape Province: Same locality as the holotype, Hale 
72251; Anenouspas, Grid 2917 BA, Hale 73232 (US); 1 km S of Nuwerus, Grid 3118 AB, Hale 72250 
(US). 

Confined to upper Namaqualand, X. springbokensis differs from closely related _X. neosynestia in 
more irregular, adnate lobes, distinctive darkish green color, strong rugosity, and presence of skyrin. 
It often grows near the base of large sheltered rocks. 


Xanthoparmelia subamplexuloides Hale, sp. nov. (Fig. 34) 

Thallus arcte adnatus, fragilis, 4-7 cm latus, lobis subirregularibus, 0.5—1.5 mm latis, superne 
continuus, emaculatus, sparse vel modice isidiatus, isidiis subcylindricis, subtus planus, pallide 
brunneus, sparse rhizinosus. 

Thallus very tightly adnate to tightly adnate on rock, fragile, 4-7 cm broad, yellowish green; lobes 
subirregular, 0.4-—1.3 mm wide, short and irregularly branched; upper surface continuous, shiny, 
subrugose and transversely cracked with age, sparsely to moderately isidiate, the isidia mostly basally 
constricted and globose to subcylindrical, easily breaking off but not erumpent, 0.06—0.08 mm in 
diameter, to 0.5 mm high, the tips pale, weakly epicorticate, unbranched; lower surface pale brown, 
sparsely rhizinate, the rhizines pale brown, simple, 0.1—0.3 mm long. Pycnidia and apothecia lacking. 

Chemistry: norlobaridone, unknowns, and usnic acid. 

Type collection: on sheltered schistose sandstone, 51 km S of Windhoek on west side of Hwy 1/5 
(opposite entrance to Bergland), elev. 1700 m, South West Africa/Namibia, Grid 2217 CC, Hale 
81144, 26 April 1988 (US, holotype; LD, PRE, isotypes). 

Additional specimens examined. $.W.A./Namibia: 47 km W of Windhoek (Khomas Highlands), 
Grid 2216 DA, Hale 81000, 81002 (US); 1 km E Neu Heusis (Khomas Highlands W of Windhoek), 
Grid 2216 DA, Hale 80956, 80957, 80959, 80961 (US); Hoogland, Waterberg Plateau National Park 
(NE of Otjiwarongo), Grid 2017 CA, Hale 80927 (US). 

Near X. amplexuloides Hale (above), this Namibian species has much smaller lobes and isidia. It 
can be considered a tightly adnate morphotype. 


Xanthoparmelia subbullata Hale, sp. nov. (Fig. 35) 

Thallus adnatus, saxicola, 5-7 cm latus, lobis subirregularibus, 1.5-2 mm latis, superne 
continuus, emaculatus, dense bullato-rugoso-isidiato, isidiis fragilibus, subtus planus, modice 
rhizinosus. 

Thallus adnate on rock, 5-7 cm broad, light yellowish green; lobes sublinear, 1-2 mm wide, 
rather short and irregularly branched, contiguous to crowded at the center; upper surface 
continuous, shiny, rugose and densely bullate-isidiate with age, the isidia fragile, 0.2-0.3 mm in 


560 


et 
aa 
eo 
a 
bool 
oR 
tool 
8 
8 


Figures 33-36. Species of Xanthoparmelia: 33, X. springbokensis (Hale 72249, holotype); 
34, X. subamplexuloides (Hale 81144, holotype (x10)); 35, X. subbullata (Hale 77072, holotype); 36, 
X. subconvoluta (Hale 74455, holotype). Scale in mm. 


561 


diameter, sometimes breaking open but not sorediate; lower surface plane, brown, moderately 
rhizinate, the rhizines brown, simple, 0.5-1 mm long. 

Pycnidia numerous; conidia bifusiform, 0.5 x 5-6 um. Apothecia adnate, 1-2 mm in diameter; 
spores not developed. 

Chemistry: salazinic, consalazinic, connorstictic, and usnic acids, chalybaeizans unknown. 

Type collection: On doleritic outcrops along road, 8.5 km W of junction Hwy R396 and Hwy R393 
near Moshesh’s Ford, elev. 1900 m, Cape Province, South Africa, Grid 3027 DD, Hale 77072, 10 Oct. 
1986 (US, holotype; PRE, isotype). 

Additional specimens examined. S. Africa. Cape Province: Same locality as the holotype, Hale 
771075 (US). Lesotho: 9 km W of Mpiti (N of Qachas Nek), Grid 3028 BA, Hale 81289, 81293 (US). 

This species is obviously related to the widespread X. chalybaeizans (Stein. & Zahlbr.) Hale but is 
differentiated by the abundant production of coarse bullate isidia. 


Xanthoparmelia subconvoluta Hale, sp. nov. (Fig. 36) 

Thallus laxe adnatus, saxicola vel terricola, coriaceus, 6-8 cm latus, lobis sublinearibus, 0.7-2 
mm latis, elongatis, superne continuus, emaculatus, sorediis isidiisque destitutus, subtus convolutus, 
brunneus, modice rhizinosus 

Thallus loosely adnate on pebbles, rarely in part free growing on adjacent soil, rather leathery, 
6-8 cm broad, light yellowish green; lobes sublinear, 0.7—-2 mm wide, elongate and little branched, 
separate to imbricate; upper surface continuous, shiny, rugulose with age, soredia and isidia lacking; 
lower surface weakly to strongly convoluted but not rolled up, brown, moderately to densely 
rhizinate, the rhizines comparatively long and protruding from below, dark brown, unbranched, 
0.5-1 mm long. Pycnidia common; conidia weakly bifusiform, 0.5 x 5-6 um. Apothecia well 
developed, substipitate, 1-2 mm in diameter with inrolled rim; spores 5-6 x 9-10 um. 

Chemistry: salazinic, consalazinic, and usnic acid. 

Type collection: Flat dolerite ridge in pasture, 6.8 km S of Sutherland on Hwy R354, elev. 1500 m, 
Cape Province, South Africa, Grid 3220 BC, Hale 74455, 29 Jan. 1986 (US, holotype; ANUC, PRE, 
isotypes). 

Additional specimens examined. South Africa. Cape Province: Same locality as the holotype, Hale 
74457, 74458 (US); Seweweekspoort, Grid 3321 AD, Hale 73426, 73435 (US); 15.9 km NE of 
Barrydale, Grid 3321 DD, Hale 74456 (US); 3.6 km N Hwy R62 on Kruisrivier road, Grid 3321 BB, 
Hale 73512 (US); Uniondalespoort, Grid 3323 CA, Hale 73442 (US); 42 km SSW of Nuwerus on 
Hwy R363, Grid 3118 AD, Hale 78845 (US). 

More or less related to the vagrant X. chlorochroa group, X. subconvoluta is loosely attached to 
pebbles and is not so strongly convoluted as to hide the dense, protruding mass of rhizines below. It 
is also close to X. amphixanthoides (Stein. & Zahlbr.) Hale, which has terete secondary laciniae. 
From North America, X. wyomingica has shorter, more congested lobes and is not as dark below. 


Xanthoparmelia subluminosa Hale, sp. nov. (Fig. 37) 

Thallus laxe adnatus, coriaceus, 5-8 cm latus, lobis subirregularibus, 2-4 mm latis, superne 
continuus, emaculatus, dense isidiatus, isidiis cylindricis, medulla pro parte ochracea, subtus planus, 
niger, sparse rhizinosus. 

Thallus adnate to loosely adnate on rock, firm but easily breaking apart, 5-8 cm broad, light 
yellowish green; lobes subirregular, 2-4 mm wide, irregularly branched, dark brown rimmed, 
imbricate; upper surface continuous, shiny, densely isidiate, the isidia cylindrical, 0.15-0.2 mm in 
diameter, to 1 mm high, black tipped, simple to coralloid branched; lower medulla in part dull 
reddish orange; lower surface plane, black, shiny, sparsely rhizinate, the rhizines black, coarse, 
unbranched, 0.4—0.8 mm long. Pycnidia common; conidia bifusiform, 0.5 x 5-6 um. Apothecia 
lacking. . 

Chemistry: salazinic, consalazinic (minor), norstictic (tr.), protocetraric (tr.), and usnic acids, 
skyrin and a second unidentified anthraquinone (det. J. A. Elix). 

Type collection: Coppins Crossing, elev. 600 m, Canberra, A.C.T., Australia. Hale 58524, 8 Jan. 
1982 (US, holotype; CBG, isotype). 

This is very close to X. luminosa (Elix) Elix & Johnst., another skyrin-containing Australian lichen 
which lacks isidia. 


Xanthoparmelia submougeotii Hale, sp. nov. (Fig. 38) 
Thallus adnatus, saxicola, 3-4 cm latus, lobis sublinearibus, 0.6-1.2 mm latis, imbricatis, superne 


562 


continuus, emaculatus, sorediatus, soraliis subterminalibus, capitatis, subtus niger, modice 
rhizinosus. 

Thallus adnate on rock, rather brittle and subpulvinate, 3-4 cm broad, dull yellowish green; lobes 
sublinear, 0.6—1.2 mm wide, rather short and irregularly dichotomously branched, brown rimmed, 
imbricate; upper surface continuous, shiny, sorediate, the soralia subterminal, capitate, 0.5-1 mm in 
diameter; lower surface plane, dark brown at the tips but black at the center, shiny, rugulose, 
moderately rhizinate, the rhizines black, simple, 0.2—0.4 mm long. Pycnidia and apothecia lacking. 

Chemistry: stictic, constictic, norstictic, and usnic acids. 

Type collection: Cordon Barril, Masafuera, Juan Fernandez, C. & I. Skottsberg 90, 1 March 1917 
(NY, holotype; US, isotype). 

This rare sorediate species is related to X. mougeotii (Schaer.) Hale, a well-known much smaller, 
very tightly adnate lichen. The only other sorediate Xanthoparmelia with a black lower surface is X. 
microspora (Mil. Arg.) Hale, which contains salazinic acid. 


Xanthoparmelia substenophylloides Hale, sp. nov. (Fig. 39) 

Thallus arcte adnatus, saxicola, 2—6 cm latus, lobis sublinearibus, 0.3-—0.6 mm latis, laciniatis, 
superne continuus, emaculatus, modice isidiatus, isidiis cylindricis, subtus planus, niger, modice 
rhizinosus. 

Thallus tightly adnate on rock, 2—6 cm broad, light yellowish green but darkening at the center; 
lobes sublinear, 0.3-0.6 mm wide, elongate, dichotomously branched, separate to contiguous, 
laciniate marginally with age, the laciniae dichotomously branched, 0.2—0.4 mm wide; upper surface 
continuous, shiny, moderately isidiate, the isidia cylindrical, 0.06—0.13 mm in diameter, 0.1-0.3 mm 
high, the tips epicorticate, mostly pale, unbranched to sparingly branched; lower surface plane, dark 
brown at the tips but blackening at the center, shiny, moderately rhizinate, the rhizines black, simple, 
0.1-0.2 mm long. Pycnidia rarely developed; conidia bifusiform, 0.5 x 5-6 wm. Apothecia 
lacking. 

Chemistry: Stictic, constictic, norstictic and usnic acids. 

Type collection: Sandstone ledges, about 5 km E of Holy Forest (NW of Thohoyandou, elev. 1000 
m., Venda, South Africa, Grid 2230 CD, Hale 79148, 14 Nov. 1986 (US, holotype). 

Additional specimens examined. South Africa. Cape Province: 28.2 km W of Mosselbaai, Grid 
3421 BB, Hale 73844 (US); Venda: 2 km E of Holy Forest, Grid 2230 CD, Hale 79149 (US); 
Transkei: Moordenaarsnek on R56, Grid 3028 DC, Hale 76913 (US). Brazil: Rio de Janeiro, 
Glaziou s.n. (US). Venezuela: Mucuy, Estado Mérida, Hale 43039a (US). 

This species is characterized by the narrow, relatively elongate, contiguous, little branched 
lobes and delicate pale-tipped isidia. Another South African species, X. pseudocongensis Hale, is 
close but has shorter, black-rimmed, crowded lobes and black-tipped isidia. The Australasian X. 
mougeotina (Nyl.) Galloway has smaller, subglobose, black-tipped isidia. 


Xanthoparmelia tsekensis Hale, sp. nov. (Fig. 40) 

Thallus arcte adnatus, saxicola, coriaceus, 6-8 cm latus, lobis sublinearibus, 0.6—1 mm latis, 
sublaciniatis, superne continuus, emaculatus, isidiis sorediisque destitutus, medulla pro parte 
ochracea, subtus planus, brunneus, modice rhizinosus. 

Thallus tightly adnate on rock, leathery, 6-8 cm broad, darkish yellow green; lobes sublinear, 
0.6-1 mm wide, irregularly branched, short and crowded, sublaciniate, the laciniae in part 
subascending, black rimmed; upper surface continuous, shiny, bullate and transversely cracked with 
age, isidia and soredia lacking; lower medulla in part ochre; lower surface plane, brown, moderately 
rhizinate, the rhizines brown, simple, 0.2—0.3 mm long. Pycnidia common; conidia bifusiform, 0.5 x 
5-6 wm. Apothecia lacking. 

Chemistry: stenosporonic, colensoic, salazinic, norstictic, and usnic acids, 2-3 unidentified 
anthraquinones. 

Type collection: On crumbling dolerite boulder along river, 5.2 km NW of Thaba Tseka junction 
on Moteng Pass-Sani Pass road, elev. ca 2900 m, Lesotho, Grid 2928 BD, Hale 81451, 7 May 1988 
(US, holotype; LD, PRE, isotypes). , 

This species, externally similar to X. conjuncta Hale (descibed above) represents a new 
combination of acids for the nonisidiate group with a pale lower surface. Another Drakensberg 
endemic, X. naudesnekia Hale, also has stenosporonic acid as the major metabolite but along with 
the hypostictic acid series, as well as broader lobes, to 1.5mm wide. Other South African 
species with stenosporonic acid (nonisidiate X. shebaiensis Nash & Elix and X. stenosporonica Hale 


563 


FEE RERPRR ERE LEE 


a 
= 
i 
* 

ad 
a 
- 
- 
Cd 
bd 
coal 
-_ 
od 

“a 
- 
coed 
= 
- 
-~- 
- 
Fel 


Figures 37-40. Species of Xanthoparmelia: 37, X. subluminosa (Hale 58524, holotype); 38, X. 
submougeotii (Skottsberg 90, holotype); 39, X. substenophylloides (Hale 79148, holotype); 40, X. 
tsekensis (Hale 81451, holotype). Scale in mm. 


564 


and isidiate X. keralensis Hale) are very tightly adnate and black below. 


Acknowledgments 
Field studies were supported by grants from the National Geographic Society and the Scholarly 
Studies Program, Smithsonian Institution. I wish to thank Dr. J. A. Elix for assistance in determining 
the chemistry of a number of specimens. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 565-576 January-March 1989 


SUGIYAMAEMYCES, A NEW GENUS OF LABOULBENIALES 
(ASCOMYCETES) ON CLIDICUS (SCYDMAENIDAE) 


ISABELLE I. TAVARES 


University Herbarium, University of California, 
Berkeley, California 94720 


JEAN BALAZUC 


30 Grande Rue Charles de Gaulle, 
94130 Nogent/Marne, France 


SUMMARY 


The genus Sugiyamaemyces is established for a taxon 
on Clidicus (Coleoptera, Scydmaenidae), in which the 
receptacular cells directly subtending the appendage 
and perithecium are laterally adnate, the perithecium 
arises from the third, rather than the second cell of 
the receptacle, and an extensively branched appendage 
is frequently present. The species S. orousettii is 
described. 


In many genera of the Laboulbeniales the exact point of 
origin of the perithecium is uncertain because of a lack 
of sufficiently immature material. Usually, the source 
has been judged to be the cell directly underlying the 
perithecial stalk cell at maturity. Except in the highly 
modified genus Coreomyces (Thaxter, 1908) and rarely in 
Herpomyces (see Tavares, 1966), the perithecium in the 
Laboulbeniales arises as a lateral branch of one of the 
receptacular cells. This cell has generally been iden- 
tified as cell II (the second cell of the typically 3- 
celled lower spore segment) or as a secondary derivative 
of cell Ii. In Amorphomyces (Tavares, 1970) and 
Rhizopodomyces (Benjamin, 1979), a septum does not sepa- 
rate two upper cells in the lower spore segment, so that 
the perithecium is an outgrowth of the upper cell of a 
two-celled receptacle. In most genera of the 
Dimorphomyceteae and apparently also in Nanomyces, peri- 
thecia are produced by outgrowths of the foot cell I 
(Tavares, 1985). 

Thalli recently found by J. Orousset on Clidicus 
formicarius Pascoe (Coleoptera, Scydmaenidae, Clidicini) 
differ from previously described genera of the 
Laboulbeniales by the production of a series of lateral 
appendages from the superposed cells of the primary appen- 
dage and by the presence of laterally adnate III and VI 


566 


cells (the receptacular cells subtending the appendage and 
perithecium). Adnate III and VI cells are a conspicuous 
characteristic of Stigmatomyces and their presence in the 
fungi on Clidicus suggests a relationship, particularly 
because the appendage cells sometimes bear phialides an- 
teriorly (on the side toward the perithecium), rather than 
branches. Furthermore, there apparently are four outer 
wall cells in each vertical row in both genera--two tall 
lower cells and two very short upper cells. However, 
Stigmatomyces is restricted to Diptera and normally has an 
unbranched appendage. 

In addition, the fungus on Clidicus differs from 
Stigmatomyces in the point of origin of the perithecium, 
as well as the degree of cell wall thickening of some 
perithecial cells. Consequently, this taxon is now being 
described as a new genus, in honor of Dr. Keiichi 
Sugiyama, Shizuoka University, Japan, who has been active, 
with his students, in investigating the Laboulbeniales of 
the southeast Asian area. 


Sugiyamaemyces I. Tavares & Balazuc, gen. nov. 


Receptaculum tribus cellulis confectum, quarum summa 
perithecii stirpem attingit. Appendicis superpositae 
cellulae ramos a latere sufferentes. Basalium perithecii 
cellularum atque perithecii stirpis cellulae parietes haud 
crassescentes. 


Typus: Sugiyamaemyces oroussetii. 


Basal and suprabasal cells of thallus superposed; 
upper cell of 3-celled receptacle laterally adnate to 
stalk cell of perithecium; appendage consisting of series 
of superposed cells, which produce lateral appendages. 
Walls of perithecial basal cells and walls of secondary 
stalk cells not thickening, so that these cells are not 
clearly visible and soon deteriorate. 


Sugiyamaemyces oroussetii I. Tavares & Balazuc, sp. nov. 


Receptaculi tres cellulae componentes perithecii 
stirpem longitudine subaequantes. Primi axis appendicis 
cellulae multo breviores, latae, a latere appendices 
proferentes, quae ipsae casu longos graciles ramos ferunt. 
Perithecium fusiforme, vel simplex, vel brevi (long. 1 um) 
prominentia ornatum. 

Parasitus Clidici formicarii Pascoe (Coleoptera, 
Scydmaenidae, Clidicinae) in insula Borneo. 


Thallus pale to deep brownish yellow; three superposed 
cells of receptacle approximately same height as peri- 
thecial stalk cell; cells of primary axis of appendage 
much shorter, broad, separated by diagonal septa, these 
cells extending anteriorly into lateral appendages; each 
lateral appendage may produce a series of long, slender, 
lateral branches. Perithecium somewhat fusiform, narrow 


567 


~200 


100 yum 


pegs. gioe. | o.,OrOussetii onuC. formicarius, 
Borneo (ZMB). Drawings by J. Balazuc. 1. Holotype 
(UC)... 2. Mature thallus with well-developed 


appendage system (Balazuc collection). 


568 


to broad, borne on stalk cell that is longer than wide and 
adnate to cell III of the receptacle; perithecium having 
no visible secondary stalk cell or basal cells. Peri- 
thecial apex simple or having extremely short outgrowth 
(ca. 1 um x 1 um) terminating one vertical row of outer 
wall cells. Trichogyne stump usually prominent. Size 
range: total length: 118-250 um; total length of peri- 
thecium above stalk cell (in thalli having 2-celled 
spores): (59)65-106(120) x (22)24-40(45) um; length of 
cell “Il: 15-37) um: width, of cell. I1:) 15-29 Aiwa 
Clidicus formicarius Pascoe, Borneo, Zoological Museum, 
Berlin, DDR (A. R. Wallace collection ?; see Pascoe, 1863) 
(UG)malfigseuel, wl Ops 

The perithecial apices vary, but there is usually a 
short, thin, apical outgrowth from one cell row (figs. 5, 
10). Often, there is also visible on sharply narrowed 
perithecial apices a shorter narrow outgrowth. Other 
perithecia have a simple, more evenly rounded apex. The 
side of the perithecium just below the apex generally is 
convex on the side opposite the trichogyne stump, which is 
on a plane or slightly concave surface. The trichogyne 
stump (figs. 1, 8, 10) is usually’ 1/2-2/3 the distance 
between the base of the third tier of outer wall cells and 
the perithecial apex, but it is sometimes closer to the 
apex or not visible at all. Although the perithecia 
appear to have four outer wall cells in each vertical row, 
septa between these cells are difficult to see. The two 
lower cells in each row are extremely tall, with lower 
septa in all rows presumably at the same levels (second 
septum shown in figs. 1, 2). There seems to be variation, 
however, in the height of the subapical and apical cells 
in each row. The upper septa of the third tier of outer 
wall cells, when they are visible at all, do not seem to 
lie at the same level in all four vertical rows. The 
visible septa sometimes are midway between the upper end 
of the second tier of cells and the perithecial apex. At 
other times, the third or subapical cells are quite short. 
In the primary appendages, the lower cells become widened 
and secondary lateral branches grow toward the perithecium 
(figs. 3, 4). There are usually about four wide cells in 
the secondary branches, which produce narrow, thin 
tertiary branches extending upward. At first, there is a 
tall I cell, and a slightly shorter II cell, wetiimage 
perithecial primordium rising from near the base of the 


Figs. 3-5. Sugiyamaemyces oroussetii on Clidicus 
formicarius. 3. Mature thallus with extensively 
branched appendage, proportionally longer III and VI 
cells (foot broken off). Orousset collection (male). 
Borneo (ZMB). X 450. 4-5. Orousset collection (type 
of host, male). Sarawak (BMNH). X 450. 4. =Small 
thallus having few spores (length indicated by lines), 
well-developed appendage. 5. Larger thallus having 
thicker cell walls; note slender, short, apical 
protuberance on perithecium (arrow). 


569 


570 


IIl cell (fig. 7, right) ...At this time, therevissageno.. 
row of sessile phialides on an unbranched appendage. It 
is from the position of these phialides that the secondary 
branches arise. In some mature thalli, the primary 
appendage axis remains narrow and any lateral outgrowths 
are broken off. 

Two males of Clidicus formicarius were found to be 
infected. On one from Borneo (Zoological Museum, East 
Berlin, DDR), thalli were found on the profemurs, pro- 
tibias, and brushes of the apex of the mesotibias. On the 
type of C. formicarius from Sarawak (British Museum of 
Natural History), thalli were found on the profemurs and 
mesotibias. Clidicus is typically an Indomalayan genus. 
Species seem to be rare in nature and are infrequently 
represented in museum collections. They live in forest 
humus and their biology is not well known. The hosts of 
Sugiyamaemyces oroussetii may be termitophiles or myrmeco- 
philes, which could account for their scarcity in museum 
collections. 

There are two quite different forms of thalli among 
the specimens observed. One has a broad perithecium and 
heavily thickened walls on lower appendage and upper 
receptacle cells; in this form, there is a tendency for 
the appendage to diverge at a strong angle. In the other 
form, the perithecium is quite slender and the appendage 
and receptacle tend to be curved in an arc; the cell walls 
are not strongly thickened. The exact position on the 
host of thalli of the two forms was not noted; it is 
probable that the form of the thallus is related to posi- 
tion of growth and that thickness of the lower appendage 


Fig. 6. Stigmatomyces on Anastrepha striata 
Schiner, Guacimo, Province of Limon, Costa Rica, April 


8, 1987, comm. I. Hedstrom. Beginning of perithecial 
development from suprabasal cell. Lactophenol-cotton 
blue. X 670.. Figs. 7-8. Sugiyamaemyces oroussetii on 
Clidicus) formicarius. 7. | Young’ thalla Atala. 
trichogyne stage, showing origin of perithecium from 
cell III--long, slender branches emerge from each of 
the superposed cells of the appendage. At right, two 
sporelings, both having perithecial outgrowths, cells 
I and II, and apparent antheridia (the top and bottom 
ones shown seem to come from the upper thallus with 
intact foot and the middle antheridium from the lower 


thallus). The long primary appendage of the upper 
thallus extends to the larger thallus at left. 
Orousset collection (from male). Borneo (ZMB). X 670. 


8. Mature thallus showing origin of VI cell from cell 
Iitl; II and the cell walls of the appendage axis have 
not broadened and II and VI have not become adnate. 
Appendage development is slight. Orousset collection, 
from type of host species (male), Sarawak (BMNH). 

X 450. 


571 


572. 


cell walls increases with age. There seem to be no dis- 
tinct characters that would indicate that these two forms 
should be placed in different species. Both forms were 
found among specimens from Sarawak and those labelled 
Borneo. 

In a well-developed appendage system, a secondary axis 
may form anteriorly (toward the perithecium) from each of 
the lower cells of the primary axis. Sometimes, cells may 
only produce phialides or a short row of unmodified cells 
that do not bear a series of anterior branchlets. Some of 
the sessile phialides of the young thallus probably grow 
out to form the regularly arranged branches. In the mate- 
rial labelled "Borneo," a well-developed secondary axis 
may resemble the first because the lower cells widen and 
become thick-walled. The Sarawak thalli had undergone no 
widening of the lower secondary axis cells. Each of the 
secondary axis cells may produce anteriorly a long, 
slender, hyaline, septate branchlet (fig. 5) that may 
sometimes be hooked at the end rather than straight. 

These slender branchlets also occur on the upper cells of 
the primary axis, generally at the end of a group of 
short, rounded cells (fig. 5). Thus, an appendage system 
can have a complex series of branches or it may be quite 
simple, usually because the long branches have broken off 
by maturity. The initial appendage, bearing lateral phia- 
lides (fig. 7), is reminiscent of that of Stigmatomyces. 
In addition, in young thalli on Clidicus, the perithecial 
primordia are attached near the base of cell III (figs. 

7, 8), rather than at the upper end of cell II as in 
Stigmatomyces (fig. 6). in the Clidicus fungiyaaae 
attachment to the upper part of cell II occurs later as 
the cells enlarge, thus obscuring the true point of origin 
of the’ perithecium (figs. 3-5), 

Furthermore, the secondary stalk cell (VII) and the 
basal cells of the perithecium have well-developed cell 
walls in Stigmatomyces that are clearly visible at matu- 
rity. In the taxon on Clidicus, the septa between the 
outer wall cells are difficult to see and the walls of the 
secondary stalk cell and perithecial basal cells are 
almost impossible to detect. 

One specimen has been found by J. Orousset on the 
lectotype of Clidicus laticeps Pic (Shembaganur, Madura 
Province, India, MNHN, Paris) that appears to be a second 
species. However, the material is inadequate and descrip- 
tion should await the discovery of more specimens. In 
this taxon, a narrow perithecial base forms a short stalk 


Figs. 9-10. 9. Sugiyamaemyces sp. on Clidicus 
laticeps, India. Orousset collection. Lectotype 

of host species (MNHN, Paris). Line indicates narrow 
stalk at: base of perithecium.  X*560.))/107 
Sugiyamaemyces oroussetii on C. formicarius, Borneo 
(ZMB). Holotype (UC). Note apical outgrowth (arrow), 
trichogyne stump (line). xX 560. 


573 


574 


rising from cell VI and the appendage is less well deve- 
loped (fig. 9). The perithecium widens subapically just 
below what appears to be a slightly inflated subapical 
tier of outer wall cells. 


DISCUSSION 


In Sugiyamaemyces, there is an unusual manner of peri- 
thecial production. The appearance of the septum across 
the top of the third cell indicates that it is probably 
the original spore septum. The two lower cells of the 
receptacle are separated by a thinner cross-wall than that 
separating the second and third cells (figs. 5, 10). This 
suggests that the lowermost septum is produced last and 
that the first septum of the receptacle is that separating 
cell III from the initial lowermost cell. Thalli having 
well-developed perithecia give the appearance of a peri- 
thecium arising from the upper end of cell II, the supra- 
basal cell; however, in some thalli, it is clear that the 
perithecium actually arises from the side of the third 
receptacle cell (figs. 7, 8). Apparently, as the recep- 
tacle and perithecial stalk cell enlarge, the lower wall 
of the stalk cell and a portion of the upper wall of the 
suprabasal cell grow together, forming a common wall. It 
was not possible to detect a clear pore between VI (the 
perithecial stalk cell) and the _third receptacle cell. 
However, the appearance of the wall and cytoplasm indi- 
cated that there is probably a central pore in the septum. 
One could regard the perithecium-producing cell as an 
undivided II-III cell and the second cell of the recep- 
tacle as a Ia cell, secondarily separated from the I cell. 
In Chitonomyces a Ia cell may be formed and division of II 
and III occurs late. In this unrelated genus, the wall 
cells of the perithecium are sometimes hard to distinguish 
as they are in Sugiyamaemyces (see Thaxter, 1924; Tavares, 
1995 )). 

In some other genera, absence of the appropriate 
' stages may have led to a misinterpretation of the origin 
of the perithecium and the perithecium may, in fact, have 
arisen from the third, rather than the second, cell of the 
receptacle. The significance of the production of the 
perithecium by the cell just below the original spore 
septum rather than the cell just above the foot cell is 
not clear, but perithecial origin might be influenced by 
the position and identity of the first septum to be formed 
in the receptacle. In those genera that have been most 
thoroughly studied, cell II produces the perithecium, but 
a generalization about origins based on a few genera is 
certain to have exceptions when sufficient material is 
available for study. 

In Sugiyamaemyces, the walls of the perithecial basal 
cells are thin and difficult to detect, as they are in 
Diplopodomyces on myriapods, Kruphaiomyces on 
Endomychidae (Coleoptera), Distolomyces on Dermaptera and 
genera in Dimorphomyceteae (having compound antheridia), 
Euphoriomyceteae (having specialized perithecial apices 


575 


and simple phialides), and Drepanomyceteae in the 
Ceratomycetaceae (see Tavares, 1985, Thaxter, 1931). The 
presence of genera having poorly developed cell walls in 
the perithecium (often within a relatively thick enve- 
loping wall) in several unrelated groups suggests that 
this character is a modification that occurs in more 
highly evolved genera and perhaps is associated with a 
reduction in thallus size or an adaptation to some condi- 
tions of the environment. When the thallus becomes more 
compact, it should become less susceptible to damage by 
contact with surrounding objects. The thickening of the 
outer wall of the base of the perithecium and the 
inclusion of perithecial basal cells and the secondary 
stalk cell within this protective wall would reduce the 
need for thick walls around each of these lower cells. 
The short, delicate stalk subtending the perithecium in 
the undescribed taxon of Sugiyamaemyces on Clidicus 
dataceps conceivably would reduce the ability of this 
taxon to withstand damage. 

With the possible exception of Acompsomyces stenichni 
(Scheloske) I. Tavares (excluded by Benjamin [in press] 
from the genus) and a possible undescribed species of 
Stemmatomyces, Sugiyamaemyces does not appear to be 
closely related to other genera occurring on Scydmaenidae. 
Diclonomyces has a simple appendage system consisting of 
long branches arising basally. The perithecium of 
Acrogynomyces has a persistent terminal trichogyne base; 
the thallus has a simple appendage bearing a few phia- 
lides. A certain amount of variation occurs in appendage 
development in Cryptandromyces, which has clearly 
delimited subequal cells in the outer perithecial walls, 
rather than strongly unequal cells (see Tavares, 1985). 
The genera that have been reported previously on 
Scydmaenidae have a greater degree of thickening of the 
cell walls of the basal cells of the perithecium, as well 
as more readily visible perithecial wall cells. 

The seriate primary axis cells in the appendage of 
Sugiyamaemyces bear a similarity to the unbranched seriate 
appendages of the undescribed Stemmatomyces on 
Scydmaenidae (Coleoptera) and of Stigmatomyces on Diptera. 
The arrangement of lateral branches produced by the 
primary axis cells in Sugiyamaemyces resembles the close, 
regular arrangement of primary axis cells and lateral 
branches in Sphaleromyces lathrobii Thaxt. and 
Corethromyces cryptobii Thaxt., both parasitizing 
Staphylinidae (Coleoptera) (see Tavares, 1985). In these 
two species, there are well-defined perithecial outer wall 
and basal cells. 

At first glance, one is struck by the similarity of 
the receptacle of Sugiyamaemyces and that of Stigmatomyces 
because of the adnate III and VI cells. Evidence indi- 
-cates that as in Laboulbenia, the VI cell of Stigmatomyces 
arises from cell II (fig. 6; see also Thaxter, 1896, pl. 
1). Nevertheless, it is possible that Sugiyamaemyces and 
Zeugandromyces, for example (parasites of Staphylinidae 
Originally placed in Stigmatomyces; see Thaxter, 1931), as 


576 


well as Stigmatomyces, Acompsomyces, and Stemmatomyces, 


may have arisen from a common ancestor. Both Scydmaenidae 


and Staphylinidae 
Sugiyamaemyces is 
gone modification 
elongation of the 
perithecial outer 
appendage system. 


belong to the Staphylinoidea. Perhaps 
a highly evolved genus that has under- 
of the lower part of the perithecium, 
lower two cells in the vertical rows of 
wall cells, and elaboration of the 
Stigmatomyces has also undergone elon- 


gation of the lower perithecial outer wall cells. The 
habits of its hosts may reduce the likelihood of damage to 
a slender, elongate thallus. 


ACKNOWLEDGMENTS 


The authors are grateful to J. Orousset and the direc- 
tors of the institutions housing the collections from 
which the specimens were obtained, as well as I. Hedstrom, 
Uppsala University, for Stigmatomyces material. We are 
also indebted to J. Hendel, Scientific Photographic 
Laboratory, University of California, Berkeley, for the 
photographs. We wish to thank Dr. R. K. Benjamin for his 
review of this publication. 


Benjamin, R. K. 


BIBLIOGRAPHY 


1979. lLaboulbeniales on 


semiaquatic Hemiptera. III. Rhizopodomyces. 
AUT SOM ta ho 4 Ole 


Pascoe) "ficar. 


1863. V.--Notices of new or little- 


known genera and species of Coleoptera. Part IV. 
Silphomorpha [Carabidae]. Westwood, Trans. Linn. 


SOGtEENV IL cud. 


on T Te Des 
TAVaGCeS ek shih. 


415. .Ji.. Entomol. 2: -26—56 ee ee 


1966. Structure and development of 


Herpomyces stylopygae (Laboulbeniales). Amer. J. 


Bots Se 5ocacL 


3233 
1970. The appendage of Amorphomyces 


(Laboulbeniales). Mycologia 62: 741-749. 


Ascomycetes). 
Braunschweig. 


1985. Laboulbeniales (Fungi, 
Mycologia Memoir no. 9. J. Cramer, 
627m. 


Thaxter, R. 1896. Contribution towards a monograph 
of the Laboulbeniaceae. Mem. Amer. Acad. Arts Sci. 


aoe fier 2, Dug 


Pls. I-XxXVI. 


- 1908. Contribution toward a monograph 
of the Laboulbeniaceae. Part II. Mem. Amer. Acad. 


ArtsvSCi. el: 


21/—469) Je GPRS woXXV Dl Te Xe, 


- 1924. Contribution towards a monograph 
of the Laboulbeniaceae. Part III. Mem. Amer. Acad. 


Arts Sci. 14: 


O94 2 Geer LS se beer 


- 1931. Contribution towards a monograph 
of the Laboulbeniaceae. Part V. Mem. Amer. Acad. 


AViSTSC 16s 


hee 4 Seem LS me ke 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 577-599 January-March 1989 


INDEX TO J. B. ELLIS’S TYPES OF PYRENOMYCETES WITH 
AMYLOID ASCAL RINGS 


KATIA F. RODRIGUES 


The New York Botanical Garden 
Bronx, 10458-5126 NY 


ABSTRACT 
An index to the types of 202 species and infraspecific taxa 
now belonging to Amphisphaeriaceae, Diatrypaceae, and 
Xylariaceae that were described by J. B. Ellis is presented. 
The names listed here are based on specimens deposited in 
herbaria at NY, FH, BPI. A list of synonyms based on the 
literature is provided. 


INTRODUCTION 

Job Bicknell Ellis (1829-1905) was born in northern New York State, but lived 
most of his life in Newfield, New Jersey. Most of his collections were made in 
Newfield or the immediate vicinity. For several years he was a teacher of classics 
and also school principal. In 1855 he began his mycological work when he started 
corresponding with Henry William Ravenel, a well established mycologist from 
South Carolina. Ravenel encouraged Ellis’s interest in fungi, introducing him to 
Systema Mycologicum (Fries, 1821-1832) and other mycological works. In addition, 
Ravenel advised Ellis to send undescribed species to European mycologists 
because of their older tradition in that field as well as their larger, world wide 
collections. He maintained a very close contact with mycologists from North 
America and Europe, such as M. J. Berkeley, M. C. Cooke, W. A. Kellerman, H. 
Rehm, F. de Thuemen, P. Sydow, P. A. Saccardo and others, who in many ways 
contributed to his knowledge of fungi (Kaye, 1986; Rogers, 1981). 

Ellis had a great interest in parasitic ascomycetes, however, he described many 
fungi in other groups as well. He issued the following exsiccatae, with the support 
of W. G. Farlow (at Harvard University), and B. M. Everhart: FUNGI NOVA- 
CAESAREENSES (Fungi of New Jersey) Ellis, NORTH AMERICAN FUNGI 
Series I - Ellis, and Second Series - Ellis & Everhart, and FUNGI COLUMBIANI 
- Ellis & Ev. Everhart, a Philadelphia busisnessman, was a long-time associate of 
Ellis, and they published new species as "Ell. et Ev". Although at first an amateur, 
Ellis was able to build up his own scientific reputation, and is widely recognized for 
his pioneer work in mycology. In 1896, Ellis sold his Herbarium, types, and library 
to the New York Botanical Garden (NY), and as result of this, most of the 
holotypes are found at NY, as are sets of North American Fungi - Series I, and 
Second Series, and Fungi Columbiani. The Herbarium Mycologicum (Everhart’s 
sets of Ell. & Ev. species), Fungi Nova-Caesareenses, and both Series of North 
American Fungi are deposited in the Farlow Herbarium (FH). The U. S. National 
Fungus Collection (BPI) has parts of both series of North American Fungi and 
Fungi Nova-Caesareenses. More detailed information about locations of the 
exsiccatae cited above are given by Stevenson (1971), and Stafleu and Cowan 


578 


(1976). The following genera were named after Ellis: Ellisiella Sacc. (1880), 
Ellisiodothis Theiss. (1914), Ellisiellina da Camara (1949), Ellisiopsis Bat. (1956), 
Ellisia Bat. & Peres (1965), and Elletevera Deighton (1969), the latter in honor of 
the Ellis & Everhart association. 

In the present paper an index to the 202 taxa named by J. B. Ellis in genera 
that are today attributed to the families Amphisphaeriaceae, Diatrypaceae, and 
Xylariaceae is presented. These families are closely related to one another, and 
have as one of their most important features an ascal ring that stains blue with 
iodine (in Melzer’s solution), readily distinguishing them from other families of 
Pyrenomycetes (although the ascal rings of many Diatrypaceae either do not 
become blue in iodine, or are so small, or the reaction so weak that it is essentially 
absent). According to Miller & Arx (1973) these families are placed within the 
order Sphaeriales, whereas Eriksson & Hawksworth (1987) place the Xylariaceae 
and Amphisphaeriaceae within the Xylariales, and the Diatrypaceae in its own 
order, Diatrypales. 

Type specimens located in the herbarium of The New York Botanical Garden 
are considered to be holotypes, except those described by Cooke & Ellis. For those 
species, the material at NY is considered isotypic, and the holotype is supposedly 
at The Royal Botanic Garden, Kew (K). Type specimens located at FH, and BPI 
are considered isotypes. Some specimens issued in North American Fungi are 
isotypes, and these are so indicated in the index. Where more than one collection 
is cited in the protologue, all types are considered to be syntypes, unless a lectotype 
has been designated. No lectotypification are expressed or implied here. 
Information within brackets [] indicates additional data provided from specimens 
located at FH, BPI, and/or the protologue. The latter type is designated by (lit.). 
The notation [lit.] is used only when specimens were not located and cites 
information strictly from the protologue. 


INDEX TO TAXA 


Amphisphaeria aspera Ell. & Ev., Bull. Torrey Bot. Club 27: 52. 1900. 
On Tetradymia spinenscens, COLORADO: Montrose, Bethel 517 (HOLOTYPE: NY). 


Amphisphaeria atrograna (Cke. & Ell.) Sacc., see Sphaeria atrograna Cke. & Ell. 


Amphisphaeria confertissima Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 
418. 1895. 
[On weatherbeaten wood (lit.)], OHIO, Morgan 1028 (HOLOTYPE: NY) 
= Gibbera confertissima (Ell. & Ev.) Sivanesan, Trans. Brit. Mycol. Soc. 65(3): 395. 1975. 


Amphisphaeria deformis Ell. & Langl., J. Mycol. 4: 123. 1888. 
On old cedar post, LOUISIANA: Pointe a la Hache, 21 Mar 1886, A. B. Langlois 1459 
(HOLOTYPE: NY; ISOTYPE: BPI) 


Amphisphaeria granulosa Ell. & Ev., J. Mycol. 10: 169. 1904. 
On old oak barrel staves lying on the ground, NEW YORK: Lyndonville, Sep 1900, Dr. C. E. 
Fairman (HOLOTYPE: NY; ISOTYPE: FH) 


Amphisphaeria hypoxylon Ell. & Ev., J. Mycol. 2: 41. 1886. 
On decaying wood, LOUISIANA: Pointe a la Hache, 30 Dec 1885, A. B. Langlois 138 
(HOLOTYPE: NY; ISOTYPE: FH) [On Carya olivaeformis (ISOTYPE: BPI)} 
= Melanomma hypoxylon (Ell. & Ev.) Cke., Grevillea 16: 53. 1887. 
= Otthia hypoxyloides (Ell. & Ev.) Ell. & Ev., N. Amer. Pyrenomyc. p. 249. 1892. 


579 


= Dimerium hypoxylon (Ell. & Ev.) Petrak, Sydowia 11: 338. 1958. 
= Immothia hypoxylon (Ell. & Ev.) Barr, Mycotaxon 29: 504. 1987. 


Amphisphaeria incrustans Ell. & Ev., N. Amer. Pyrenomyc. p. 201. 1892. 
On old wood, NEBRASKA: Milford, Oct 1887, H. G. Webber 27 (HOLOTYPE: NY) 
= Microthelia incrustans (Ell. & Ev.) Corlett & Hughes, New Zealand J. Bot. 16(3): 360. 1978. 
= Kirschteiniothelia aethiops (Berk. & Curt.) Hawksworth, Bot. J. Linn. Soc. 91(1-2): 185. 1985. 


Amphisphaena langloisii Ell. & Ev., N. Amer. Pyrenomye. p. 205. 1892. 
On decaying log of (Carya?), LOUISIANA: near St. Martinsville, 30 Jul 1889, A. B. Langlois 
2171 (HOLOTYPE: NY; ISOTYPE: FH) 


Amphisphaeria melantera Ell. & Ev., Bull. Torrey Bot. Club 24: 278. 1897. 
On bark of Quercus undulata, COLORADO: Deansbury, 24 Apr 1897, Bethel 230 
(HOLOTYPE: NY) 
= Kirschteiniella applanata (Fr.) Petrak, fide Sivanesan, Trans. Brit. Mycol. Soc. 65: 398. 1975. 
= Splanchnonema melanterum (Ell. & Ev.) Barr, Mycotaxon 15: 354. 1982. 


Amphisphaeria nuda Ell. & Ev., Erythea 2: 18. 1894. 
On Celtis occidentalis, KANSAS: Rockport, (HOLOTYPE: NY) [10 Aug 1893, Bartholomew 
1034 (ISOTYPE: FH)] 
= Lojkania nuda (Ell. & Ev.) Barr, Mycotaxon 20: 17. 1984. 


Amphisphaeria oronoensis (Ell. & Ev.) Sacc., see Sphaeria oronoensis Ell. & Ev. 
Amphisphaeria pilosella Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 418. 
1895. 


On rotten wood, OHIO, Summer 1894, [Morgan 1103 (lit.)] (HOLOTYPE: NY) 
= Gibbera pilosella (Ell. & Ev.) Sivanesan, Trans. Brit. Mycol. Soc. 65: 395. 1975. 


Amphisphaena platani Ell. & Ev., N. Amer. Pyrenomyc. p. 201. 1892. 
On bark of Platanus, LOUISIANA: near St. Martinsville, 27 Jan 1890, A. B. Langlois 2213 
(HOLOTYPE: NY; ISOTYPE: FH) 


Amphisphaeria separans Ell. & Ev., Bull. Torrey Bot. Club 24: 130. 1897. 
On old cottonwood shingle, KANSAS, 2 Apr 1896, Bartholomew 2104 (HOLOTYPE: NY) 
= Herpotrichia separans (Ell. & Ev.) Sivanesan, Mycol. Pap. 127: 10. 1971. 
= Lojkania separans (Ell. & Ev.) Barr, Mycotaxon 20: 17. 1984. 


Amphisphaeria subiculosa Ell. & Ev., J. Mycol. 2: 103. 1886. 
On decaying poplar log, LOUISIANA: Pointe a la Hache, 29 Jan 1886, A. B. Langlois 382 
(HOLOTYPE: NY; ISOTYPES: BPI, FH) 
= Herpotrichia rhodosticta (Berk. & Br.) Sacc., fide Sivanesan, Mycol. Pap. 127: 16. 1971. 
= Byssosphaeria rhodomphala (Berk.) Cke., fide Barr, Mycotaxon 20: 32. 1984. 


Anthostoma acerinum Ell. & Fairm., Proc. Rochester Acad. Sci. 4: 189. 1905. 
On Acer sp., NEW YORK: Lyndonville, Apr 1904, C. E. Fairman (ISOTYPE: FH). 


Anthostoma flavoviride Ell. & Holw., in Arth. et al., Bull. Geol. Nat. Hist. Surv. 3: 
32. 1887. 
On dead Populus tremuloides. MINNESOTA: Vermilion Lake, 25 Jul 1886, Holway 263 
(HOLOTYPE: NY). 
_ = Anthostomella flavoviridis (Ell. & Holw.) P. Martin, J. S. African Bot. 42:71. 1976. 


Anthostoma formosum Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 344. 
1894, 
On dead limb of Celtis occidentalis. KANSAS: [Rockport (lit.)] 14 Jun 1894, Bartholomew 1492 
(HOLOTYPE: NY). 


580 


= Lopadostoma formosum (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 75. 1976. 


Anthostoma microecium Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 344. 
1894, 
On dead limbs of Asimina triloba, WEST VIRGINIA: Nuttallburg, 12 Feb 1894, L. W. Nuttall 
(ISOTYPE: FH) 
[Nuttall (433) 1377 (ISOTYPE: BPI)] 
= Lopadostoma microecium (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 75. 1976. 


Anthostoma mortuosum (EIll.) Sacc., see Sphaeria mortuosa Ell. 


Anthostoma ontariense Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 228. 
1890. 
On willow [dead limbs of Salix (lit.)], CANADA: London, 1 Feb 1890, J. Dearness 1390 
(HOLOTYPE: NY, ISOTYPE: FH) 
= Eutypa ontariensis (Ell. & Ev.) Tiffany & Gilman, Iowa State Coll. J. Sci. 40: 126. 1965. 
= Cryptosphaeria subcutanea (Wahl. : Fr.) F. Rappaz, Mycotaxon 20; 581. 1984. 


Anthostoma picaceum (Cke. & Ell.) Ell. & Ev., see Sphaeria picacea Cke. & EI. 


Anthostoma saprophilum Ell. & Ev., J. Mycol. 3: 43. 1887. 
On rotten maple wood, NEW JERSEY: Newfield, May 1878 [May 1876 (lit.)], J. B. Ellis s.n. 
(HOLOTYPE & ISOTYPE: NY; ISOTYPE: FH). 


Anthostoma stictoides Ell. & Ev., nom. inval. Martin (1976) attributed this name to 
Ell. & Ev., although they never published it. 


Anthostomella albocincta Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 424. 
1895. 
On dead culms of Anindinaria, LOUISIANA: St. Martinsville, 6 Aug 1895, A. B. Langlois 
2407 (HOLOTYPE: NY; ISOTYPES: BPI, FH) 
= Entosordaria albocincta (Ell. & Ev.) Hoehn., Sber. Akad. Wiss. Wien Math.-nat. 129: 166. 1920. 


Anthostomella brachystoma Ell & Ev., Bull. Washburn Coll. Lab. Nat. Hist. 1: 5. 
1884. 


On Tsuga pattoniana, WASHINGTON: Mt. Paddo, Adams, Sep 1883, W. N. Suksdorf 113 
(ISOTYPES: BPI, FH) 
= Xylosphaeria brachystoma (Ell. & Ev.) Cke., Grevillea 17: 86. 1889. 


Anthostomella cornicola Ell. & Ev., Erythea 1: 198. 1893. 
WASHINGTON: Seattle, Aug 1892, A. M. Parker 115 (HOLOTYPE: NY) [On twigs of 
Comus, North Bend, Kings Co. (ISOTYPE: FH)] [On Comus pubescens (ISOTYPE: BPI)]} 
= Entosordaria comicola (Ell. & Ev.) Hoehn., Sber. Akad. Wiss. Wien Math.-nat. 129: 166. 1920. 


Anthostomella eructans Ell. & Ev., in. Fairm., Proc. Rochester Acad. Sci. 1: 48. 
1890 
On decorticated branch on ground, NEW YORK: Lyndonville, May 1889, Dr. C. E. Fairman 
42 (HOLOTYPE: NY) 

Anthostomella hypsophila Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 338. 
1894, 


On dead stems of Lonicera involucrata, NW COLORADO: Cameron Pass, 24 Jul 1894, C. F. 
Baker 243 (HOLOTYPE: NY; ISOTYPE: FH) 


Anthostomella leucobasis Ell. & Mart., see Sphaeria leucobasis Ell. & G. Martin. 


Anthostomella ludoviciana Ell. & Langl., Proc. Acad. Nat. Sci. Philadelphia 1890: 
228. 1890. 


581 


On dead Smilax sp. LOUISIANA: St. Martinsville, 21 Jan 1889, A. B. Langlois 1696 
(HOLOTYPE: NY; ISOTYPE: BPI) 


Anthostomella magnoliae Ell. & Ev., J. Mycol. 4: 122. 1888. 
On dejected Magnolia leaves, LOUISIANA: Pointe a La Hache, 14 Aug 1888, A. B. Langlois 
1480 (HOLOTYPE: NY; ISOTYPE: BPI) [Jul 1888, 4. B. Langlois 1480 (ISOTYPE: FH)] 
= Entosordaria magnoliae (Ell. & Ev.) Hoehn., Sitzb. Akad. Wien 129: 166. 1920. 
= Anthostomella unguiculata (Mont.) Sacc., fide Francis, Mycol. Pap. 139: 60. 1975. 


Anthostomella mammoides Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 139. 
1893. 


On Ostrya, CANADA: London, Deamess 1801 (HOLOTYPE: NY) [On Ostrya virginica, Jun 
1892 (ISOTYPE: FH)] 


Anthostomella melanosticta Ell. & Ev., J. Mycol. 3: 44. 1887. 
On old leaves of Sabal palmetto. LOUISIANA: Pointe a la Hache, Nov 1886, A. B. Langlois 
830 (HOLOTYPE: NY; ISOTYPE: FH) [23 Nov 1886 (ISOTYPE: BPI)] [Dec 1886 (lit.)] 


Anthostomella minor Ell. & Ev., J. Mycol. 3: 43. 1887. 
On petioles of Sabal serrulata, FLORIDA, Dec 1886, W. W. Calkins 746 (HOLOTYPE: NY) 
= Anthostoma minor (Ell. & Ev.) Muller, fide Muller & Dennis, Kew Bull. 19: 369. 1965. 


Anthostomella ostiolata Ell. & Ev., Bull. Torrey Bot. Club 11: 42. 1884. 
On Laurus benzoin, NEW JERSEY: Newfield, J. B. Ellis s. n. (HOLOTYPE: NY) 
= Anthostoma ostiolatum (Ell. & Ev.) Cke., Grevillea 17: 90. 1889. 


Anthostomella pholidigena (Ell.) Ell., see Sphaeria pholidigena Ell. 


Anthostomella sabalensioides (Ell. & G. Martin) Sacc., see Sphaeria sabalensioides 
Ellis & G. Martin. 


Anthostomella suberumpens Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 
338. 1894. 
On inner side of weathered elm bark, KANSAS: Rockport, 13 Nov 1893, E. Bartholomew 1244 
(HOLOTYPE: NY; ISOTYPES: BPI, FH) 


Anthostomella thyridioides Ell. & Ev., J. Mycol. 9: 167. 1903. 
On dead Populus deltoides, KANSAS: Rooks Co., 18 Jul 1902, E. Bartholomew 2969 
(ISOTYPES: BPI, FH) 


Ceriospora alabamiensis Ell. & Ev., N. Amer. Pyrenomyc. p. 391. 1892. 
On herbaceous stems, ALABAMA, # 1770 [Atkinson (lit.)] (HOLOTYPE:NY) 
= Urosporella alabamiensis (Ell. & Ev.) Barr, Mycologia 58: 690. 1966. 


Clypeosphaeria imperfecta Ell. & Ev., N. Amer. Pyrenomyc. p. 410. 1892. 
On bark of living birch trees, NEW YORK: Syracuse, Jan 1887, L. M. Underwood, N. A. F. 2: 
1960 (HOLOTYPE: NY, ISOTYPES: FH, BPI) 
= Pyrenula imperfecta (Ell. & Ev.) R. C. Harris, Mich. Botanist 12: 43. 1973. 


Clypeosphaeria minor Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 138. 
1893. 
On bark of birch roots, CANADA: London, Apr 1892, J. Dearness (HOLOTYPE: NY; 
ISOTYPE: FH) 


Clypeosphaeria sanguinea Ell. & Ev., N. Amer. Pyrenomyc. p. 409. 1892. 
On dry hard wood, PENNSYLVANIA: Jan 1880, Dr. J. W. Eckfeldt; KANSAS, Winter 1884, F. 
W. Cragin 134 (SYNTYPES: NY) 


582 


Clypeosphaeria ulmicola Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 138. 
1893. 


On Ulmus, CANADA: London, Apr 1892, Dearness 1776 (HOLOTYPE: NY; ISOTYPE: FH) 
[On dead elm limbs (ISOTYPE: BPI)} 


Cryptosphaeria fissicola (Cke. & Ell.) Sacc., see Sphaeria fissicula Cke. & Ell. 


Cryptosphaeria juglandina Ell. & Holw., Bull. Lab. Nat. Hist. Univ. 3(3): 41. 1895. 
On Juglans cinerea, IOWA: Decorah, Jun 1892, E. W. D. Holway (ISOTYPE: BPI) 
= Engizostoma juglandinum (Ell. & Holw.) Kuntze, Rev. Gen. Pl. 3(2): 474. 1898. 


Diatrype acervata Ell. & Ev., J. Mycol. 4: 75. 1888. 
On spots in leaves of Yucca filamentosa, NEW JERSEY: Newfield, 20 Jul 1888, J. B. Ellis s. n. 
(HOLOTYPE: NY; ISOTYPE: FH), and N. A. F. 2: 2124 (ISOTYPES: 
NY, BPI, FH) 
= Diaporthe acervata (Ell. & Ev.) Ell. & Ev., N. Amer. Pyrenomyc. p. 738. 1892. 
= Valsa acervata (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3 (2): 539. 1892. 
= Mycosphaerella acervata (Ell. & Ev.) Barr, Contr. Univ. Michigan Herb. 9: 596. 1972. 


Diatrype aethiops Cke. & Ell., Grevillea 6: 10. 1877. 
On Morus, NEW JERSEY: Newfield, near Malaga, Feb 1877, J. B. Ellis s. n. (ISOTYPE: BPI) 
= Valsaria aethiops (Cke. & Ell.) Sacc., Syll. Fung. 1: 745. 1882. 
Melogramma aethiops (Cke. & Ell.) Cke., Grevillea 13: 109. 1885. 
Valsaria insitiva (Tode : Fr.) Ces. & De Not., fide Ell. & Ev., N. Amer. Pyrenomye. p. 555. 
1892. 


Diatrype americana Ell. & Berl., Icon. Fung. 3: 94. 1902. 
On Magnolia, NEW JERSEY: Newfield, Jun 1874, J. B. Ellis s.n. (TYPE: BPI) 


Diatrype celastrina Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 343. 1894. 
On dead Celastrus scandens, KANSAS, 29 May 1894, Bartholomew 1472 (HOLOTYPE: NY; 
ISOTYPE: FH) 

= Valsa celastrina (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 539. 1898. 


Diatrype collariata Cke. & Ell., Grevillea 5: 32. 1876. 
On hickory, NEW JERSEY: Newfield, J. B. Ellis 539 ISOTYPES: NY, FH) 
= Valsa caryigena B. & C, fide Ell. & Ev., N. Amer. Pyrenomyc. p. 576. 1892. 
= Valsa collariata (Cke. & Ell.) Kuntze, Rev. Pl. 3(2): 539. 1898. 
= Eutypella collariata ( Cke. & Ell.) Berl., Icon. Fung. 3: 76. 1902. 


Diatrype comptoniae Ell. & Ev., J. Mycol. 2: 89. 1886. 
On old stems of Comptonia asplenifolia, NEW JERSEY: Newfield, May 1886, J. B. Ellis s. n. 
(HOLOTYPE: NY; ISOTYPE: FH) 
= Thyrnidaria comptoniae (Ell. & Ev.) Berl. & Voglino, fide Sacc., Syll. Fung. Addit. 1-4: 417. 1886. 
= Diatrypella comptoniae (Ell. & Ev.) Ell. & Ev., Cat. Pl. New Jersey p. 536. 1889. 
= Pseudovalsa comptoniae (Ell. & Ev.) Ellis & Ev., N. Amer. Pyrenomyc. p. 542. 1892. 


Diatrype cornuta Ell. & Ev., N. Amer. Pyrenomye. p. 568. 1892. 
On dead bark of Ailanthus glandulosus, # 259, NEW YORK: Lyndonville, Apr 1891, Charles 
E. Fairman (HOLOTYPE: NY) 


= Valsa comnuta [as "cornula"] (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 539. 1898. 
= Eutypella leprosa (Pers. : Fr. : Fr.) Berl., fide Rappaz, Mycologia Helvetica 2(3): 461. 1987. 


Diatrype disciformis var. americana Ell., see Diatrype americana Ell. & Berl. 


Diatrype fibritecta Cke. & Ell., Grevillea 5: 31. 1876. 


583 


On Juniperus virginiana, NEW JERSEY: Newfield, J. B. Ellis s.n. ISOTYPE: NY) 
= Valsa fibritecta (Cke. & Ell.) Kuntze, Rev. Gen. Pl. 3(2): 540. 1898. 


Diatrype hochelagae Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 224. 1890. 
On elm, "Hochelegae is an Indian name of the St. Lawrence River," [CANADA: London, Mar 
1890, J. Dearness (lit.)] (HOLOTYPE: NY) 

= Valsa hochelagae (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 540. 1898. 


Diatrype hullensis Ell. & Ev., N. Amer. Pyrenomyc. p. 567. 1892. 
On rotten wood, CANADA: Quebec, West of Hull, 14 Sep 1891, # 187 (HOLOTYPE: NY) 
[Macoun (lit.)]. 
= Eutypa flavovirens (Pers. : Fr.) Tul., fide Glawe & Rogers, Mycotaxon 20: 439. 1984. 
= Diatrype flavovirens (Pers. : Fr.) Fr., fide Rappaz, Mycologia Helvetica 2(3): 406. 1987. 


Diatrype infuscans Ell. & Ev., N. Amer. Pyrenomye. p. 571. 1892. 
On Smilax, TEXAS: Houston, 14 Apr 1869, H. W. Ravenel (HOLOTYPE: NY) 
= Eutypella quadrifida (Schwein.) Ell. & Ev., fide Rappaz, Mycologia Helvetica 2(3): 488. 1987. 


Diatrype irregularis Cke. & Ell., Grevillea 6: 92. 1878. 
On old pear tree limbs lying on the ground, NEW JERSEY: Newfield, 14 Sep 1878, J. B. Ellis 
s. n. (ISOTYPE: NY) 
= Diatrypella irregularis (Cke. & Ellis) Sacc., Syll. Fung. 1: 207. 1882. 


Diatrype lateritia Ell., Bull. Torrey Bot. Club 9: 19. 1882. 
On Carpinus, PENNSYLVANIA: W. Chester, Aug 1879, J. H. Wright (HOLOTYPE: NY) 
= Thyridaria lateritia (Ell.) Sacc., Syll. Fung. 2: 141. 1883. 
= Melogramma lateritia (Ell.) Cooke, Grevillea 13: 109. 1885. 
= Melogramma vagans De Not., fide Berl., Icon. Fung. 1: 46. 1891. 
= Melogramma campylosporum Fr., fide LaFlamme, Sydowia 28: 240. 1975 


Diatrype linearis Ell. & Ev., Bull. Torrey Bot. Club 24: 134. 1897. 
On Eucalyptus globulus, CALIFORNIA: Compton, 11 Dec 1896, McClatchie (HOLOTYPE: 
NY) 


= Diatrype prominens Cke. & Harkn., fide Rappaz, Mycologia Helvetica 2(3): 430. 1987. 


Diatrype maclurae Ell. & Ev., N. Amer. Pyrenomyc. p. 570. 1892. 
On Osage orange, CANADA: London, 11 May 1891 (HOLOTYPE: NY) [Dearness 579 
(ISOTYPE: FH)] 
= Eutypella leprosa (Pers. ex Fr. : Fr.) Berl., fide Rappaz, Mycologia Helvetica 2(3): 461. 1987. 


Diatrype macounii Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 224. 1890. 
On maple bark, BRITISH COLUMBIA, 1889, Macoun 127 (HOLOTYPE: NY) [20 May 1889 
(ISOTYPE: FH)] 

= Valsa macounii (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 540. 1898. 
= Diatrype bullata (Hoffm. : Fr.) Fr., fide Glawe & Rogers, Mycotaxon 20: 419. 1984. 


Diatrype megastoma Ell. & Ev., J. Mycol 1: 141. 1885. 
On dead alders, NEW JERSEY: Newfield, 20 Jul 1885 (ISOTYPE: FH) N. A. F. 2: 1556 
(ISOTYPES: NY, BPI) [Alnus sernulata (lit.)]. 
= Valsa megastoma (Ell. & Ev.) Kuntze, Rev. Gen. PI. 3(2): 540. 1898. 
= Eutypella cerviculata (Fr. : Fr.) Sacc, fide Rappaz, Mycologia Helvetica 2(3): 527. 1987. 


Diatrype microspora Ell., Bull. Torrey Bot. Club 8: 74. 1881. 
On alder, MAINE: Wells, Rev. J. Blake, Herb. Ellis ISOTYPE: BPI) 
= Anthostomea ellisii Sacc., Syll. Fung. 1: 308. 1882. 
= Anthostoma microsporum Karst., fide Ell. & Ev., N. Amer. Pyrenomyc. p. 582. 1892. 
= Camarops microspora (P. Karst.) Shear, Mycologia 30: 588. 1938. 


584 


Diatrype microstega Ell. & Ev., N. Amer. Pyrenomyc. p. 574. 1892. 
On bark, CALIFORNIA, Dr. Harkness 1877 (HOLOTYPE: NY) 
= Valsa microstega (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 540. 1898. 
= Diatrype prominens Cke. & Harkn., fide Rappaz, Mycologia Helvetica 2: 430. 1987. 


Diatrype minima Ell. & Ev., J. Mycol. 1: 91. 1885. 
On Magnolia, Apr 1885, J. B. Ellis s. ns (HOLOTYPE: NY) [NEW JERSEY: Newfield 
(ISOTYPE: FH)] 
= Valsa minima (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 540. 1898. 


Diatrype nigerrima Ell. & Ev., J. Mycol. 10: 170. 1904. 
On bark of Vitis, ILLINOIS: Glencoe, Jun 1893, E. T. & S. A. Harper (HOLOTYPE: NY) [Ex 
Herb. E. T. & S. A. Harper, # 904, Jun 1903 (ISOTYPE: FH)] 
= Eutypella aequilinearis (Schwein. : Fr.) Starb., fide Rappaz, Mycologia Helvetica 2: 473. 1987. 


Diatrype olivacea (Cke. & Ell.) Ell., see Diatrypella olivacea Cke. & EIl. 


Diatrype phaeosperma Ell., Amer. Naturalist 17: 195. 1883. 
On Amelanchier canadensis, IOWA: Decorah, 23 Aug 1882, E. W. Holway 228 (HOLOTYPE: 
NY; ISOTYPE: FH) 
= Anthostoma phaeospermum (Ell.) Sacc., Syll. Fung. 2: xiv. 1883. 
= Fuckelia phaeospermum (Ell.) Cke., Grevillea 12: 52. 1883. 


Diatrype pustulans Ell. & Ev., J. Mycol. 4: 80. 1888. 
On dead stems of Arundinaria, LOUISIANA: St. Martinsville, Rev. A. B. Langlois 1215 
(partly) (ISOTYPE: FH) 
= Valsaria pustulans (Ell. & Ev.) Sacc., Syll. Fung. 9: 758. 1891. 


Diatrype quercina Fr. var. lignicola Cke. & Ell., Grevillea 5: 54. 1876. 
On decorticated oak branches, NEW JERSEY: Newfield, J. B. Ellis # 2412 (lit.) 
= Cryptovalsa eutypaeformis Sacc., fide Ell. & Ev., N. Amer. Pyrenomyc. p. 517. 1892. 
= Allescherina eutypaeformis (Sacc.) Berl., Icon. Fung. 3: 110. 1905. 


Diatrype radiata Ell., Amer. Naturalist 17: 195. 1883. 
On dead limbs of elm, IOWA: Decorah, Sep 1882, E. W. Holway 266, (HOLOTYPE: NY; 
ISOTYPE: FH) 
= Valsa radiata (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 540. 1898. 
= Eutypella leprosa (Pers. ex Fr. : Fr.) Berl., fide Rappaz, Mycologia Helvetica 2: 461. 1987. 


Diatrype rhuina Cke. & Ell., Grevillea 7: 8. 1878. 
On Rhus venenata, NEW JERSEY: Newfield, Feb 1878 & Dec 1878, J. B. Ellis s. n. 
(SYNTYPES: NY, BPI) 
= Calospora rhoina (Cke. & Ell.) Sacc., Syll. Fung. 2: 234, 1883. 
= Diaporthe rhoina (Cke & Ell.) Ell. & Ev., N. Amer. Pyrenomyc. p. 424. 1892. 


Diatrype sphaerospora Ell. & Ev., J. Mycol. 3: 42. 1887. 


On Magnolia glauca, NEW JERSEY: Newfield, Jun 1878, J. B. Ellis s. n. (HOLOTYPE: NY; 
ISOTYPE: FH) 


Diatrype texensis Ell. & Ev., J. Mycol. 2: 40. 1886. 
On fallen limbs, TEXAS: Houston, Apr 1869, H. W. Ravenel (ISOTYPES: BPI, FH) 
= Thynidaria texensis (Ell. & Ev.) Berl. & Vogl., Syll. Fung. Addit. 1-4: 164. 1886. 
= Pseudovalsa texensis (Ell. & Ev.) Ell. & Ev., N. Amer. Pyrenomyc. p. 541. 1892. 


Diatrype tiliacea Ell., Amer. Naturalist 17: 195. 1883. 
On Tilia americana, IOWA: Ames, Oct 1882, J. C. Arthur 86 (HOLOTYPE: NY) 


585 


= Hercospora tiliacea (Ell.) Sacc., Syll. Fung. 9: 702. 1891. 

= Melanconis tiliacea (Ell.) Ell. & Ev., N. Amer. Pyrenomyc. p. 524. 1892. 

= Diaporthe tiliacea (Ell.) Hoehn, fide Wehm., Univ. Michigan Stud. Sci. Ser. 9: 185. 1933. 
= Phragmodiaporthe tiliacea (Ell.) Barr, Mycologia Mem. 7: 155. 1978. 


Diatrype tremellophora Ell., Amer. Naturalist 16: 239. 1882. 
On Magnolia glauca, NEW JERSEY: Newfield, 1875, J. B. Ellis s. n., "as Diatrype disciformis 
Fr. var. magnoliae Thm.” (HOLOTYPE: NY) 
= Diatrype disciformis (Hoffm.) Fr. var. magnoliae Thuem., Bull. Torrey Bot. Club 6: 95. 1896. 


Diatrype trifida Ell. & Macbr., Bull. Lab. Nat. Hist. Univ. Iowa 4: 71. 1896. 
On wood, MEXICO: Tehuantepec, 1895, C. L. Smith (HOLOTYPE: NY) 


Diatrype tumida Ell. & Ev., N. Amer. Pyrenomye. p. 567. 1892. 
On elm bark, CANADA: London, Feb 1890, J. Dearness (ISOTYPE: FH) 
= Valsa tumida (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 540. 1898. 
= Eutypella tumida (Ell. & Ev.) Wehm., Pap. Michigan Acad. Sci. 5: 179. 1926. 


Diatrype vitis (Ell. & Ev.) Sacc., see Diatrypella vitis Ell. & Ev. 


Diatrypella citricola Ell. & Ev., N. Amer. Pyrenomyc. p. 587. 1892. 
On dead orange twigs, FLORIDA: Sandford, Apr 1891, L. M. Underwood 2377 (HOLOTYPE: 
NY; ISOTYPES: NY, BPI, FH) 
= Cryptovalsa citricola (Ell. & Ev.) Berl., fide Abbado, Malpighia 16: 320. 1902. 


Diatrypella decipiens Ell. & Ev., J. Mycol. 4: 80. 1888. 
On bark of Umbellularia californica, OREGON: Coos Co., Feb 1884, W. S. Carpenter 
(HOLOTYPE: NY; ISOTYPE: FH) 


Diatrypella demetrionis Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 225. 
1890. 
On dead limbs of Salix chlorophylla, in a wet mountain valley, COLORADO, Jul 1888, 
Demetrio 205 (HOLOTYPE: FH) 


Diatrypella deusta Ell. & Martin, Amer. Naturalist 16: 809. 1882. 
On petioles of Sabal serrulata, FLORIDA, Winter 1882, Dr. Geo. Martin (HOLOTYPE: NY) 
[Green Cove Springs, Herb. S. M. Tracy (ISOTYPE: BPI)] N. A. F.: 1184 (ISOTYPES: NY, 
BPI, FH) 
= Allescherina deusta (Ell. & Martin) Berl., fide Abbado, Malpighia 16: 303. 1902. 
= Cryptovalsa deusta (Ell. & Martin) Petrak, Sydowia 7: 103. 1953. 


Diatrypella fraxini Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 426. 1895. 
On dead Fraxinus viridis, KANSAS, 22 Jul 1895, Bartholomew 1783 (HOLOTYPE: NY) 


Diatrypella herbacea Ell. & Ev., J. Mycol. 3: 42. 1887. 
On dead (short?) stems of high weeds [Ambrosia trifida (lit.)], 1 Sep 1886, LOUISIANA: St. 
Gabriel, Langlois 505 (HOLOTYPE: NY; ISOTYPES: BPI,FH). 


Diatrypella hysterioides Ell. & Ev., J. Mycol. 2: 99. 1886. 
On piece of poplar in a willow jungle, # 380, LOUISIANA, 29 Jan 1886, (HOLOTYPE: NY, 
ISOTYPE: BPI) [Langlois 380 (ISOTYPE: FH)] 

Diatrypella irregularis (Cke. & Ell.) Sacc., see Diatrype irregularis Cke. & Ell. 


Diatrypella missouriensis Ell. & Ev., N. Amer. Pyrenomyc. p. 586. 1892. 
On Corylus, MISSOURI: Concordia, Jan 1888, Rev. C. H. Demetrio 75. (HOLOTYPE: NY) 


586 


Diatrypella obscurata Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 27. 1895. 
On limbs of some shrub or tree, PENNSYLVANIA: Bethlehem, Schw. [lit.]. 


Diatrypella olivacea Cke. & Ell., Grevillea 6: 9. 1877. 
On Nyssa, NEW JERSEY: Newfield, J. B. Ellis s. n. ISOTYPE: FH) 
= Diatrype olivacea (Cke. & Ell.) Ell., Prel. Cat. Fl. New Jersey p. 198. 1881. 
= Diatrypella subfulva (B. & C.) Sacc., sec. Cke., fide Ell. & Ev. N. Amer. Pyrenomyc. p. 594. 
1892. 


Diatrypella paupera Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 404. 1893. 
On dead branch, NICARAGUA: Castillo Viejo, Feb 1893, C. L. Smith (HOLOTYPE: NY; 
ISOTYPE: BPI) 


Diatrypella populi Ell. & Holw., J. Mycol. 1: 4. 1885. 
On poplar, IOWA: Decorah, Aug 1883, Holway 349 (HOLOTYPE: NY) 


Diatrypella prunicola Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 344. 1894. 
On Prunus pennsylvanica, NEW YORK: Alcove, Feb 1894, C. L. Shear 298 (HOLOTYPE: 
NY); New York Fungi # 337 (ISOTYPES: NY, BPI, FH). 


Diatrypella pulcherrima Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 144. 
1893. 


On willow twigs and buds, Dearness 1875 (HOLOTYPE: NY) [On dead twigs of Salix, 
CANADA: London, 24 May 1892 (ISOTYPE: FH)] [possible ISOTYPE in BPI, ex Herb. 
Ellis) 

= Valsella pulcherrima (Ell. & Ev.) Berl, Icon. Fung. 3: 106. 1905. 


Diatrypella pustulata Ell. & Ev., J. Mycol. 3: 116. 1887. 
On Lonicera, NEW JERSEY: Newfield, May 1887, J. B. Ellis s. n. (HOLOTYPE: NY; 
ISOTYPE: FH) 
= Cryptovalsa pustulata (Ell. & Ev.) Ell. & Ev., N. Amer. Pyrenomyc. p. 516. 1892. 


Diatrypella ramularis Ell. & Ev., J. Mycol. 3: 42. 1887. 
On dead vines of Lonicera japonica, LOUISIANA: Pointe a la Hache, 9 Dec 1886, A. B. 
Langlois 861 (HOLOTYPE: NY; ISOTYPES: BPI, FH) 


Diatrypella sassafras Ell. & Ev., N. Amer. Pyrenomyc. p. 588. 1892. 
On dead limbs of Sassafras, NEW JERSEY: Newfield, 29 Mar 1891, J. B. Ellis s. n. 
(HOLOTYPE: NY; ISOTYPE: FH) 
= Cryptovalsa sassafras (Ell. & Ev.) Berl., fide Abbado, Malpighia 16: 311. 1902. 


Diatrypella tocciaeana De N. var. subeffusa Ell. & Ev., J. Mycol. 4: 62. 1888. 
On bark of dead alder, MASSACHUSETTS, Rev. Jos. Blake [lit.]. 
= Diatrypella verrucaeformis (Ehr. : Fr.) Nits., fide Ell. & Ev., N. Amer. Pyrenomyc. p. 584. 1892. 


Diatrypella vetusta Ell. & Ev., J. Mycol. 9: 168. 1903. 
On decorticated sticks, ILLINOIS: River Forest, Oct 1902, E. J. Harper (ISOTYPE: FH) 


Diatrypella vitis Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 225. 1890. 
On dead vines of Vitis bipinnata, LOUISIANA: Pointe a la Hache, Bayou Chene, 25 Oct 1888, 
A. B. Langlois 1508 (HOLOTYPE: NY; ISOTYPES: BPI, FH) 
= Diatrype vitis (Ell. & Ev.) Sacc., Syll. Fung. 9: 476. 1891. 
= Valsa vitis (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 540. 1898. 


Diatrypella xanthostroma Ell. & Ev., J. Mycol. 9: 225. 1903. 
On Pirus japonica, CANADA: London, 3 Nov 1903, Dearness 2045 (HOLOTYPE: NY; 
ISOTYPE: FH) [On Cydonia japonica, Rehm: Ascomycetes, # 1985 (ISOTYPE: BPI)] 


587 


Eutypa echinata Ell. & Ev., J. Mycol. 3: 43. 1887. 
On dead shoots of Fraxinus, LOUISIANA: Plaquemines Co., 29 Dec 1886, A. B. Langlois 952 
(HOLOTYPE: NY; ISOTYPE: FH) 
= Eutypa heteracantha Sacc., fide Ell. & Ev. N. Amer. Pyrenomyc. p. 100 502. 1892. 
= Engizostoma echinatum (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 474. 1898. 
= Peroneutypa heteracantha Sacc., fide Berl., Icon. Fung. 3: 81. 1968. 
= Eutypella scoparia (Schwein. : Fr.) Ell. & Ev., fide Rappaz, Mycologia Helvetica 2: 537. 1987. 


Eutypella aesculina Ell. & Ev., Erythea 1: 146. 1893. 
On Aesculus californica, CALIFORNIA: Berkeley, 23 Apr 1893, W. C. Blasdale 111 
(HOLOTYPE: NY; ISOTYPE: BPI) 
= Engizostoma aesculinum (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 473. 1898. 
= Eutypella scoparia (Schwein. : Fr.) Ell. & Ev., fide Rappaz, Mycologia Helvetica 2: 538. 1987. 


Eutypella alpina Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 425. 1895. 
On dead trunks of Alnus incana, COLORADO: Larimer Co., 11 Jul 1895, C. F. Baker 363 
(ISOTYPES: FH, BPI) N. A. F.: 3331 (ISOTYPES: NY, BPI) 
= Engizostoma alpinum (Ell. & Ev.) Kuntze, Rev. Gen. PI. 3(2): 473. 1898. 
= Eutypella cerviculata (Fr. : Fr.) Sacc., fide Rappaz, Mycologia Helvetica 2: 527. 1987. 


Eutypella amorphae Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 140. 1893. 
On Amorpha fruticosa, 12 Dec 1892, # 822 (HOLOTYPE: NY) [KANSAS: Rockport, 
Bartholomew 822 (ISOTYPES: BPI, FH)] N. A. F. 2: 2931 (ISOTYPES: NY, BPI) 

= Engizostoma amorphae (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 473. 1898. 
= Eutypella tetraploa (Berk. & Curt. ex Berk. & Broome) Sacc., fide Rappaz, Mycologia Helvetica 
2: 511. 1987. 


Eutypella canodisca (Ell. & Holw.) Sacc., see Valsa canodisca Ell. & Holw. 
Eutypella capillata (Ell. & Ev.) Ell. & Ev., see Valsa capillata Ell. & Ev. 


Eutypella carpinicola Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 342. 1894. 
On Carpinus americana, NEW YORK: Alcove, Oct 1893, C. L. Shear (HOLOTYPE: NY; 
ISOTYPE: BPI) [On Carpinus caroliniana (ISOTYPE: FH)] N. A. F.: 3028 (ISOTYPES: NY, 
FH) 

= Engizostoma carpinicola ( Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 473. 1898. 
= Massalongella carpinicola (Ell. & Ev.) Berl., Icon. Fung. 3: 1. 1968. 


Eutypella coryli Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 143. 1893. 
On Corylus, CANADA: London, Deamess 1872 (HOLOTYPE: NY) [Jun 1892 (ISOTYPE: 
FH)] 
= Engizostoma coryli (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 473. 1898. 


Eutypella densissima Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 341. 1894. 
On dead limbs of Aralia spinosa, WEST VIRGINIA: Fayette Co., 24 Feb 1894, L. W. Nuttall 
363 (HOLOTYPE: NY; ISOTYPE: FH) 
= Engizostoma densisimmum (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 473. 1898. 


Eutypella deusta (Ell. & Ev.) Ell. & Ev., see Valsa deusta Ell. & Ev. 


Eutypella exigua Ell. & Ev. ex Berl., Icon. Fung. 3: 58. 1902. 
On dead elm branches, CANADA: London, 22 Feb 1890, J. Deamess 1443 (ISOTYPE: FH) 
= Eutypella scoparia (Schwein. : Fr.) Ell. & Ev., fide Rappaz, Mycologia Helvetica 2: 538. 1987. 


Eutypella fici Ell. & Ev., Bull. Torrey Bot. Club 24: 133. 1897. 
On a dead piece of fig tree, LOUISIANA: St. Martinsville, 26 Mar 1896, A. B. Langlois 2443 
(HOLOTYPE: NY; ISOTYPES: BPI, FH) 


588 


= Eutypella scoparia (Schwein. : Fr.) Ell. & Ev., fide Rappaz, Mycologia Helvetica 2: 538. 1987. 


Eutypella herbicola Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 426. 1895. 
On Aster cordifolius, OHIO, Morgan 1122 (ISOTYPE: FH) 
= Engizostoma herbicola (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 474. 1898. 
= Eutypella goniostoma (Schwein.) Sacc., fide Rappaz, Mycologia Helvetica 2: 491. 1987. 


Eutypella juglandina (Cke. & Ell.) Sacc., see Valsa juglandina Cke. & Ell. 


Eutypella lutescens (Ell.) Sacc., see Valsa lutescens Ell. 
Eutypella maclurae (Cke. & Ell.) Ell. & Ev. See Valsa maclurae Cke. & Ell. 
Eutypella microcarpa (Ell. & Ev.) Sacc. See Valsa microcarpa Ell. & Ev. 


Eutypella populi Ell. & Ev., Amer. Naturalist 31: 342. 1897. 
On poplar twigs, CANADA, 29 Sep 1896, # 113, Macoun (HOLOTYPE: NY) [On dead 
limbs of Populus (lit.)] 


Eutypella rugiella (Cke. & Ell.) Sacc., see Valsa rugiella Cke. & EII. 


Eutypella sarcobati Ell. & Ev., Bull. Torrey Bot. Club 24: 462. 1897. 
On Sarcobatus vermiculatus, COLORADO: Alamosa, 12 Jul 1897, Bethel 324 (HOLOTYPE: 
NY; ISOTYPES: BPI, FH) 


Eutypella tiliae Ell. & Ev., Bull. Torrey Bot. Club 24: 280, 1897. 
On dead wood, Canada, # 432, 22 Oct 1896, Macoun (YHOLOTYPE: NY). [On bark of dead 
Tilia americana, (lit.)]. 


Eutypella venusta (Ell.) Sacc., see Valsa venusta Ell. 
Hypoxylon albocinctum Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 229. 
1890. 


On bark of dead Crataegus, OHIO: Preston, Hamilton Co., 10 Jan 1890, A. P. Morgan 884 
(HOLOTYPE: NY). [On bark of Ostrya virginica, (ISOTYPE: FH)] 

Ostrya virginica was originally given as the host on the holotype packet (NY); Crataegus was 
later written over that name. 


Hypoxylon atrorufum Ell. & Ev., N. Amer. Pyrenomyce. p. 742. 1892. 
On bark, MICHIGAN, Jan 1892, G. H. Hicks 165 (HOLOTYPE: NY) 
= Hypoxylon cohaerens Pers. ex Fr., fide Miller, A Monograph of the world species of Hypoxylon, 
p. 46, 1961. [According to Ellis (1892) the host is oak(?). According to Cash (1953), and Shear 
(1946), the host is Fagus]. 


Hypoxylon atroviride Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 346. 1894. 
On bark of dead trees (birch or oak), WEST VIRGINIA: Nuttallburg, L. W. Nuttall 275. 
(HOLOTYPE: NY) [16 Dec 1893 (ISOTYPE: FH)] [L. Nuttall 1320 VJ. B. E. 275) ISOTYPE: 
BPI] 

= Bolinia tubulina (Alb. & Schw. : Fr.) Sacc., Syll. Fung. 1: 352. 1882. 
= Camarops tubulina (Alb. & Schw. : Fr.) Shear, Mycologia 30: 568. 1938. 
= Camarops ohiensis (Ell. & Ev.) Nannfeldt, Svensk Bot. Tidskr. 66(4): 365. 1972. 


Hypoxylon bicolor Ell. & Ev., J. Mycol. 2: 88. 1886. 
On dead branches of Quercus virens, LOUISIANA: Pointe a la Hache, A. B. Langlois 344 
(HOLOTYPE: NY; ISOTYPE: FH) 
= Hypoxylon rubiginosum Pers. : Fr., fide Miller, A Monograph of the world species of Hypoxylon, 
p. 29, 1961. 


589 


= Hypoxylon bicoloratum P. Martin, J. S. African Bot. 42: 72. 1976. 
Hypoxylon californicum Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 426. 
1895. 


On wood of Adenostylum fasciculatum, CALIFORNIA: [Pasadena (lit.)], 5 Aug 1894, A. J. 
McClatchie 755 (HOLOTYPE: NY) 


= Hypoxylon rubrostromaticum Mill., A Monograph of the world species of Hypoxylon, p. 24, 
1961. 


Hypoxylon cinereum Ell. & Ev., N. Amer. Pyrenomyc. p. 647. 1892. 
On decaying pieces, LOUISIANA: near St. Martinsville, 28 Mar 1890, A. B. Langlois 2278 
(HOLOTYPE: NY; ISOTYPES: BPI, FH) 
= Hypoxylon hypophlaeum (Berk. & Rav.) Mill., Mycologia 33: 75. 1941. 


Hypoxylon commutatum Nits. subsp. holwayanum Sacc. & Ell., Michelia 2: 570. 
1882. 


On oak, # 3679, IOWA: Decorah, Jul 1883, E. W. Holway (HOLOTYPE: NY). [On plum, 21 
Jul 1883 (ISOTYPE: FH)] 
= Hypoxylon rubiginosum Pers. : Fr., fide Miller, A Monograph of the world species of Hypoxylon, 
p. 29. 1961. 


Hypoxylon cylindrophorum Ell. & Ev., Bull. Iowa Univ. Lab. Nat. Hist. 2: 407. 1893. 
On dry hard wood, NICARAGUA: take Nicaragua, Island of Ometepe, Winter 1893, Central 
American Fungi (C. A. F.) # 82, C. L. Smith (HOLOTYPE: NY) 

= Camarops polyspermum (Mont.) Miller, Trans. Brit. Mycol. Soc. 15: 151. 1930. 
= Numulariola cylindrophora (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 76. 1976. 


Hypoxylon discoideum Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 450. 
1893. 


On dead stem of climbing Rhus toxicodendron, NEW JERSEY: Newfield, Dec 1892, J. B. Ellis 
s.n. (HOLOTYPE: NY) [In the swamp (ISOTYPE: FH)] 

= Hypoxylon investiens (Schw.) Curt., fide Miller, A Monograph of the world species of 
Hypoxylon, p. 49. 1961. 


Hypoxylon fibuliforme Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 4: 71. 1896. 
[On dead wood (lit.)], NICARAGUA: Indian River, Mar 1896, C. L. Smith 12 (HOLOTYPE: 
NY) [near Greytown, Charles L. Smith 131, Central American Fungi (ISOTYPE: BPI)]. 
According to Miller (1961) the type specimen is too old to make a determination and the 
species is excluded. 


Hypoxylon holwayii Ell., Amer. Naturalist 17: 193. 1883. 
On poplar, IOWA: Decorah, Jul 1882, E. W. Holway 145 (HOLOTYPE: NY), and N. A. F.: 
1182 (ISOTYPES: NY, FH). 
= Hypoxylon mammatum (Wahl.) Miller, A Monograph of the world species of Hypoxylon, p.64. 
1961. 


Hypoxylon lucidum Ell. & Ev., Bull. Iowa Univ. Lab. Nat. Hist. 4: 72. 1896. 
On dead wood, NICARAGUA (C. L. Smith) [lit.]. 
= Hypoxylon haematostroma Mont., fide Miller, A Monograph of the world species of Hypoxylon, 
p. 36. 1961. 


Hypoxylon morgani Ell. & Ev., N. Amer. Pyrenomyc. p. 648. 1892. 
On rotten wood, OHIO, A. P. Morgan 890 (HOLOTYPE: NY) 


Hypoxylon multiforme Fr. var. effusum Cke. & Ell., Grevillea 5: 33. 1876. 
On maple, NEW JERSEY: Newfield, Oct 1876, J. B. Ellis 569 ISOTYPE: NY) 
= Sphaeria multiformis Fr. f. effusa Fr., Syst. Mycol. 2: 334. 1823. 


590 


= Hypoxylon rubiginosum Pers. : Fr., fide Miller, A Monograph of the world species of Hypoxylon, 
p. 29. 1961. 


Hypoxylon nicaraguense Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 407. 1893. 
On wood, NICARAGUA: Ometepe (a), and near Castillo Viejo (b), Feb 1893, C. A. F. 26, B. 
Shimek (SYNTYPES: NY, BPI) 
= Hypoxylon sclerophaeum Berk. & Curt., fide Miller, A Monograph of the world species of 
Hypoxylon, p. 42. 1961. 


Hypoxylon nuttallii Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 346. 1894. 
On Magnolia fraseri, WEST VIRGINIA: Nuttallburg, 4 May 1894, L. W. Nuttall 477 
(HOLOTYPE: NY; ISOTYPE: BPI) 

= Hypoxylon rubiginosum Pers. : Fr., fide Miller, A Monograph of the world species of Hypoxylon, 
p. 30. 1961. 


Hypoxylon occidentale Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 345. 
1894, nom Ell. & Morg. ex Martin, J. S. African Bot. 42: 82. 1976. 
On dead limbs, CALIFORNIA: Pasadena, Mar 1894, Prof. A. J. McClatchie (HOLOTYPE: 
NY; ISOTYPE: FH) and N. A. F. 2: 3127 (ISOTYPES: NY, FH). 
= Hypoxylon thouarsianum (Lév.) Lloyd, Myc. Writ. 5: 26. 1919. 


Hypoxylon occidentale Ell. & Morgan ex Martin, J. S. African Bot. 42: 82. 1976. 
On Liriodendron, OHIO: Morgan 621 (HOLOTYPE NY) 


Hypoxylon ohiense Ell. & Ev., N. Amer. Pyrenomyc. p. 648. 1892. 
On rotten wood, OHIO: A. P. Morgan 883 (LECTOTYPE: NY) and 965 (PARATYPE: NY) 
= Bolinia tubulina (Alb. & Schw.: Fr.) Sacc., Syll. Fung. 1: 352. 1882. 
= Camarops ohiensis (Ell. & Ev.) Nannfeldt, Svensk Bot. Tidskr. 66(4): 365. 1972. 


Hypoxylon pallidum Ell. & Ev., J. Mycol. 4: 68. 1888. 
{On bark of dead oak limbs, LOUISIANA (lit.)] Catahoula, Langlois 1273, ex Herb. A. B. 
Langlois (ISOTYPE: BPI) 
= Hypoxylon notatum Berk. & Curt., fide Miller, A Monograph of the world species of Hypoxylon, 
p. 20. 1961. 


Hypoxylon papillatum Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 408. 1893. 
On decaying wood of various deciduous trees, KANSAS, Winter 1884, F. W. Cragin 140. 
OHIO, Autumn 1883, A. P. Morgan 294. DELAWARE, Jul 1893, A. Commons 2160. 
NICARAGUA, Winter 1893, C. L. Smith 57 (SYNTYPES: NY) 


Hypoxylon piceum Ell., Amer. Naturalist 17: 194. 1883. 
On rotten wood, IOWA: Decorah, Oct 1882, E. W. Holway 289 (HOLOTYPE: NY) [E. W. 
Holway 287 (lit.)]. 
= Hypoxylon rubiginosum Pers.: Fr., fide Miller, Bothalia 4: 258. 1942. 
= Hypoxylon sclerophaeum Berk. & Curt., fide Miller, A Monograph of the world species of 
Hypoxylon, p. 42. 1961. 


Hypoxylon platystomum Ell. & Ev., N. Amer. Pyrenomyce. p. 649. 1892. 
On cross cuttings of Melia, LOUISIANA: St. Martinsville, 25 Oct 1890, A. B. Langlois 2333 
(HOLOTYPE: NY; ISOTYPE: FH) 


= Hypoxylon stygium (Lév.) Sacc., fide Miller, A Monograph of the world species of Hypoxylon, p. 
91. 1961. 


Hypoxylon subchlorinum Ell. & Calkins, J. Mycol. 4: 86. 1888. 
On bark of dead limbs of some deciduous tree, FLORIDA: Jacksonville, Winter 1886, W. W. 
Calkins, N. A. F. 2: 2115 (ISOTYPES: NY, FH) 


= Hypoxylon rubiginosum Pers. : Fr., fide Miller, A Monograph of the world species of Hypoxylon, 
p. 27. 1961. 


591 


Hypoxylon subluteum Ell. & Ev., N. Amer. Pyrenomyc. p. 648. 1892. 
On decaying soft wood, LOUISIANA: Near St. Martinsville, 24 Aug 1890, A. B. Langlois 2276 
(HOLOTYPE: NY) 
= Hypoxylon rubiginosum Pers. : Fr., fide Miller, Bothalia 4: 258. 1942. 
= Hypoxylon serpens (Pers. : Fr.) Kickx, fide Miller, A Monograph of the world species of 
Hypoxylon, p. 80. 1961. 
= Hypoxylon bipapillatum Berk. & Curt., fide Petrini & Rogers, Mycotaxon 26: 408. 1986. 


Hypoxylon vernicosum Ell. & Ev., Amer. Naturalist 31: 426. 1897. 
On rotten wood, sent from Ohio as H. marginatum, OHIO: A. P. Morgan (HOLOTYPE: NY) 
= Hypoxylon truncatum (Schw. : Fr.) Mill., Trans. Brit. Mycol. Soc. 17: 130. 1932. 


Kretzschmaria pusilla Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 410. 1893. 
On fallen tree trunks, NICARAGUA: Castillo Viejo, Feb 1893, C. A. F. 29, C. L. Smith 
(HOLOTYPE: NY) [C. L. Smith 72 (ISOTYPE: BPI)] 
= Kretzschmaria clavus (Fr.) Sacc., Syll. Fung. 2: 29. 1883. 


Kretzschmania spinifera Ell. & Macbr., Bull. Iowa Univ. Lab. Nat. Hist. 4: 71. 1896. 
On decaying bark, MEXICO, 1895, C. L. Smith (HOLOTYPE: NY; ISOTYPES: NY, BPI) 


Nummulania albosticta Ell. & Morg., Bull. Torrey Bot. Club 24: 135. 1897. 
On hickory, OHIO: [Preston (lit.)], A. P. Morgan 1178 (HOLOTYPE: NY) [On dead Carya 
(ISOTYPE: FH)] 
= Hypoxylon albostictum (Ell. & Morg.) Miller, Mycologia 25: 325. 1933. 
= Numulariola albosticta (Ell. & Morg.) P. Martin, J. S. African Bot. 42: 75. 1976. 


Nummulania lateritia Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 144. 1893. 
On bark of dead Fraxinus sambucifolia, CANADA: London, Mar 1892, J. Dearness 1283 
(ISOTYPE: FH). 

= Kommamyce lateritia (Ell. & Ev.) Niewland, Amer. Midl. Naturalist 4: 375. 1916. 
= Hypoxylon rubiginosum Pers. : Fr., fide Miller, A Monograph of the world species of 
Hypoxylon, p. 27. 1961. 


Nummularia pezizoides Ell. & Ev., Bull. Torrey Bot. Club 11: 74. 1884. 
On bark, CANADA: Ottawa, Oct 1883, Macoun (HOLOTYPE: NY) 
= Biscogniauxia pezizoides (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 2: 398. 1891. 
= Nummularia repanda (Fr.) Nits., fide Jong & Benjamin, Mycologia 63: 869. 1971. 


Nummularia repanda (Fr.) Nits. var. zonata Ell. & Ev., Bull. lowa Univ. Lab. Nat. 
Hist. 2: 405. 1893. 
On dead tree(?), NICARAGUA: Castillo Viejo, Feb 1893, C. A. F. 18, C. L. Smith 
(HOLOTYPE, ISOTYPE: NY) [C. L. Smith 105 (ISOTYPE: BPI)]} 


Nummularia rufa Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 406. 1893. 
On bark, NICARAGUA: Ometepe, Winter 1893, C. L. Smith 54 (HOLOTYPE: NY; 
ISOTYPE: BPI) 
= Hypoxylon stygium (Lév.) Sacc., fide Miller, A Monograph of the world species of Hypoxylon, p. 
92. 1961. 


Nummularia subapiculata Ell. & Ev., J. Mycol. 5: 23. 1889. 
On bark, KANSAS: Topeka, 1884, F. W. Cragin 267 (HOLOTYPE: NY) 
- = Hypoxylon tinctor (Berk.) Cke., fide Miller, A Monograph of the world 
species of Hypoxylon, p. 119. 1961. 


Nummularia vernicosa Ell. & Ev., Bull. Iowa Univ. Lab. Nat. Hist. 2: 406. 1893. 
On dead branch, NICARAGUA: Castillo Viejo, Feb/Mar 1893, C. A. F. 21, C. L. Smith 
(HOLOTYPE: NY) [Smith 117 (ISOTYPE: BPI)] 


ees 


= Hypoxylon melanaspis Mont., fide Miller, A Monograph of the world species of Hypoxylon, p. 
107. 1961. 


Phomatospora wistariae Ell. & Ev., J. Mycol. 8: 68. 1902. 


On Wistaria frutescens, ALABAMA: Carver 686 (HOLOTYPE: NY) [22 Sep 1900 (ISOTYPE: 
FH)] 


Podosordaria mexicana Ell. & Holw., Bot. Gaz. 24: 37. 1897. 


On cow dung, MEXICO: Cuernavaca, Sep 1896, ex- Herb. Holway, E. W. Holway 
(HOLOTYPE: NY) 


= Xylaria chardoniana (Toro) Miller, fide Dennis, Kew Bull. 1957: 306. 1957. 


Poronia leporina Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 229. 1890. 
On rabbit dung, MISSOURI: Emma, Sep 1889, Demetrio 250 (HOLOTYPE: NY; ISOTYPE: 
BPI) and N. A. F.: 2354 (ISOTYPES: NY, FH). 
= Podosordaria leporina (Ell. & Ev.) Dennis, Kew Bull. 1957. 306. 1957. 
= Poronia minuta Petch, fide Krug & Cain, Canad. J. Bot. 52: 596. 1974. 


Poronia turbinata Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 410. 1892. 
On bark, NICARAGUA: Castillo Viejo, Feb/Mar 1893, C. A. F. 32, C. L. Smith 
(HOLOTYPE: NY; ISOTYPE: BPI) 
= Kretzschmaria turbinata (Ell. & Ev.) Lloyd, Myc. Writ. 6: 1033. 1921. 


= Penzigia turbinata (Ell. & Ev.) Miller, A Monograph of the world species of Hypaxylon, p. 142. 
1961. 


= Kretzschmaria turbinata (Ell. & Ev.) P. Martin, J. S. African Bot. 36: 79. 1970. 


Rosellinia abietina Fuckel var. trichota (Cke. & Ell.) Sacc., see Sphaeria abietina 
Fcekl. var. trichota Cke. & Ell. 


Rosellinia albolanata Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 227. 1890. 
On old rails, MISSOURI: Emma, Nov 1889, Rev. C. H. Demetrio 269 (HOLOTYPE: NY; 
ISOTYPE: FH) 


= Hypoxylon albolanatum (Ell. & Ev.) P.Martin, J. S. African Bot. 42: 72. 1976. 
Rosellinia arctispora (Cke. & Ell.) Sacc., see Sphaeria arctespora Cke. & Ell. 


Rosellinia bakeri Ell., Torreya 5: 87. 1905. 


On Urera, NICARAGUA, 1903, C. F. Baker 3990 (HOLOTYPE: NY) [Chinandega 
(ISOTYPE: FH)] [Dec 1903 (lit.)]. 


= Hypoxylon investiens (Schw.) Berk. f. bakeri (Ell.) Miller, fide Dennis, Kew Bull. 14(3): 450. 
1960. 


Rosellinia bicolor Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 327. 1894. 
On rotten pieces, LOUISIANA: near St. Martinsville, 26 Aug 1889, A. B. Langlois 2139 
(HOLOTYPE: NY) 


Rosellinia bigeloviae Ell. & Ev., Amer. Naturalist 31: 341. 1897. 


On Bigelovia graveolens, COLORADO: E. Bethel (HOLOTYPE: NY) [Golden, Dec 1896 
(ISOTYPE: BPI)] 


Rosellinia caespitosa Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 327. 1894. 
On dead hackberry limbs, KANSAS: Rockport, 22 Nov 1893, E. Bartholomew 1252 
(HOLOTYPE: NY) 


= Lopadostoma caespitosum (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 75. 1976. 


Rosellinia compressa Ell. & Dearn., Proc. Canad. Inst. n. ser. 1: 89. 1897. 
On elm, CANADA: Ontario, near Granton, J. Deamess 1791 (HOLOTYPE: NY) 


ao 


Rosellinia confertissima Ell. & Ev., Bull. Torrey Bot. Club 24: 126. 1897. 
On rotten wood, OHIO: Morgan 1172 (ISOTYPE: FH) [Morgan 1173 (lit.)] 


AGE geasteroides Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 415. 
1895. 


On the base of decaying stem of Arundinaria, LOUISIANA: near St. Martinsville, 17 Jul 1895, 
A. B. Langlois 2404 (HOLOTYPE: NY; ISOTYPES: BPI, FH) 

= Astrocystis mirabilis B. & Br., fide Diehl, Mycologia 17: 188. 1925. 

= Hypoxylon geasteroides (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 72. 1976. 


Rosellinia gigantea Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 401. 1893. 
On bark, NICARAGUA: Castillo Viejo, Winter 1893, C. L. Smith 34 (HOLOTYPE: NY; 
ISOTYPE: FH), C. A. F. 10 (ISOTYPES: NY, BPI) 
= Hypoxylon giganteum (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 72. 1976. 


Rosellinia gigaspora Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 401. 1893. 
On bamboo cane(?), NICARAGUA: Castillo Viejo, Feb/Mar 1893, C. L. Smith 
(HOLOTYPE: NY) 


= Hypoxylon gigasporum (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 72. 1976. 
Rosellinia glandiformis Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 227. 
1890. 


On live oak stump (Quercus), LOUISIANA: St. Martinsville, Feb 1889, A. B. Langlois 1768 
(HOLOTYPE: NY; ISOTYPE: FH) 
= Hypoxylon glandiforme (Ell. & Ev.) P. Martin, J. S. African Bot. 34: 190. 1968. 


Rosellinia hystrix Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 444. 1894. 
On old hickory nuts lying on the ground, NEW JERSEY: Newfield, 18 May 1893, J. B. Ellis s. 
n. (HOLOTYPE: NY; ISOTYPE: FH) 


Rosellinia kellermannii Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 227. 
1890. 


On very rotten wood of Negundo aceroides, KANSAS: Manhattan, 24 Mar 1889, # 1378 
(HOLOTYPE: NY) [Kell. & Swingle 1378 (ISOTYPE: FH)] 
= Coniochaeta kellermannii (Ell. & Ev.) Munk, Dansk Bot. Ark. 15(2): 54. 1953. 


Rosellinia langloisii Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 228. 1890. 
On much rotten vine of Vitis vulpina, LOUISIANA: St. Martinsville, 26 Mar 1889, A. B. 
Langlois 1679 (HOLOTYPE: NY; ISOTYPES: BPI, FH). [A. B. Langlois 1779 (lit.)] 

= Hypoxylon langloisii (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 73. 1976. 


Rosellinia limoniispora Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 326. 
1894. 


On dead limbs of Fraxinus, KANSAS: Rockport, 4 Aug 1894, Bartholomew 1545 
(HOLOTYPE: NY) 


= Hypoxylon limoniisporum (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 73. 1976. 


Rosellinia macouniana Ell. & Ev., Bull. Torrey Bot. Club 11: 74. 1887. 
On rotten wood, CANADA: Ottawa, Oct 1883, Macoun 253 [lit.]. 
= Byssosphaeria macouniana (Ell. & Ev.) Cke., Grevillea 15: 122, 1887. — 


Rosellinia macra Ell. & Ev., Bull. Torrey Bot. Club 24: 126. 1897. 


On leaves of monocot plant, FLORIDA, 1895, A. P. Morgan 1134 (HOLOTYPE: NY) [On 
leaf of (Sabal?) ISOTYPE: FH] 


Rosellinia megaloecia Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893. 128. 
1893. 


594 


On Salix, MONTANA: One mile west of Sheridan, Mill Creek, 20 Jan 1892, W. M. Fitch 9 
(HOLOTYPE: NY) 


Rosellinia melaleuca Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 402. 1893. 
On wood, NICARAGUA: Ometepe, Jan 1893, B. Shimek, C. A. F. 11 (HOLOTYPE: NY; 
ISOTYPE: BPI). 


Rosellinia muriculata Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 416. 1895. 
On fibrous cottonwood bark (Popiilus monilifera), KANSAS: Rooks Co., 21 Dec 1894, E. 
Bartholomew 1613 (HOLOTYPE: NY; ISOTYPES: BPI, FH) 


Rosellinia obliquata (Somm.) Sacc. var. americana Ell. & Ev., N. Amer. Pyrenomyc. 


p. 169. 1892 
On dead cones of Pinus ponderosa var. scopulorum, MONTANA: Belt Mts., Sep 1889, F. W. 
Anderson 613 (HOLOTYPE: NY; ISOTYPES: NY, BPI) 


Rosellinia ostiolata Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 327. 1894. 
On bark of Ulmus americana, KANSAS: Rockport, 2 Apr 1894, E. Bartholomew 1429 
(HOLOTYPE: NY; ISOTYPE: FH) 


Rosellinia ovalis (Ell.) Sacc., see Sphaeria ovalis Ell. 


Rosellinia parasitica Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1890: 227. 1890. 
On dead limbs of Symphoricarpus occidentalis, MONTANA: Helena, Rev. F. D. Kelsey, N. A. 
F, 2: 2351 (ISOTYPES: NY, BPI, FH) [Elisens (1985) mentioned a holotype in NY, but it was 
not found]. 


Rosellinia pinicola Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1894: 327. 1894. 
On weathered pine board, KANSAS: Rockport, 15 Feb 1894, E. Bartholomew 379 
(HOLOTYPE: NY) 


Rosellinia poliosa Ell. & Ev., Bull. Torrey Bot. Club 22: 439. 1895. 
On decaying post, MEXICO: Monterey, Jun 1895, Dr. B. F. G. Egeling 50 (HOLOTYPE: NY) 
= Hypoxyion poliosum (Ell. & Ev.) P. Martin, J. S. African Bot. 42: 73. 1976. 


Rosellinia pulcherima Ell, & Ev., Erythea 1: 197. 1893. 
On detached fir bark partly covered with soil, WASHINGTON: Seattle, 18 Sep 1892, A. M. 
Parker 160 (HOLOTYPE: NY; ISOTYPE: FH) 
= Hypoxylon pulchrum P. Martin, J. S. African Bot. 42: 82. 1976. 


Rosellinia subcompressa Ell. & Ev., Bull. Torrey Bot. Club 24: 277. 1897. 
On dead cottonwood twigs, SOUTH DAKOTA: Aberdeen, Apr 1897, D. Griffiths 7 
(HOLOTYPE: NY; ISOTYPE: FH) [On Populus deltoidea Marsh. (ISOTYPE: BPI)] 


Rosellinia thelena (Fr.) Rabh. var. terrestris Ell. & Ev., Proc. Acad. Nat. Sci. 
Philadelphia 1895: 22. 1895. 
On ground, NEW YORK, (Schw-.) [lit.]. 


Rosellinia trichota (Cke. & Ell.) Ell. & Ev., see Sphaeria abietina (Fckl.) var. 
trichota Cke. & Ell. 


Rosellinia xylarispora (Cke. & Ell.) Sacc., see Sphaeria xylariaespora Cke. & Ell. 


Sphaeria abietina Fuckel var. trichota Cke. & EIll., Grevillea 6: 13. 1877. 
On Jersey pine lumber lying on the ground, NEW JERSEY: Gloucester Co., Newfield, 27 Apr 
1876, Cooke 2401. On Jersey pine pole in the grape trellis, NEW JERSEY, Mar 1877. On 


smooth cut surface of a white cedar stump, NEW JERSEY: Gloucester Co., Newfield, Apr 
1877 (SYNTYPES: NY) 


a), 


= Rosellinia abietina Fuckel var. trichota (Cke. & Ell.) Sacc., Syll. Fung. 1: 272. 1882. 
= Rosellinia trichota (Cke. & Ell.) Ell. & Ev., N. Amer. Pyrenomye. p. 172. 1892. 


Sphaeria antiqua Ell. & Ev., Bull. Torrey Bot. Club 10: 90. 1883. 
On loose bark of grape-vines, NEW JERSEY: Newfield, May 1883, J. B Ellis s. n. (ISOTYPE: 
FH) 
= Thyridium antiquum (Ell. & Ev.) Berl. & Vogl., Syll. Fung. Addit. 1-4: 185. 1886. 
= Pleospora antiqua (Ell. & Ev.) Ell. & Ev., Cat. Pl. N. J. p. 523. 1890. 
= Xylosphaeria antiqua (Ell. & Ev.) Petr., Sydowia 4: 18. 1950. 
= Mycothynidium antiquum (Ell. & Ev.) Petr., Sydowia 15: 289. 1961. 


Sphaeria arctespora Cke. & Ell., Grevillea 5: 93. 1877. 
On Andromeda (inside the bark), NEW JERSEY: Vineland, 12 Nov 1876, J. B. Ellis s. n., ex 
Herb. W. G. Farlow, ‘type gathering’ (ISOTYPES: NY, FH, BPI) 
= Roseilinia arctispora (Cke. & Ell.) Sacc., Syll. Fung. 1: 268. 1882. 


Sphaeria atrograna Cke. & Ell., Grevillea 8: 15. 1879. 
On rotten Liquidambar, NEW JERSEY: Malaga, 14 Sep 1878, # 3179 (ISOTYPE: NY) 
= Amphisphaeria atrograna (Cke. & Ell.) Sacc., Syll. Fung. 1: 722. 1882. 
= Melanomma atrogranum (Cke. & Ell.) Cke., Grevillea 16: 52. 1887. 
= Gibbera atrograna (Cke. & Ell.) Sivanesan, Trans. Br. Mycol. Soc. 65: 396. 1975. 


Sphaeria bisphaerica Cke. & Ell., Grevillea 7: 41. 1878. 
On an old apple tree limb lying on the ground, NEW JERSEY: Newfield, 10 Jun 1878, # 3068, 
“no duplicate". (~HOLOTYPE: NY) 
= Amphisphaeria bisphaerica (Cke. & Ell.) Sacc., Syll. Fung. 1: 721. 1882. 


Sphaeria fissicula Cke. & Ell., Grevillea 6: 94. 1878. 
On Rosa, NEW JERSEY: Newfield, # 2683, J. B. Ellis (lit.) 
= Cryptosphaenia fissicola (Cke. & Ell.) Sacc., Syll. Fung. 1: 185. 1882. 
= Engizostoma fissicola (Cke. & Ell.) Kuntze, Rev. Gen. Pl. 3(2): 474. 1898. 


Sphaeria leucobasis Ell. & G. Martin, Amer. Naturalist 16: 809. 1882. 
On petioles of Sabal serrulata, FLORIDA, Winter 1882, Dr. G. Martin (HOLOTYPE: NY) 
[Green Cove Springs (ISOTYPE: BPI)] N. A. F.: 1199 (ISOTYPES: NY, BPI, FH) 
= Anthostomella leucobasis Ell. & G. Mart., N. Amer. Pyrenomyc. p. 418. 1892. 


Sphaeria mortuosa Ell., Bull. Torrey Bot. Club 9: 73. 1882. 

On Eupatorium purpureum, on old Polygonum, on old half decayed Andropogon, NEW 
JERSEY: Gloucester Co., Newfield, Oct 1879, Apr 1880, 23 Sep 1879, respectively 
(SYNTYPES: NY) N. A. F.: 897 (ISOSYNTYPE: NY, FH, BPI) 

= Anthostoma mortuosum (EIl.) Sacc., Syll. Fung. 9: 519. 1891. 

= Xylosphaeria mortuosa (Ell.) Cke., Grevillea 17: 86. 1889. 

= Anthostomella xylostei (Sacc.) P. Martin, J. S. African Bot. 35: 404. 1969. 

= Anthostomella tomicoides Sacc., fide Francis, Mycol. Pap. 139: 52. 1975. 


Sphaeria oronoensis Ell. & Ev., J. Mycol. 3: 117. 1887. 
On rotten wood of some coniferous tree, MAINE: Orono, Nov 1886, F. L. Harvey 57 
(HOLOTYPE: NY; ISOTYPE: BPI) 
= Amphisphaeria oronoensis (Ell. & Ev.) Sacc., Syll. Fung. 9: 746. 1891. 


Sphaeria ovalis Ell., Bull. Torrey Bot. Club 8: 125. 1881. 
On sage brush, UTAH: Pleasant Valley, 1881, S. J. Harkness (HOLOTYPE: NY), and N. A. 
F.: 896 (ISOTYPE: NY, BPI, FH) 
= Rosellinia ovalis (Ell.) Sacc., Syll. Fung. 1: 269. 1882. 
= Rosellinia pulveracea (Ehr.) Fckl., fide Riley, Torreya 1: 22. 1901. 


596 


Sphaeria pholidigena Ell., Bull. Torrey Bot. Club 10: 54. 1883. 
On cones of red pine, UTAH: Pleasant Valley, Feb 1882, S. J. Harkness 90 (HOLOTYPE: 
NY) and N. A. F. 2: 1664 (ISOTYPES: NY, FH) 
= Anthostomella pholidigena (Ell. ) Berl. & Vogl., Syll. Fung. Addit. 1-4: 44. 1886. 
= Anthostoma pholidigena (Ell.) Cke., Grevillea 17: 90. 1889. 
= Lopadostoma conorum (Fckl.) P. Martin, J. S. African Bot. 35: 400. 1969. 


Sphaeria picacea Cke. & EIll., Grevillea 7: 9. 1878. 
On decorticated Vaccinium and Acer, NEW JERSEY: Newfield, J. B. Ellis s. n. ISOTYPE: 
NY), and N. A. F.: 183 (ISOTYPE: NY, FH, BPI) 
= Anthostomella picacea (Cke. & Ell.) Sacc., Syll. Fung. 1: 293. 1882. 
= Anthostoma picaceum (Cke. & Ell.) Ell. & Ev., N. Amer. Pyrenomyc. p. 578. 1892. 
= Anthostomella inconspicua P. Martin, J. S. African Bot. 35: 398. 1969. 


Sphaeria sabalensioides Ell. & G. Martin, Amer. Naturalist 16: 810. 1892. 
On Sabal, FLORIDA: Green Cove Springs, Winter 1881-1882, Dr. Martin (HOLOTYPE: NY) 
[On Sabal serrulata (lit.)} 
= Anthostomella sabalensioides (Ell. & G. Martin) Sacc., Syll. Fung. 2: xiii. 1883. 
= Entosordaria sabalensioides (Ell. & G. Martin) Hoehn., Sitzb. Akad. Wien 192: 166. 1920. 


Sphaeria xylariaespora Cke. & Ell., Grevillea 6: 94. 1877. 
On decorticated Andromeda, NEW JERSEY: Newfield, J. B. Ellis 2800 (lit.). 
= Rosellinia xylarispora (Cke. & Ell.) Sacc., Syll. Fung. 1: 272. 1882. 
= Coniochaeta xylarispora (Cke. & Ell.) Cke., Grevillea 16: 16. 1887. 


Thyridium americanum Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893. 139. 
1893. 


On Xanthoxylum americanum (prickly ash), CANADA: London, Apr 1892, # 1780 [Dearness 
(lit.)] (HOLOTYPE: NY) 

= Xylosphaeria americana (Ell. & Ev.) Petrak, Sydowia 4: 18. 1950. 

= Mycothyridium americanum (Ell. & Ev.) Petrak, Sydowia 15: 288. 1961 (1962). 


Thyridium antiquum (Ell. & Ev.) Berl. & Vogl., see Sphaeria antiqua Ell. & Ev. 


Thyridium canadense Ell. & Ev., N. Amer. Pyrenomyc. p. 416. 1892. 
On old logs, CANADA: Ontario, Lake Nigiron, 26 Jun 1887, [Macoun (lit.)] (HOLOTYPE: 


NY) 
= Thyridella canadensis (Ell. & Ev.) Sacc., Syll. Fung. 11: 351. 1895. 
= Peltosphaeria canadensis (Ell. & Ev.) Berl, Icon. Fung. 2: 110. 1897. 
= Peltosphaeria canadensis (Ell. & Ev.) Riedl, Sydowia 15: 304. 1961 (1962). 


Thyridium pallidum Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1895: 424. 1895. 
On dead Rhus glabra, KANSAS: Rooks Co., May 1895, Bartholomew 1711 (HOLOTYPE: 
NY) [25 May 1895 (ISOTYPE: FH)] 
= Xylosphaeria pallida (Ell. & Ev.) Petrak, Sydowia 4: 19. 1950. 
= Mycothyridium pallidum (Ell. & Ev.) Petrak, Sydowia 15: 289. 1961 (1962). 


Thyridium stilbostomum Ell. & Ev., J. Mycol. 9: 223. 1903. 
On old maple limbs, CANADA: Ontario, 21 Oct 1903 [Deamess 3001 (lit.)] (HOLOTYPE: 
NY; ISOTYPE: FH) 
= Xylosphaeria stilbostoma (Ell. & Ev.) Petrak, Sydowia 4: 19. 1950. 
= Mycothyridium stilbostomum (Ell. & Ev.) Petrak, Sydowia 15: 290. 1961 (1962). 


Thyridium syringae Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 139. 1893. 
On dead limbs of lilac (Syringa), CANADA: London, Mar 1892, Dearness 1654 (HOLOTYPE: 


NY) 
= Xylosphaeria syringae (Ell. & Ev.) Petrak, Sydowia 4: 19. 1950. 


oa 


= Mycothyridium syringae (Ell. & Ev.) Petrak, Sydowia 15: 290. 1961 (1962). 


Thyridium vitis Ell. & Ev., Bull. Torrey Bot. Club 27: 53. 1900. 
On dead shoots of Vitis riparia, KANSAS: Rocks Co., Bartholomew [lit.] 
= Xylosphaeria vitis (Ell. & Ev.) Petr., Sydowia 4: 20. 1950. 
= Mycothyridium vitis (Ell. & Ev.) Petr., Sydowia 15: 290. 1961 (1962). 


Valsa canodisca Ell. & Holw., Proc. Acad. Nat. Sci. Philadelphia 1890: 223. 1890. 
On Salix, IOWA: Decorah, 9 May 1886, E. W. Holway (HOLOTYPE: NY) 
= Eutypella canodisca (Ell. & Holw.) Sacc., Syll. Fung. 9: 463. 1891. 
= Engizostoma canodisca (Ell. & Holw.) Kuntze, Rev. Gen. Pl. 3(2): 473. 1898. 
= Diatrype albopruinosa (Schwein.) Cke. var. salicina Rehm, fide Rappaz, Mycologia Helvetica 2: 
503. 1987. 


Valsa capillata Ell. & Ev., J. Mycol. 4: 74. 1888. 
On decaying pieces laying in a damp place, LOUISIANA: St. Martinsville, 28 May 1888, A. B. 
Langlois 1254 (HOLOTYPE: NY; ISOTYPE: FH). 
= Eutypella capillata (Ell. & Ev.) Sacc., Syll. Fung. 9: 462. 1891 
= Engizostoma capillatum (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3(2): 472. 1898. 
= Peroneutypella capillata (Ell. & Ev.) Berl., Icon. Fung. 3: 84. 1902. 
= Eutypella scoparia (Schwein. : Fr.) Ell. & Ev., fide Rappaz, Mycologia Helvetica 2: 537. 1987. 


Valsa deusta Ell. & Ev., J. Mycol. 4: 74, 1888. 
On a piece of rotten Carya, LOUISIANA: St. Martinsville, 16 Jun 1888, Langlois 1334 
(HOLOTYPE: NY; ISOTYPE: FH) [On Carya olivaeformis (ISOTYPE: BPI)] 
= Eutypella deusta (Ell. & Ev.) Ell. & Ev., N. Amer. Pyrenomyc., p. 489. 1892. 
= Engizostoma deustum (Ell. & Ev.) Kuntze Rev. Gen. PI. 3(2): 473. 1898. 
= Eutypella scoparia (Schwein. : Fr.) Ell. & Ev., fide Rappaz, Mycologia Helvetica 2: 537. 1987. 


Valsa juglandina Cke. & Ell., Grevillea 5: 92. 1877. 
On Juglans regia, # 2421, NEW JERSEY: Newfield, Oct 1876, J. B. Ellis ISOTYPE: NY). 
= Eutypella juglandina (Cke. & Ell.) Sacc., Syll. Fung. 1: 154. 1882. 
= Eutypella juglandicola (Schw.) Ell. & Ev. var. juglandina (Cke. & Ell.) Ell. & Ev., N. Amer. 
Pyrenomyc. p. 495. 1892. 
= Engizostoma ellisii Kuntze, Rev. Gen. Pl. 3(2): 472. 1898. 


Valsa lutescens Ell., Bull. Torrey Bot. Club 9: 111. 1882. 
On dead limbs of Quercus coccinea, NEW JERSEY: Newfield, Feb 1881, N. A. F.: 876, as 
cited in the original description. (ISOTYPE: NY, FH) 
= Eutypella lutescens (EIlL.) Sacc., Syll. Fung. 2: vii. 1883. 
= Engizostoma lutescens (Ell.) Kuntze, Rev. Gen. Pl. 3(2): 474. 1898. 


Valsa maclurae Cke. & EIll., Grevillea 8: 14. 1879. 
On Maclura, NEW JERSEY: Newfield, Sep 1878, J. B. Ellis 3104 (ISOTYPE: NY) N. A. F.: 
873 (ISOTYPES: NY, BPI, FH) 
= Eutypella maclurae (Cke. & Ell.) Ell. & Ev., N. Amer. Pyrenomyc. p. 496. 1892. 
= Cytospora maclurae Ell. & Barth., Erythea 5: 48. 1897. 
= Enzigostoma maclurae (Cke. & Ell.) Kuntze, Rev. Gen. Pl. 3(2): 474. 1898. 


Valsa microcarpa Ell. & Ev., J. Mycol. 4: 122. 1888. 
On decayed pieces of peach tree, LOUISIANA: St. Martinsville, 18 Aug 1888, A. B. Langlois 
_ 1481 (HOLOTYPE: NY; ISOTYPES: BPI,FH) 
= Eutypella microcarpa (Ell. & Ev.) Sacc., Syll. Fung. 9: 462. 1891 
= Peroneutypella microcarpa (Ell. & Ev.) Berl, Icon. Fung. 3: 83. 1902. 
= Eutypella scoparia (Schwein. : Fr.) Ell. & Ev., fide Rappaz, Mycologia Helvetica 2: 538. 1987. 


Valsa rugiella Cke. & Ell., Grevillea 5: 92. 1877. 


598 


On maple [On Acer rubrum (lit.)], NEW JERSEY: Newfield, Mar 1876, J. B. Ellis s. n. 
(ISOTYPE: NY) N. A. F.: 176 (possible, ISOTYPE: NY, FH) 

= Eutypella rugiella (Cke. & Ell.) Sacc., Syll. Fung. 1: 156. 1882. 

= Engizostoma rugiellum (Cke. & Ell.) Kuntze, fide Rev. Gen. Pl. 3(2): 475. 1898. 


Valsa venusta Ell., Bull. Torrey Bot. Club 9: 112. 1882. 
On Robinia pseudacacia, NEW JERSEY: Gloucester Co., Newfield, Jun 1881 (HOLOTYPE: 
NY) N. A. F.: 875, as cited in the original description (ISOTYPE: NY, FH) 
= Eutypella venusta (Ell.) Sacc., Syll. Fung. 2: vii. 1883. 
= Engizostoma venusta (Ell.) Kuntze, Rev. Gen. Pl. 3(2): 475. 1898. 


Xylaria cylindrica Ell. & Ev., Bull. lowa Univ. Lab. Nat. Hist. 2: 414. 1892. 
On bark, NICARAGUA: Castillo Viejo and Ometepe, Winter 1893, B. Shimek & C. L. Smith 
12 (HOLOTYPE: NY; ISOTYPE: FH) 
= Xylaria teres Sacc., Syll. Fung. 11: 284. 1895. 
= Xylaria multiplex (Kunze) Dennis, Bull. Jard. Bot. Etat 31: 125. 1961. 


ACKNOWLEDGEMENTS 


I am grateful to the following individuals who have freely given me unpublished information, 
advice, encouragement, and time: Drs. J. R. Boise, R. E. Halling, D. H. Pfister, C. T. Rogerson, A. Y. 
Rossman, G. J. Samuels, and B. M. Thiers. Part of this project was completed with a grant from the 
Friends of the Farlow Reference Library and Herbarium of Cryptogamic Botany. I deeply appreciate 
the critical comments offered by Profs. M. E. Barr Bigelow and J. D. Rogers in their presubmission 
reviews of this manuscript. 


LITERATURE CITED 


Batista, A. C. 1956. Systematic revision of the genera Ellisiella Sacc. and Ellisielina Camara, and the 
new genus Ellisiopsis. Ann. Soc. Biol. Pernambuco 14 1/2: 16-25. 

, & G. E. P. Peres. 1965. Novos Deuteromycetes da Micogeografia intercontinental. Mycopath. 
Mycol. Appl. 25: 161-172. 

Cash, E. K. 1953. A record of the fungi named by J. B. Ellis. USDA Special Pub. 2(2): 1-345. 

Da Camara, E. S. 1949. Mycetes Aliquot Lusitaniae IX. Agron. Lusit. 11: 39-73. 

Deighton, F. C. 1969. Microfungi. IV: Some hyperparasitic Hyphomycetes, and a note on 
Cercosporella uredinophila Sacc. Mycological Paper 118: 1-41. Commonwealth Mycological 
Institute, Kew. 

Elisens, W. J. 1985. The Montana collections of Francis Duncan Kelsey. Brittonia 37: 382-391. 

Eriksson, O. E. & D. L. Hawksworth. 1987. Outline. of the Ascomycetes. Systema Ascomycetum 6: 
259-337. 

Fries, E. M. 1821 - 1832. Systema Mycologicum. Lund, Berlin. Griefswald. Moritz. 3 vols. 

Harris, R. C. 1973. The corticolous pyrenolichens of the Great Lakes region. Mich. Botanist 12: 3-68. 

Kaye, G. C. 1986. A Symposium on the history of North American Mycology: Job Bicknell Ellis. 
Mycotaxon 26: 29-45. 

Martin, P.M. D. 1976. Studies in the Xylariaceae: supplementary note. J. S. African Bot. 42: 83. 

Miller, E. & J. A. von Arx. 1973. Pyrenomycetes: Meliolales, Coronophorales, Sphaeriales. In: The 
Fungi: An Advanced Treatise, G. C. Ainsworth, F. K. Sparrow and A. S. Sussman (eds.), vol. IV 
A, pp. 87-132. Academic Press, NY. 

Rogers, D. P. 1981. A Brief History of Mycology in North America, pp. 13-14. Mycol. Soc. Amer., 
Cambridge, Mass. 

Saccardo, P. A. 1880. Conspectus generum fungorum Italiae inferiorum, nempe ad sphaeropsideas, 
melanconieas et hyphomyceteas pertinentium systemate sporologico dispositorum. Michelia 2: 
1-38. 

Shear, C. L. 1946. Studies of types and authentic specimens of Hypoxylon - I. Lloydia 8: 245-262. 


ae, 


Stafleu, F. A. & R. S. Cowan. 1976. Taxonomic Literature, 2nd ed., Bohn Scheltema, Utrecht. 1: 742- 
743. 

Stevenson, J. A. 1971. An account of fungus exsiccati containing material from the Americas. Beih. 
Nova Hedwigia 36: 99-164. 

Theissen, F. von. 1914. Uber Polystomella, Microcyclus u.a. Ann. Mycol. 12: 63-75. 


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MY COTAXON 


Vol. XXXIV, No. 2, pp. 601-614 January-March 1989 


NOTES ON TROPICAL AND WARM TEMPERATE BASIDIOMYCETES 
A.L.WELDEN and C.L. OVREBO 


Department of Biology 
Tulane University 
New Orleans, LA 70118 


SUMMARY. Eight noteworthy fungi in seven genera from the tropical and 
warm temperate zones are discussed and illustrated Four, De//exu/é 
subsimplex, tlachnocladium schweinfurthiana, end Remearieé 
cyanocephala and R. zippe/i7 ere clavarioid; one, fcehinochaete 
brachyporus is poroid; one Gyrodontium versico/or is tydnoid; one, 
Gomphus cavipesis cantharelloid; andone, 7he/ephore aurantiotincta 
is thelephoroid. All are either rarely reported, or occur outside their known 
ranges, or they are poorly understood, or a combination of these. 


Key Words: Deflexula, Lachnocledium, Ramaeria, Echinochaete, Gyrodontium, 
Gomphus, Thelephora 


Deflexula subsimplex (P.Henn.) Corner, Ann. Bot. n.s. 16: 279. 1952. 
Fig. A 1. Plate 1. 


Basidiocarps springing from small inconspicuous mycelial discs 2-3mm 
wide,consisting of small and large clusters of unbranched pendent spines 
- lem; width of spines ca. 300 um, whitish buff when fresh, dark flesh color 
when dry. 

On corticated wood of an undet. hardwood (Guzman 24483 in ENCB & NO); 
on tree bark (C. Ovrebo 2083 & A. Meier in NO). The former collection is 
from Mexico; the latter from Costa Rica. 

Spores 10-11 x 5.4-6.3 im, amygdaliform, with slightly thickened 
walls, 8 prominent apiculus end often a large guttule, angled and light yellow 
in drying, IKI-; basidis not observed; besidioles ca. 23-30 x 6-9 ym, 
prominently guttulate, stoutly clavate or cylindrical with swollen spices; 
hymenium not developed for ca. 300 pm from the tip of spine on one side and 

ca. 900 pm from the tip on the other; dimitic with skeletal hyphae; 
subhymenium of generative hyphae ca. 4 pm in diam arranged into 4 
pseudoparenchyma; cystidia and other sterile organelles absent; generative 
hyphae of context 4-6 um in diam, with very long cells, with clamps and 


602 


often secondary simple septa; skeletal hyphae unstained in phloxine, with 
slightly thickened walls, prominent in context. 


This fungus was reported by Corner (1967) as 2. n/vea(Pat.) Corner 
from Guadeloupe. Later, Corner (1970) synonymized that taxon with 2. 
subsimplex end reported collections from Costa Rica, Panama, Brazil, 
Bolivia (our spore sizes reported here are close to those reported by Corner 
for Singer's collection from this region), Argentina, Indis, Borneo, and 
Solomon Islands. Although this taxon is seldom listed in tropical collections, 
it appears to be a typical member of the pantropic mycoflora. 


Echinocheete brachyporus (Mont.) Ryv., Bull. Jard. Bot. Nat. Belg. 
48:101. 1978. Fig A 2-7. 


Basidiocarp laterally stipitate, somewhat spongy coriaceous; pileus 4.6 cm 
long x 6.5 cm wide, undulate to shallow lobed, stipe junction clearly marked 
by narrow ridge on upper surface; abhymenial surface minutely tomentose, 
orange yellow with vinaceous tints, especially toward margin, extreme edge 
may be deep wine red; adhymenial surface near pale orange yellow but reddish 
black at pore edges (bruising ?); pores shallow, 2-3/mm in well-developed 
areas, some appear pulled apart in drying, other small areas devoid of pores; 
pores irregularly polygonal and with brown protruding setoids (under lens), 
decurrent over ca. one-half stipe surface. 

Stipe 0.8 cm long from ridge at pileus junction to base x 0.8 cm wide, with 
traces of pore outlines caused by minutely fimbriate ridges which darken 
with KOH (setoids). 

Lignicolous. Mexico: Cobs, Quintana Roo, 12 May 1986, leg. A.L.W. s.n. 
(NO). 

Spores 10-14 x 3.5-6.5 pm (x Il= 12.7 x 4.6 um; e=2.7), 
cylindrical to cylindrical-allantoid, colorless, often with 1-several large 
guttules and many smaller ones, IKI- , acyanophilous. 

In section pileus with cuticle and context; adhymenially the context 
extends into trama with only slight change in hyphal organization; cuticle 
27-90 pm thick, of densely compact basically parallel hyphae, with an 
occasional hyphal end projecting sbhymenially; context of interwoven hyphae 
often more compact in trama; generative hyphee in + perallel bundles; 
subhymenium inconspicuous. Hyphal structure dimitic with binding hyphee; 


FIG. A. 1. Deflexu/a subsimp/ex. Hymenial detail, clamped generative 
hyphae, and spores. 2-7. £ch/nocheete brachyporus. 2. Habit sketch x 
0.8. 3. Mature setoid elements from hymenium. 4. Young setoid element. 5. 
connection between wide thick-walled generative hypha and narrow 
thin-walled generative hypha. 6. Hymenial detail and spores. 7. Binding 
hyphae. Bar= 10um. 


603 


604 


generative hyphae thin-walled clamped, ca. 4.5 um in diam, some -12 um in 
diam, staining evenly in phloxine and usually with long cells (vascular 
hyphae 7); binding hyphee unstained in phloxine, walls -1 pm thick, main 
body 6-7 ym in diam, sparingly branched slong its length, with 1-few 
tapering branches at tip, these with narrow lumen, 2-3 um in diam, -400 
uum between a branch and its tip. Hymenium of basidioles, basidia, and 
spinulose heavily pigmented setoids; setoids metamorphosed basidioles, 
basidia ca. 7 um in diam, clavate, 4-ster igmate. 


This taxon is pantropical. It has been reported from Cayenne in South 
America (Saccardo, 1888) and from Uganda, Tanzania, Malawi, Kenya, and 
Burundi. It would be interesting to know the forest type of those collections. 
The Quintana Roo collection was made in a Semi-deciduous Forest (Rzedowski, 
1986) on limestone soil. The forest is subject to an annual dry season of 
some sever ity. 


Gomphus cavipes Corner, Nov. Hedw. 18: 808. 1969. Fig. B 1-4. Plate 1. 


Basidiocarps fleshy-coriaceous, petalloid, flabellate, ochraceous 
cinnamon, some with recurved margins giving an infundibuliform 
(pseudoinfundibuliform) appearance, 5-8 cm tall KOH-, greenish gray in 
ferric sulfate solution; abhymenial surface smooth, glabrous, opaque or 
watery translucent, light ochraceous, not discoloring; adhymenial surface 
smooth, dull, pale, not discoloring; context 3-4 mm thick midway from 
pileus margin to stipe; stipe hollow, rounded to somewhat flattened, 
concolorous except for white basal mycelium, sometimes 2-3 stipes joined at 
base. 

On soil. Colombia: Mpio. Guatape, Dpto. Antioquia. In premontane 
rainforest, ca. 1850 m alt. leg. C. Ovrebo 2520 (NY, HUA). 

Spores 8.5- 16.2 x 4.5-7.2 pm (x 30= 9.8 x 5.9 um; e=1.6), oval to 
cylindrical, light ochraceous, ochraceous buff in deposit, rugose, 
thick-walled, amygdaliform , cyanophilous, apiculus 1.5-2 jim long. 

Basidia ca. 56 x 7-9 pm (base obs. in one), clamped, cylindrical-clavate 
or clavate with long slender stalks 3-4 um in diam, enlarging toward apex, 
4-sterigmate, each sterigma ca. 4.5 pm long, straight; paraphysoid hyphse 
5-4 Mm in diam, among basidia and basidioles; clamped generative hyphae 
5-9.4 jim in diam, thin-walled, some giving rise to numerous, unbranched, 
1-2x coiled oleiferous hyphae 3-9.5 um in diam, found only in context. 


This is second report of G cavipesand extends its range slightly from 
Trinidad to Colombia. It differs in minor ways from the holotype. The 
adhymenial surface is smooth but in some areas it drys into irregular blocks 
which exposes the context. The holotype shows a number of shallow folds 


605 


and little or no cracking. The collection was ochraceous cinnamon when 
fresh; the holotype white. There was no color change when handled, but (and 
this was inadvertently omitted when the holotype was sent to Professor 
Corner) notes accompanying a second Trinidad specimen (ALW 1756) refer 
to a color change to vinaceous when bruised. More seriously, the Colombian 
material lacks the smaller lateral pilei of the holotype when dry and appears 
pleuropodial rather than mesopodial. In some respects this material 
resembles the description of G swbclavaeformis (Berk.)Corner 
(1970). However, the lack of gloeocystidial endings in the hymenium and 
its hollow stipe firmly places this fungus in G cav/pes. The apparent 
pleuropodial condition in its dried state may be explained in part by the 
tendency of the hollow base to collapse when drying and in part to the evident 
flattening and coalescing of the distal parts into a dorsiventral pileus. The 
surfaces surrounding the hollow stem tend to flatten as they grow distally. 
As these surfaces come together , the narrowing hollow is filled with loosely 
interweaving vegetative and oleiferous hyphee surrounded by densely 
compact longitudinally parallel hyphae, also with oleiferous elements.. Still 
more distally the hollow disappears and the two surfaces coalesce. In section 
this latter region appears as a typical dorsiventral pileus. Hymenial 
development is unilateral even before coalescence into a flattened pileus. But 
before these features can be determined accurately, freshly collected 
material preserved in a weak fixative such as recommended by Corner 
(1967) must be available. 


Gyrodontium versicolor (Berk. & Br.) Maas G., Persoonia 3(2): 190. 
1964. Fig. C 1. Plate 1. 


Basidiocarp sessile-pileate; pileus 35-45 mm longx S0-75 mm wide, 
+ reniform; abhymenial surface matted-cottony, slightly moist, light 
yellow when young, becoming darker ochraceous brown, finally brown, often 
darker brown toward base of pileus, not discoloring; context spongy, 3 mm 
thick, light greenish yellow; adhymenial surface with terete, obtuse, 
yellowish green spines 2-3 mm long toward the base, shorter and smaller 
toward the margin, ca. 2/mm; margin with whitish sterile zone 1-2 mm 
wide. 

Costa Rica: La Selva Biological Station, on base of dead tree in lowland 
tropical rainforest, 2 Jul 1986, leg. C. Ovrebo 2084; |.c., on same 
substrate, leg. 7 Jul 1986, C. Ovrebo 2126. 

Spores 4-5S.4 x 2.7-3.1 pum (x 10= 4.0 x 2.8 jum; e= 1.4), oval, 
_ brown, thick-walled, with prominent apiculus, print dark olivaceous green 
whenfresh, olive brown when dry. 

With clampless generative hyphae 2-6 yim in diam, thin-or slightly 
thick-walled, pale yellowish; context of compact anastomoasing fibrils of 


606 


hyphae separated by loosely arranged, branching individual hyphae, more 
compact adhymenially where densely interwoven hyphae grow into spines 
bearing the hymenium ; abhymenial surface covered by amorphous material 
with numerous embedded spores often mixed with loosely arranged hyphae; 
basidia and basidioles completely covering the spine surface; basidia 20-24 
x 3.6-5.4 um with 4 short needle-like sterigmata each ca. 2 um long. 


Maas Geesteranus (1964) has painstakingly untangled the synonymy of 
this taxon, which he considers monotypic. Specimens of @ vers/co/orare 
known from Africa, Australia, the Bonin Islands, and Ceylon. It is listed in 
Hawksworth, et al (1983) as widespread. To our knowledge this is the first 
report of this species from the New World. 

Gyrodontium spores are typically coniophoroid. In any single mount 
they are erratically cyasnophilous. The positively cyanophilous spores are 
the younger ones. The fruitbody is often spotted deep brown. This is caused 
by heavy deposits of spores. 


Lachnocladium schweinfurthiana P. Henn., Bot. Jahrb. 17:21. 1893. 
Fig. B. S-7. 


Basidiocarps densely caespitose, -4.5 cm tall, arising from ochraceous 
rhizomorphic strands and white mycelium incorporating debris; much 
branched dichotomously and polychotomously almost to the base; 
anastomosing freely by outgrowths of cortical hyphae into a compact mass, 
apices often suppressed and cristate. 

U.S.A.: Florida, in humus closely associated with rotting hardwood stump 
and grass rhizomes, Univ. of Florida campus near Florida State Museum, 
Gainesville, 11 Aug 1985, leg ALW s.n. (NO). 

Spores 3.5-5.0 x 25-45 pm (x10=3.7 x 3.1. um; e=1.1), 
subspheerical, hyaline, thin-walled, IKI-, acyanophilous, 0- |-quttulate; 
basidia 20-37 x 4-5 ym, cylindrical (ventricose-cylindrical ?), 
4-sterigmate, each sterigma ca. 6 um long, straight, needle-like. 

Basidiocarp in section showing a central rounded to flattened darker 
medulla - 180 jim in diam surrounded by a paler cortex; medullary hyphae 
2-5 um in diam, thin-walled, hyaline, or lightly pigmented with thickened 
walls, dichohyphidia occasionally present; cortex filled with dichophyses, 


Fig. B. 1-4. Gomphus cavipes. \. Habit x 0.8. 2. Basidia and basidioles. 3. 
spores. 4. Oleiferous hyphs. 5-7. Zechnoc/sdium schweinfurthiene. S. 
Habit x .8 and schematic drawing showing branching of medulla. 6. Basidia, 
spore, and gloeocystidia. 7. Dichohyphidium from cortex. Bar= 10 pm. 


607 


608 


faintly stratose, dichohyphidia at base 3-4.5um in diam, branching 
dichotomously several times, ultimate branches ending in short acutely 
tipped dichotomies, walls pigmented, thickened toward the base, those toward 
the periphery smaller but not coralloid, faintly dextrinoid. 


In the field the densely anastomosing fungal mass appears reddish brown 
with yellowish tips. On closer inspection this appearance is caused by the 
absence of the pinkish gray hymenium from the ochraceous tips, which 
contrasts with the darker parts. 

Species of Lachnoc/adium are characteristic elements of wet tropical 
forests. This particular taxon has been reported under 6 variety of names 
(Corner, 1967, 1970) from Blumenau end Amazonia, Brazil ,£ & S. Africa, 
and from the Congo to Malaya. The present collection demonstrates that this 
taxon, under appropriate conditions, may grow in warm temperate zones as 
well. 

Corner (1970) restricts Lachnocladtaceae to this genus, rejecting 
Reid's suggestion that O/chop/euvropus Reid, Veraria Karst., 
Scytinostrome Donk, and Asterostrome Mass. be included within the 
family. Boidin (1980) and his colleagues and students have intensively 
Studied Vararije and have divided the genus into Vererve Kerst. and 
Dichostereum Pil. A seventh genus, Dichocenthere//us Corner 
(1966) bears some resemblance , i.e., dichophyses and gloeocystidia, to this 
group. It is tempting to bring these genera together into 8 single family. 
However, because of the number of features which appear sporadically 
throughout the various species, e.g., clamp-connections, sulfocystidia 
(macrocystidia) amyloid spores, oleiferous hyphae, ornamented spores, it is 
more discrete at our present level of knowledge to be conservative in our 
taxonomy. 


Ramar ia cyanocephala (B. & C.) Corner, Ann. Bot. Mem. 1: 568. 1950. 
Fig. C 2-4. 


Basidiocarps of many stout cinnamon brown branches very compactly 
arranged, 7. cm tall x 5 cm wide x 3 cm deep; branching at or near the base, 
dichotomous with first and second internodes rather long, third internodes 
relatively quite short and immediately forming dichotomies, frequently one 
of the ultimate dichotomies suppressed giving the sppeerance of 6 
trichotomy; branch tips in dried material pale tan (color of fresh material 
not noted); medullary flesh whitish, pale tan or cream, 4/u/sh green iin 
ferric sulfate solution. 

Coste Rica: Guanacaste Province near Bagaces, 29 Nov 1983, L.D.Gomez & 
R. Alfaro 22106 in NO. 

Spores 10.8 - 15.3 x 6.3-9 ym, thick-walled, yellow brown, with long 
acute spines, amygdaliform, prominently epiculete. Hyphae thin-walled, 


609 


clamped, 3-6.5 yim in diam. Basidia 8-12 ym in diam toward apex, 3 um 
wide toward base, more than 54 ‘ym long (base not obs.), bisterigmate. 


The general impression of the fructification is that its growth was 
arrested, although spores ere abundant. This is not an unreasonable 
assumption, for the specimen was collected in s Tropical Dry Forest Moist 
Providence Transition (see Tosi map in Janzen, 1983) at the end of 
November when the region is entering one of its driest periods. 


A second collection of this taxon has a radically different appearance. 


Basidiocarps of many slender cinnamon brown branches laxly erranged, 
- 15cm tall x Scm wide, stem -5 cm long x 5S mm wide, each major segment 
with 3-6 dichotomies with relatively long internodes , ultimate branches 
tapering to acute tips; nodal areas flattened (in drying?); tips darkened or 
concolorous; medullary flesh whitish to pale tan , /v/sh green in ferric 
Sulfate solution. 

Costa Rica: Puntarenas Province, Finca El Eden, Km183, Rt. 2, 6 Sep 
1983 , Gomez 22966 (NO) 

Spores 9.9-13 x 63-7.2 wm Hyphae 2-10 pm in 
diam ,thin-walled,clamped. Basidis 6.3-9 wm in diam, with two stout 
Sterigmata -9 um long. Microscopically similar to Gomez & Alfaro 22106. 


Macroscopically the two specimens are similar only in their color. 
Microscopically they are practically identical. The taller, more lax 
collection (Gomez 22966) was made in southern Costa Rica in a region of 
Tropical Moist Forest in a wet month. Climatic difference may help explain 
the difference in macroscopic appearances of these specimens (assuming the 
determinations ere correct). Clavarioid fungi have an open growth plan, and 
more abundant moisture would allow maximum elongation of the branches, 
which would help develop s less compact, more lax and open fructification. 
The reverse would be true in a regimen of declining moisture. 


Ramar ia zippelii (Lév.) Corner, Ann. Bot. Mem. 1: 632. 1950. 


Basidiocarps- S cm tall, bushy, branches tan to brown mixed with purple 
brown toward the base, some purple brown upon injury, tips greenish yellow 
to pinkish; medullary flesh tan to brownish, 7o¢ reacting to ferric 
Sulfate solution. 

Trinidad: near Arima in the Northern Range. ALW & Lemke 1664, 1745, 
1759, 1771, all kindly determined by Professor Corner as A. grands 
(Pk.) Corner and later synonymized by him with A. zeape///. 

Spores typical of the subgen. £c//norameriaser. Grandisporée, 


610 


10-16x 6.5-9.5 um. Basidia bister igmate. 


Corner (1967,1970) has emphasized the difficulty of differentiating A. 
cyanocephals and A. zeppe/i/ without field notes on fresh material, 
especially the color of the branch tips. In his key toser. Grand7sporee 
Corner lists 4 taxa which have predominantely bisterigmate basidia. 
Remaria nigrescens (Brinkm.) Donk and 2. ap/ehyana (Speg.) Corner 
were eliminated because of their orange color and the blackening of the flesh 
in the former species. Specimens of A. cyanocephe/a with purple tips 
collected from Jamaica (ALW 487) and Trinidad (ALW 1719), the former 
confirmed by Professor Corner , have 6 whitish medulla and become bluish 
green in ferric sulfate solution. Our specimens of A. 2/fpe@/77 \ack this 
reaction and the medulla is darker. 

That at least some ramarias react to ferric sulfate solution (Donk, 1964; 
Marr & Stuntz, 1973) is well-known. According to the latter authors only 6 
few taxa among those from Washington state (A. amy/oices, &. 
celevirescens, R. claviemulate, and PR. velocimutens, all new 
species) react instantly and dramatically. In addition to A. cyanocephals 
and A. 2/ppe//7 there are seven other taxa in NO. Reacting immediately to 
ferric sulfate solution are. A //evobrunnescens (Atk.) Corner, A. 
mollerena(Bres. & Roum.) Corner, A. str/cta(Fr.) Quel., and A. verne 
(Coker) Corner, whose reaction occurs in the region at the base of the cortex 
and not in the medulls proper. Not reacting, or reacting faintly at best, are: 
R. eurea(Fr.) Quel., A cir. formosa (Fr.) Quel., and A. guayensis 
(Pat.) Corner. Our material is dried and some of our specimens are several 
years old. Specimens of ”. cyvanocepha/a and R. 27ippe/7s trom Trinidad 
were collected 28 years ago, and A. cyaenocepha/a still reacts instently to 
ferric sulfate. Even though little is known of the relationship of this reaction 
to basidiocarp age, and that such tests are best performed on fresh material, 
it is noteworthy that among the specimens available to us the reaction 
distinguishes the two taxa. We recognize the Costa Rican specimens as A. 
cyanocepha/abecause: (a) their microscopic features are essentially those 
of A. cyanocepha/a, (b) they react instantly and definitively to ferric 
sulfate, (c) the only other similar taxon, A. 2/p9e//7, does not react, and 
(d) known specimens of &. cyanocepha/a of the same age as 2. 2/ppe/// 
specimens also react instantly to ferric sulfate. 


Fig. C. 1. Gyrodontium versico/or. Basidia, basidioles, and spores. 2-4. 
kamera cyahocephala 2. Generative hyphae, some inflated. 3. Spores. 4. 
Basidia, middle one with a minutely branched sterigma, and generative 
hyphae. 5-7. /helephorea surantiotincta. 5. Habit x 0.8. 6. Generative 
hypha with occluded lumen. 7. Basidia, basidiole, and spores. Bar = 10um. 


611 


612 


Thelephora aurantiotincta Corner , Beih. Nov. Hedw. 27:44. 1968. Fig. C 
5-7. 


Basidiocarp -7 cm tall x 6.6 cm wide (a portion of the base broken off), 
with a very strong smell of fenugreek , foliaceous, imbricating from a common 
stem, intricately fused, segments thick, not cyanescent in KOH but darkening 
(pale brown) in KOH and ferric sulfate solution; abhymenial surface matte, 
off-white to pale cream, spiculose-striate, striae caused by groups of repent 
hyphae, these sometimes purplish brown, especially toward the margin, 
azonate; margin thick, blunt, as if arrested in development, stained purplish 
brown, streaked this color toward base; adhymenial surface pallid white 
except toward margin and where papillae develop; papillae distinct, ca. 0.5 
mm high, but tips soon purplish brown giving a light brown tone to the older 
parts, younger portions fading toward margin from orange through pale 
orange to white. 

Mexico: ca. Atocuapan, Km 2.5 Totula-Xalapa highway , alt. 1350 m, 
Veracruz, terricolous ?, 23 Jul 1984, leg. A. Sempieri 915 (XAL , NO). 

Spores 5.8 -7.2 ym, lobate, angles minutely crispate, apiculus 1- 1.5 um 
long, fuscous vinaceous, IKI-. 

Monomitic with clamped hyphee 3-7 um in diem, thin-walled or with 
sclerified segments, these with capillary to occluded lumens, wider 
thin-walled hyphae often with simple secondary septa, sclerotized hyphae 
occuring singly or in small groups traversing the context. Basidia $4-56 x 
7.2 um, 4-sterigmate, cylindrical clavate; cystidia absent. 


Corner (1968) described this taxon as growing in humus of a mountain 
forest (Mt. Kinabalu) in N. Borneo at ca. 1600 m alt. As he notes, 7. 
aurantiotincta is aremarkable member of its genus. It would immediately 
catch the eye of any collector passingly familiar with 7/e/ephore A second 
intriguing feature is the similarity of habitat in widely separated areas. This 
report represents only the second time this taxon has been reported. It will 
be interesting to learn in the future if similar habitats in other areas of the 
tropics also support this fungus. 

A sterile floccose hymenochaetoid fungus was growing over the abhymenial 
surface of this collection. The complete lack of spores and basidia precludes 
determination. 

ACKNOWLEDGEMENTS 


The authors sre grateful to s number of agencies end individuals for making 
it possible to assemble not only specimens of the fungi reported here, but for 
other specimens which have been invaluable in our comperative 
Studies. The agencies sre the National Science Foundation (ALW to collect in 


613 


the Lesser Antilles, Central America, and Mexico) and the Tinker Foundation 
through the Mesoamerican Ecology Institute of the R.T. Stone Center for Latin 
American Studies of Tulane University (ALW & CLO). C.L.Ovrebo’'s collecting 
in Colombia was supported by R. E. Halling (NSF grant BSR - 860024). 
Individuals who have sent specimens are Dr. Gaston Guzman of Mexico and L. 
D. Gomez of Costa Rica. Prof. £.J.H. Corner kindly identified a large number of 
Clavar ioid fungi. 
LITERATURE CITED 

Boidin J.&P.Lanquetin. 1980. Contribution a l’etude du genre Dichostereum 
Pilat (Basidiomycetes Lachnocladiaceae).Bull. Soc. Mycol. France 96( 4): 
581-406. 


Corner, E.J.H. 1966. A monograph of cantharelloid fungi. Oxford Univ. Press. 
Vii + 255 pp. 


—_________ 1967. A monograph of C/avar/eand allied genera. Facsimile 
reprint. Ann. Bot. Mem. 1. xv+ 740pp. 


—________.._ 1968. A monograph of 7/4e/ephora (Basidiomycetes). Beih. 
Nov. Hedw. 27: 1-110. 


1969. Notes on cantharelloid fungi. Nov. Hedw. 18: 783-818. 


—_______._ 1970. Supplement to 8 monograph of C/avar/e and allied 
genera. Beih. Nov. Hedw. 33:1-299. 


Donk M.A. 1964. A conspectus of the families of the Aphyllophorsles. 
Persoonia 3(2): 199-324. 


Hawksworth, D.L., B.C.Sutton, & G.C. Ainsworth. 1983. Ainsworth and Bisby’s 
Dictionary of the fungi. Commonwealth Mycological Inst. England. 


Janzen, Daniel H. ed. 1983. Costa Rican Natural History. Univ. of Chicago, IL. 


Meas Geesteranus,R.A. 1964. Notes on  Hydnums-II.Persoonia 
3(2): 155-192. 


Marr, Currie & Daniel E. Stuntz. 1973. Aamersvaof western Washington. 
Bib). Mycol. 38: 1-232. 


Rzedowski, J. 1986. Vegetacion de Mexico. Editorial Limusa. Mexico. 432p. 


saccardo, P.A. 1888. Sylloge Fungorum. 6: 228. 


614 


Plate 1. Gomphus cavipes. Upper leftca. x 1. Deffexu/a substmp lex. 


Upper right ca. x 1. Gyrodontium versicolor. Below ca. x 1. Photos by 
C. L. Ovrebo. 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 615-622 January-March 1989 


AMANITA PROTECTA—A NEW SPECIES 
FROM COASTAL SOUTHERN CALIFORNIA 


Rodham E. Tulloss 
21 Lake Drive 
Roosevelt, New Jersey 08555 


Greg Wright 
4517 Live Oak Drive 
Claremont, California 91711 


Summary 


Amanita protecta is described as new from the coastal region of 
southern California. 


Amanita protecta Tulloss & Wright sp. nov. Holotypus: California, Los Angeles 
County, Santa Monica Mountains, Stunt Canyon, 3.i.1987 Barry Silver [Wright 1383A 
= Tulloss 1-3-87-BS1] (NY’). 


Etymology: protectus, protected—treferring to the thick universal veil in this 
species. 


Pileus griseolus vel griseus vel brunneo-griseus, 40 - 115 mm in mensura 
diametrica, margine striata, nonappendiculata; materies volvica fibrillo-coacta, alba 
vel subcinerea, et, juxta marginem, pallida, rara, pulverulenta. Lamellae confertae, 
liberae vel anguste adnatae, margine grisea vel griseo-brunnea. Stipes 80 - 130 x 10 - 
22.5 mm, pallidus, fibrillis griseis vel griseo-brunneis. Volva circumsciscens vel 
lobata, alba vel albida, a stipe remotissima, 20 - 40+ mm alta. Caro albida vel 
subgriseola, decolorationibus denique ochraceis. Basidiae tetrasterigmaticae, (53.5-) 
60.5 - 92 x 13.5 - 16.5 (-17.5) uum; fibulae absentes. Sporae (8.7-) 9.4 - 12.6 (-14.7) x 
(7.3-) 8.4 - 11.2 (-13.3) um, globulosae vel subglobulosae vel late ellipsoideae vel 
ellipsoideae, nonamyloideae. 


if 
F - Herbarium of the Field Museum of Natural History, Chicago, Illinois, U.S.A. 
NY - Herbarium of The New York Botanical Garden, Bronx, U.S.A. 


All collections with no herbarium location cited are in Tulloss’ private herbarium. 


616 


Amanita protecta (Fig. 1) is a member of section Vaginatae. It is a medium-sized 
mushroom with gray to gray-brown pileus, an exannulate stipe covered for much of its 
length with dark gray to black fibrils, and a robust universal veil leaving thick 
felted/fibrillose patches over a layer of fine pulverulence on the pileus and a thick, 
lobed, volva at the stipe base. The volval limb is rather widely separated from the stipe 
by a gap partially due to the fact that the inner 1-2 mm of the volva is carried up on the 
expanding pileus. All parts of the basidiocarp have a tendency to develop ochraceous 
stains when damaged and due to aging. The context of the pileus and stipe, as well as 
the universal veil tissue, display many terminal hyphal segments and terminal inflated 
cells (sometimes in chains) with thick walls. In the portion of the universal veil below 
its desiccated surface, the hyphae are coiled and twisted. The basidia of this species can 
be quite long for the genus. Clamps were observed only in the stipe context where they 
were very rare. 


PILEUS: 40 - 115 mm diam, from pale to dark gray or “‘brownish grey’’ or deep 
brownish gray (11D-E27), viscid when moist, becoming shiny when dry, convex 
expanding to plano-convex with deflexed margin to finally concave, occasionally 
deeply rimose; margin weakly striate to striate (0.1R - 0.25R); universal veil up to 3 
mm thick, occasionally appearing to have two layers (then the ‘‘outer layer’’ thin, 
whitish with ochraceous staining, rather fragile, submembranous), otherwise whitish or 
pale grayish at first, eventually cream or grayish with ochraceous stains, subatomate to 
atomate, in large floccose-felted patches becoming smaller toward margin where the 
remnants are reduced to a fine pulverulence, detersile; context 4+ - 10 mm thick at disk 
thinning evenly to margin or to about 90% of radius and then a membrane to margin, 
firm, whitish to pale grayish, becoming faintly pink or ochraceous on exposure, 
occasionally dark grayish below the pileipellis in disk. LAMELLAE: close, 4.5 - 12.5 
mm broad, free to very narrowly adnate with a white decurrent line on stipe, white with 
a dark gray or gray-brown margin, unchanging, after drying 4A-B3 or 5A4 or 2.5Y 
8/2-4 or 7.5YR 8/6; margin minutely fimbriate (lens); lamellulae truncate, frequently 
with line of truncation concave, occasionally anastomosing with neighboring lamellae. 
STIPE: 80 - 130 x 10 - 22.5 mm, white to pale grayish at apex, creamy white to 
grayish with ochraceous stains for 10 - 20 mm above attachment of volval sac 
(sometimes with pallid silky fibrils in this area), otherwise medium gray to very dark 
gray to gray-brown fibrils (sometimes in chevron pattern) on a sordid ground, 
subcylindrical, flaring at apex, without bulb, rounded to somewhat pointed at base, 
staining ochraceous with exposure; context white above and white, fulvous, tannish, or 
salmon-cinnamon in base, faintly becoming pink or staining ochraceous, the 
discoloration becoming more intense toward the base, stuffed, central cavity diam 
about one third to one half that of stipe, or rarely solid, at times becoming hollow near 
base; exannulate; universal veil saccate, subcircumscissile to limbate, highest point of 
limb reaching to 20 - 40+ mm from stipe base, up to 5 mm thick at point of connection 
to stipe, surface white to whitish with ochraceous stains, occasionally the surface 


2 
Color codes of the form ‘‘11D2’’ and associated color names in double quotation marks are from 
(Korerup & Wanscher, 1978). Color codes of the form ‘*7.5YR 8/6”’ are from (Munsell Color, 1975). 


617 


Fig. 1 Amanita protecta. Tulloss 1-3-87-BS1 [x0.9]. 


618 


dessicates producing a thin, submembranous to hardened or shell-like ‘“‘outer layer’ 
sometimes having the texture of a chicken eggshell; interior pale grayish white to 
grayish, floccose-felted, at times largely carried up onto pileus so that the ‘‘outer 
layer’’ forms a limb separated considerably from the stipe, occasionally leaving only 
one or more concentric ridges between the volval sac and the stipe (Fig. 2), 
occasionally remaining as a separate second limb (not a limbus internus) which may 
even project above the limb formed by the ‘‘outer layer’’ (Fig. 3); no noticeable limbus 
internus. Odor slightly sweet (Wright) to faintly unpleasant in age (Tulloss). Taste 
slightly sweet (Wright). Wright experienced no symptoms of poisoning after eating 
about 30 g of the cooked mushroom. Meixner test (Vergeer, 1986) negative for 
amatoxins—unreactive or dull orangish or orange-reddish with a yellow ring around 
the acid drop. Paracresol spot test for tyrosinase (Marr, 1979) positive except in 
universal veil, lamellae, marginal pileus context, and very base of stipe. Syringaldazine 
spot test for laccase (Marr, 1979) negative. Phenol on stipe: wine-red. KOH on pileus: 
no reaction. 


PILEIPELLIS: up to 560 um thick, composed of interwoven, undifferentiated, 
filamentous, frequently branching hyphae 2.1 - 3.5 um diam, gelatinizing near surface, 
not having a clear boundary with pileus context. PILEUS CONTEXT: a tangle of 
interwoven, filamentous, undifferentiated, branching hyphae 3.5 - 10.5 um diam, with 
lengthy, thick-walled (wall thickness here and below to 1.2 tum except as noted), 
terminal or subterminal segments inflated up to 28 um diam; inflated cells thick-walled, 
plentiful, terminal singly or in chains, subglobose to pyriform to clavate to elongate to 
irregular, up to 106 x 63 um (the larger cells more often elongated than the smaller); 
branching oleiferous hyphae 2.5 - 5.0 im diam, common. LAMELLA TRAMA: 
bilateral; branching, undifferentiated, filamentous hyphae 3.5 - 5.6 um diam; inflated 
cells to 70 x 24.5 um, thin-walled, intercalary; oleiferous hyphae present, but not 
common, 3.5 - 4.9 um diam. SUBHYMENIUM: slightly inflated ramose to 
subcellular to cellular. BASIDIA: (53.5-) 60.5 - 92 x 13.5 - 16.5 (-17.5) tum, narrowly 
to very narrowly clavate, 4-spored, rarely 3- or 2-spored, thin-walled; clamps not seen. 
UNIVERSAL VEIL: On the stipe base at the surface: composed largely of gelatinizing, 
undifferentiated, filamentous, branching hyphae 4.2 - 5.9 um diam, disorderly, 
intertwining or in intertwining fascicles, some with intercalary and/or terminal 
segments slightly inflated to 19.6 tum diam; clavate terminal cells up to 77 x 25 um; 
pyriform to lachrimiform cells up to 65.1 x 44.1 tum; occasional oleiferous hyphae 3.5 - 
4.2 um diam; tissues difficult to reinflate. On the stipe base below the surface: Below 
a depth of about 1 mm, before inflation, the tissue seems mainly composed of 
intertwining fascicles of hyphae which surround minute lacunae; when reinflated, the 
tissue is seen to be composed of undifferentiated, filamentous, branching, tightly coiled 
or twisted hyphae 3.1 - 8.0 um diam with terminal and subterminal sequences of hyphal 
segments often inflated up to 14.0 um diam; scattered oleiferous hyphae 4.9+ tum diam; 
frequent inflated cells thin- or thick-walled, terminal, subglobose to ellipsoid up to 64 x 
50.5 jum and clavate up to 126 x 33 um. On the pileus: nearest pileipellis, dominated 
by globose to subglobose to ovoid inflated cells, terminal, singly or in chains of up to 4, 
thin- and thick-walled (these with wall thickness to 1.5 um), up to 61.6 x 52.5 um; 
undifferentiated, filamentous, extensively branching hyphae 2.1 - 8.4 um diam which, 
in a very thin layer just above the pileipellis, sometimes gelatinize leaving only inflated 
cells in that region. Further from pileus, hyphae becoming more curved and showing 


619 


some coiling as in veil at stipe base, with terminal and subterminal segments inflated to 
14 um diam; terminal subglobose to clavate cells thick-walled, up to 56 x 29.4 um. 
STIPE CONTEXT: acrophysalidic; branching, filamentous, undifferentiated hyphae, 
2.8 - 4.9 um diam; oleiferous hyphae 10.5 - 12.6 um diam, not common; dominated by 
thick-walled acrophysalides up to 215 x 52 um (most narrower); clamps very rare. All 
tissues pale yellow in 2% KOH or 10% NH,OH. 


Figs. 2-5 Amanita protecta. 2. Diagram of stipe base and universal veil with inner 
layers of volva largely carried away on the pileus. 3. Diagram of stipe base and 
universal veil with most of the interior of the universal veil remaining attached to its 
dessicated external surface. 4. Universal veil elements from patch on pileus (Tulloss 
1-3-87-BS1). 5. Elements of hymenium and subhymenium (Tulloss 1-3-87-BS1). The 
bars in Figs. 4 and 5 represent 20 um. 


620 


BASIDIOSPORES: [140 spores from 7 specimens measured in Melzer’s reagent] 
(8.7-) 9.4 - 12.6 (-14.7) x (7.3-) 8.4 - 11.2 (-13.3) pum, (average length (per specimen) = 
10.3 - 11.4 um; average length (overall) = 11.0 um; average width (per specimen) = 9.3 
- 10.2 xm; average width (overall) = 9.7 um; Q = 1.0 - 1.23 (-1.43); average Q (per 
specimen) = 1.09 - 1.19; average Q (overall) = 1.13), inamyloid, thin-walled, hyaline, 
smooth, globose to subglobose to broadly ellipsoid, occasionally ellipsoid, sometimes 
slightly expanded at one end, frequently adaxially flattened or slightly so; contents 
guttulate; apiculus sublateral, truncate conic; white in deposit. 


Distribution and habitat: Solitary to gregarious under Quercus agrifolia Née, 
December to March, in coastal southern California to 800 m elev. 


Collections examined: CALIFORNIA, Los Angeles Co., Santa Catalina Island, 
Grand Canyon, 27.xii.1920 L. W. Nuttall 988 (F, as A. velosa), Santa Monica Mtns., 
Stunt Canyon, 3.i.1987 Barry Silver [Tulloss 1-3-87-BS1, Wright 1383A] (holotype, 
NY), 10.1.1987 B. Silver [Wright 3701], Riverside Co., Santa Ana Mins., El Cariso, 
17.11.1987 G. Wright 3701A, Slaughterhouse Canyon 3.iii.1987 G. Wright 3712, Santa 
Barbara Co., Santa Ynez Mtns., Los Prietos Campground, 30.11.1979 Florence Nishida 
[Wright 1383]. 


DISCUSSION 


The most remarkable macroscopic feature of A. protecta is its universal veil. In 
some specimens the veil appears to have a tough outer layer covering a second layer of 
floccose-felted material which is separated from the pileus by a third, pulverulent layer. 
Microscopically, the sometimes hardened surface is revealed to be a desiccated layer of 
tissue nearly identical to that underlying it; the portion of the veil nearest the pileus is 
composed mostly of small inflated cells, sometimes in chains, which make up the fine 
pulverulent layer visible where no volval patch is present. In Section Vaginatae, this 
form of universal veil is known only from the present species. 


The upper limit for the length of basidia in A. protecta considerably exceeds that 
for any taxa described by Jenkins (1986). 


To our knowledge, the presence of thick-walled inflated cells in the pileus and stipe 
contexts and in the universal veil has not been commonly recorded in Amanita. We at 
first considered that this might be a character which evolved to resist desiccation; 
however, the situation cannot be so simple; the character is found also in A. mortenii 
Knudsen & Borgen (known only from southwestern Greenland) (Knudsen & Borgen, 
1987), in at least the hyphae of A. peltigera Reid (Reid, 1980), and in A. phalloides 
(Fr.) Link in Wildenow in which the character has been observed in English, North 
American, and Australian collections according to Reid (1980) and Tulloss (1988). In 
Tulloss’ current studies of Amanita section Amidella, thickened cell walls have proven 
to be rather common. 


In the field, if the limb of the volval sac were completely broken away, one might 
mistake this species for a member of Section Amanita close to A. pantherina (DC. : Fr.) 
Krombh. because the specimen would appear to have a cothurnate, even abrupt, bulb; 
however, the peculiar character of the volval material remaining on the pileus, the dark 
gill margins, and the dark fibrils on the exannulate, completely elongating (bulbless) 


621 


stipe should serve to prevent such an error. 


Amanita constricta Thiers & Ammirati is a species of section Vaginatae with 
grayish to brownish pileus, grayish stipe, and grayish margins to its lamellae (Thiers & 
Ammirati, 1982). This entity can be distinguished from A. protecta by having a 
sulcate to tuberculate sulcate pileus margin and a thinner, constricted and flaring, 
membranous universal veil that reaches up one third to one half of its stipe and bruises 
reddish or salmon when moist. Arora (1986: 289) provides a good photograph of the 
universal veil of A. constricta. The lamellae of A. constricta turn gray or grayish with 
age. 


Amanita pseudovaginata Hongo has some macroscopic similarities to A. protecta 
(gray material on stipe and gill margins and a relatively robust stipe base), but is 
smaller according to Hongo (1983). Also, the spores of A. pseudovaginata are much 
narrower (7 - 9 (-9.5) tum) and the basidia shorter (40 - 58 um). 


Nuttall 988 was determined originally to be A. velosa (Pk.) Lloyd; this latter species 
has a pallid, yellowish to orangish pileus; a relatively pointed stipe base; narrower 
spores; no marked tendency to ochraceous staining in the tissues; a more membranous 
volva which does not break up into pulverulence and floccose patches on the pileus and 
which tends to leave a single membranous calyptra over the disk; etc. 


ACKNOWLEDGMENTS 


We are grateful to Dr. David T. Jenkins, University of Alabama at Birmingham, for 
his review of this paper and for the opportunity to discuss matters of presentation in 
some depth. We thank Dr. Cornelis Bas, Rijksherbarium, Leiden, The Netherlands, for 
reviewing the article and for advice on selecting the specific epithet. Dr. Gregory M. 
Mueller, Field Museum of Natural History, Chicago, Illinois, graciously hosted Tulloss 
at F and provided a loan of material for which we extend our thanks. We thank Mr. 
Barry Silver, Arcata, California, for supplying two collections; Ms. Allison Petrilla, 
Roosevelt, New Jersey, for assistance with Latin; Ms. Mary A. King, Roosevelt, New 
Jersey, for assistance with document preparation; and Mr. Neal Macdonald, Princeton, 
New Jersey, for preparing the illustrations. 


LITERATURE CITED 
Arora, D. 1986. Mushrooms Demystified. 2nd ed. Ten Speed Press, Berkeley. xii + 959 


Bas, C. 1969. Morphology and subdivision of Amanita and a monograph of its section 
Lepidella. Persoonia 5(4): 285-579. 

Hongo, T. 1983. Notulae mycologicae (18). Memoirs of the Faculty of Education, 
Shiga University, Natural Science 33: 37-41. 

Jenkins, D. T. 1986. Amanita of North America. Mad River, Eureka. vi + 198 pp. 

Knudsen, H. and T. Borgen. 1987. Agaricaceae, Amanitaceae, Boletaceae, 
Gomphidiaceae, Paxillaceae and Pluteaceae in Greenland. Arctic and Alpine 
Mycology II. G. Laursen, J. F. Ammirati and S. A. Redhead, eds. Plenum, 
New York: 235-253. 

Kornerup, A. and J. H. Wanscher. 1978. Methuen handbook of colour. Methuen, 
London. 252 pp. 


622 


Marr, C. D. 1979. Laccase and tyrosinase oxidation of spot test reagents. Mycotaxon 9: 
244-276. 


Munsell Color. 1975. Munsell soil color charts. Baltimore. 

Reid, D. A. 1980. A monograph of the Australian species of Amanita Pers. ex Hook. 
(Fungi). Australian Journal of Botany. Suppl. Ser. 8. 96 pp. 

Thiers, H. D. and J. F. Ammirati, Jr. 1982. New species of Amanita from western North 
America. Mycotaxon 15: 155-166. 


Tulloss, R. E. 1988. Amanita sinicoflava: a new species from eastern North America. 
Mycotaxon 32: 421-431. 


Vergeer, P. P. 1986. The Meixner test evaluated. Mcllvainea 7(2): 61-68. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 623-635 January-March 1989 


CONTRIBUTION TO THE STUDY OF THE 
MYVAUMYCETES IN. SPAIN, 1). * 


GABRIEL MORENO, CARLOS ILLAWA & MICHEL HEYKOOP 
Department of Plant Biology (Botany). 
University of Alcala de Henares. Madrid. Spain 


ABSTRACT 


44 species of Myxomycetes are reported from several localities of peninsular Spain, The following 
taxa are new to Spain; Badhamta dubia Nann,-Srem,, Oiderma chondrfoderma (de Bary & Rostaf, 36, 
Lister and Qsdymiuw karstensiz Nann,-Brem, Two new species are described: QYacheupsis nannengae, 
characterized by ifs spherical and warted spores (9-11 pm diam,) and its abundant brown filamentous 
capillitium, and Dsdymium marinert, characterized by its limeless stipes, its whitish convex 
columella and its warted spores with a darker zone (9-10 pm diam,}, All these tava are fully 
described and figured with microphotographs of their most important taxonomical features, 


INTRODUCTION 


In addition to our recent research (MORENO & al., 1987; HEYKOOP & 
al., 1988) we start with this paper a series of studies on the 
taxonomy and chorology of the Myxomycetes of the spanish peninsula. 

The microphotographs were made under a Nikon microscope model 
Optiphot, with an incorporated system of automatic photography and 
under a Zeiss stereoscopic microscope model DRC. The microscopical 
examinations were made in a 5% KOH solution with a pretreatment in 96* 
alcohol. 

The material examined is kept in the herbarium of the Department of 
Plant Biology (Botany) of the University of Alcala de Henares (H. AH). 


CATALOGUE OF SPECIES LISTED ALPHABETICALLY 
Amaurochaete atra ‘Alb. & Schwein.) Rostaf. 


AVILA: Casavieja, on a dead branch of Ainus pinaster, 4-V1-1938 
Sanchez & C, Illana, H,AH 11044, 


&, Fransesch, P, 


; 


Arcyria cinerea (Bull.) Pers. 
MADRID: Somasierra, on dead wood of Setula alba, 24-Vi-1988, ¢, Jilana, G, Moreno & 
F, Esteve-Aaventds, H,AX 13020 and 11036, 


Arcyria incarnata (Pers.) Pers. 
GUADALAJARA: Centenera, on a carpophore of Jrameteas trogiy, 6-X1-19287, 2, Mustieles $ 
AR, Maratilla, #,AK 10952, 


MADRID: Somosierra, on dead wood of Corvlus avel/ana, 24- VI-1988, ¢, JiJana, H,AH 


! 


* This paper was presented at the "IV Reunion Conjunta de Micologia" 
celebrated in La Manga, Murcia (3-5 October 1988). 


624 


i Alcala de Henares (Finca La Oruga), on soli in a meadow in poplar grove, 30-I- 
1988, ¢, J//ana, R,AN 10947, 


Arcyria obvelata ‘Oeder) Onsberg 
SEGOVIA: Mountain gass of la Quesera, on dead wood of Fagus sylvatica, 16-1k-1982, &, 
Noreng, H,AH 11033, 


Arcyria pomiformis ‘(Leers) Rostaf. 
CIUDAD REAL: Sierra de Riofrio, on a branch of Betula parvioracteata, VS-VI1I-1982, &, 
Morano, §, Rivas-Martinez &8 4, Pernado, 4,AR- 19935, 7 


Badhamia dubia Nann.-Brem. (Fig. 1-4) 
FONTEVEORA: La Lanzada, internal part of the bark of Fyrnuws sp,, }4-V-1983, JZ, 
Gonzalez, H,AH 11002, 


20um 


10um 


Figs, 1-4, Badhamta dubia: Sporocysts and spores (HK,AH 13002) 


Comments: this species is characterized by its crowded, sessile, 
greyish-whitish with abundant CaCO= sporocysts, 0,5-1,5 mm dian. 
g- 


Spores in tight clusters, globose to subglobose, il “fim diem, 


625 


strongly warted. The capillitium is typically "badhamioid" and 
whitish. 


This is a new contribution to the national catalogue of Myxomycetes, 
Gracia (pers. comm.) has collected it in Catalufia on bark of Olea 
europaea. 


Badhamia foliicola Lister 

CASTELLON; Qropesa del Mar, on a branch of Frunus amygdalus, 3-1-1984, @, Moreno 
H,AH 10931, 

MADRID; Mountain pass of Canencia, on dead leaves of CYstus Jadanifer, e-Xti-1983,_ &, 
Moreno & F, Esteve-Raventds, H,AH 10932, Hdmera, on rests of Foaceae, 28-1X-1986, 8, 
Moreno, H,AH 11032, 


Badhamia panicea (Fr.) Rostaf. 
MADRID; Cercedilla, stem of AvJanthus altissima, 16-X1-1982, @, Moreno, H,AH 10930, 


Badhamia utricularis (Bull.) Berk, 
MADRID; Mountain pass of Canencia, on the bark of Quercus pyrenaica, ct RRR: Lk ey 
Morena & F, Esteva-Raventds, H,AK 10916 and 10923, 


Badhamia versicolor A. Lister (Fig. 5-8) 

MADRID; Miraflores de la Sierra, on the bark of (mus minor among lychens and mosses, 
B-X1I-1983, 6, Moreno & F, Esteve-Raventds, 4,AH 10909 (duplo in the private herbarium 
of Nannenga~Bremekamp sub n2 16120) and 10922, 


Figs, 5-8, -Badhamia versicolor: Sporocysts and spores (H,AH 10922) 


Comments: our collections present sessile sporocysts, scattered to 
gregarious, yellow to whitish-yellowish, 0,3-1 mm diam. Spores in 
hollow clusters, warted on the outside nearly smooth on the inside of 
the cluster, which desintegrate in part under slight pressure into 
free spores. Comparison with other specimens seems to indicate that 


626 


there are two forms of B. versicolor, viz. one with spores in hollow 
regular and rather irregularly ovoid clusters of ovoid spores, and the 
other with more or less spherical only, hollow clusters of relatively 
shorter, turbinate spores (Nannenga-Bremekamp, pers. comm. ). 

This species is very close to B. nitens Berk. because they both have 
spores with "caps" of warts on the outside and they are smoother to 
smooth on the inside of the cluster. It differs from it because B. 
nitens presents more or less solid (and not hollow) clusters of, 
usually darker, spores. i 


Craterium leucocephalum (Pers.) Ditmar 


MALLORCA: Gorg Blau, on leaves of @uercus riex, 2-V-1983, /7,2, Srgurer, H,AH 10933, 


Cribraria vulgaris Schrader 
PONTEVEDRA: La Lanzada, on branch of Arnussp,, 14-V-1983, A, Cruz, H,AH 10934, 


Diacheopsis nannengae Moreno, Illana & Heykoop, sp. nov. (Fig. 9-13) 
Etymology: in honour of Mrs. N.E. Nannenga-Bremekamp, because of her 
contribution on the knowledge of Myxomycetes. 


Sporangia 0,4-1 mm diam., globosa, sessilia aut seiuncta aut 
gregaria, argentea, iridescentia. 
Sporae sphaericae 9-11 ym diam., violaceo-brunneae, verrucosae. 


Capillitium filamentis brunneis intermixtis constitutum aut laevibus 
aut nodosis, 1-2 ym diam. Plasmodium ignotun. 

Habitat: tn cortice ramorum mortuorum Pini sylvestris, Montejo de la 
Sierra (Madrid), 15-IV-1963, leg. G. Moreno, Holotypus H.AH 11005. 


Sporocysts globose, 0,4-1 mm diam., sometimes shortly prolate, 
sessile, scattered to gregarious. Peridium very fine, membranous, 
silvery or iridescent with blueish, purplish or violaceous tints. 
Dehiscence irregular and generally from the apex. 

Spores spherical, 9-11 pm diam., violet-brown to dark brown in mass, 
in transmitted light showing distinct warts which are irregularly 
distributed, small crests can be observed under phase-contrast 
microscope, which sometimes coalesce into a broken reticulation. 
Capillitium abundant, brown, connected to the base of the sporocyst, 
forming, under the microscope, a tangle of wide and irregular net of 
anastomosing filaments , sometimes with big concolorous ellipsoidal 
nodes (e.g. 20 x 5 um. Plasmodium not observed. 

Habitat: on the bark of a dead branch of Finus sylvestris, Montejo 
de la Sierra (Madrid), i5-I1V-1983, G. Moreno, Holotypus H.AH 11005. 
(Isotypus in the private herbarium of WNannenga-Bremekamp sub n® 
15064). 

Comments: Diacheopsis nannengae is characterized by its rather small 
spores (9-11 m diam.) combined with its abundant brown capillitium 

It is close to D. picninica Krzemieniewska, but the latter has even 
smaller spores (6-7 pm diam.) and its sporocysts are cylindrical (vide 
KOWALSKI, 1975). With D. depressa Thind & Lakhanpal, as described from 
India, it shares the spore size (9-10 pm diam.) but it differs from it 
in its brown not hyaline capillitium, (KOWALSKI, 1975), 

We place this new species in the genus Diacheopsis instead of in 
Colloderma because the latter presents a gelatinous peridium. No trace 
of this gelatinous layer has been observed in the sporocysts of our 
species, not even under phase-contrast microscope. 


627 


Figs, 9-13,- Ofacheapsis nannengae: Sporocysts, spores and capillitium (Kolotypus! 


628 


629 


Diderma chondrioderma (de Bary & Rostaf.) G. Lister (Fig. 14-20) 
MADRIO; Alcala de Henares, on bark rests of Fopulus alba, 15-X1-1987 
11013, 


Comments: our collection presents stalked sporocysts, 0,4-0,8 mm 
diam., discoid to umbilicate, depressed around the stipe apex, whitish 
to slightly greyish and with abundant lime deposits. Stipe whitish ta 
pale straw-yeliowish, up to 0,5 mm of length. Columeila globose, 
whitish to very paie straw colour. Spores smooth, spherical, 11-12,5 
pm diam., purple-grey and densely warted. Capillitium hyaline to very 
paie brown, 1-3 um dian. 

This is a new contribution to the national catalogue of Myxomycetes. 


Diderma spumarioides ‘Fr.) Fr. 

MAORTD; Alcala de Henares (Puente ¢uiema), on Avaceae in pine grove of PYnus 
Aalepensis, 5-11-)988, J, Prieto, HAR 10396, 

MALLORCA: Road from Liuch ta Soller, on leaves and branches of @uercun flax, 25-I- 


$963, J, Siguier, 4,AH 10937, 


Didymium bahiense Gottsberger em. Nann.-Brenm. 
MADRIO; Alcala de Henares (Quinta Cervantes}, on spike of Je3 mays, 9-X1-1987 
fliana, H,AR 10945, 


al 


i L, 


Figs, 21-24.- Ofdvarum karstensii: Sporocysts and soores (HAN 10993) 


‘ 


Didymium clavus (Alb. & Schwein.) Rabenh. 


630 


SEGOVIA; Mountain pass of Los Leones, on a dead branch of Saatolina rasmarinifolla, 
8-1V-1984, 6, Moreno, H,AH 10926, 


Didymium difforme (Pers.) Gray 

MADRID: Alcala de Hanares (Quinta Cervantes), on stems of A/thaea rosea, 15-X11-1993, 
C, [ilana, HAR 10919, Alcala de Henares (Quinta Cervantes), on dead wood of Cercis 
sildiguastrum, \3-V-1388, ¢, Ji/ana, H,AH 11030, 


Didymium karstensii Nann.-Brem. (Fig. 21-24) % 
MADRID: Alcala de Henares (Finca La Qruga), on leaves of Populus alba, 7-V-1988, F, 
Esteve-Aaventds & C, Iilana, 4,AX 10993, 


Comments: This species is characterized by its sessile whitish to 
greyish-whitish sporocysts with a tipically wrinkled-rugose double (at 
least below) peridium. The spores are subglobose, 10-12 um diams;" and 
their ornamentation agrees exactly with the one described by NANNENGA- 
BREMEKAMP (1974). 

This is a new contribution to the national catalogue of Myxomycetes. 


Didymium marineri Moreno, Heykoop & Illana, sp. nov. (Fig. 25-33) 

Etymology: in honor of Prof. Sebastian Mariner-Bigorra, who died 
recently and collaborated with us during fifteen years making several 
latin diagnosis. 


Figs, 25-27,- Oidyvarum marinerz: Sporocysts (Holotypus) 


Sporangia 0,4-1 mm diam., globosa, stipitata, albo-cinericia. 


631 


Figs, 28-33,- Ofdymium mariners, Spores and capillitium (Holotypus) 


632 


Columella convexa et alba. Stipes 0,4-0,7 mm longus, brunneo-ruber. 
Sporae sphaericae 9-10 ym diam., fortiter verrucosae aliquibus zonis 
obscurioribus. Capillitium 1-2 ym diam., aut hyalinum aut pallide 


luteo-brunneun, nodulis trregularibus et dichotomice desinens. 
Peridium calcis crystallis aggregatis constitutum. Plasmodium ignotum. 
Habitat: in truncis. Phoenicis canariensis, Oropesa ~ del Mar 


(Castellon), 19-I1V-1987, leg. G. Moreno, Holotypus H.AH 11040. 


Sporocysts deeply umbilicate below at their union to the stipe, 0,4- 
lmm diam., scattered to gregarious. Peridium granulose, whitish from 
the abundant white lime crystals on it, with irregular and generally 
apical dehiscence. Columella whitish variable, often flat or convex, 
rarely globose. Stipe 0,4-0,7 mm long, reddish-brown, paler at the 
apex and without lime crystals. Plasmodium not observed. 

Spores brown in mass, 9-10 wpm diam., spherical and densely warted 
with a well defined darker zone. Capillitium hyaline to pale 
yellowish-brown, 1-2 ym diam., with some irregular nodes, branched 
dichotomously, united to the peridium in part with some expanded tips. 

Habitat: on rests of petioles (‘still on’ the stem) ote ricenz: 
canariensis, QOropesa del Mar (Castellon), 3-1-1984, G. Moreno, HAH 
11014 .y 11015. ‘Ibidem, 4-1-1984, HAH” 11041,  Ibidemiyeiyatya ear, 
Holotypus H.AH 11040. (Collected on several palm-trees very distant 
from each other, from different gardens of this locality). Paratype in 
the private herbarium of Nannenga-Bremekamp sub n& 16.066, and in MA- 
funed. 

Comments: Didymium marineri is characterized by its well defined 
whitish convex columella and by its warted spores (9-10 pum diam.) with 
a darker zone. This species is similar macroscopically )ta Dd: 
squamulosum ‘Alb. & Schwein.) Fr. which differs in its typically 
whitish and corrugated strongly calcareous stipe. D. minus (Lister) 
Morgan and D, melanospermum (Pers.) 7. Macbride are different because 
they have a brown to dark brown, and not whitish, columella and 
peridium. D. bahiense Gattsberger em. Nann.-Brem, can be segregated by 
its long stipes which are ¢ to 3 times longer than the diameter of the 
sporocyst, and by its slightly bigger spores (9)10-12¢14,5) pam, 
NANNENGA-BREMEKAMP (1974). 


Didymium melanospermum (Pers,) T, Macbride 

MADRID: Montejo de la Sierra, on a branch of FProws sylvestris, 15-1V-1983 
H,AH 11004, Mountain pass of la Quesera, on dead leaf of Fagus sylvatica, 3 
Marana, HAR 11000, 


&, Morena, 
~X- 


I-X-1986, 6, 


Didymium minus (Lister) Morgan 
BURGOS: San Juan del Monte, on humus of Quercus rotwadifolia, Pinus halapensis and 
Cistus sp,, 3-X-1987, F, Esteve-Aaventds, HAH 11042, 


Didymium squamulosum ‘Alb. & Schwein.) Fr. 

MADRIO: Montejo de la Sterra, on leaves of Saliv sp,, 1Q-X-1987, @, Moreno, HAH 
10997, Miraflores de la Sierra, on leaves of (mus minor, 8-X-1984, G, Moreno, HAH 
11003, Alcala d@ Henares (Quinta Cervantes), on dried foaceae, 9-X1-1987, ¢, JlJana, 
HAH 10942, Ibidem, on fruit of Ceres siliguastrum and leaves of UVlaws minor, 17-XI- 
1987, HAH 10343, Ibidem, on dried Foareze, 18-VI-88, ¢, J/i/ana, H,AH 11617, Ibidem, 
15-V-1988, ¢, JiJana, HAR 11018, Ibidem, on needles of Fraws pines and fruits of 
Cercis siliguastrum, \7-X1-\987, ¢, J//ana, HAH 11019, Robregordo (Mountain pass of 


633 


Bumosierra), on leaves of Betula fontqueri, 1-X-1986, R. Galan, fF. 
Esteve-Raventos & G. Moreno, H.AH 11016 


Didymium vaccinum ‘Durieu & Mont.) Buchet 
MADRID: Alcald de Henares (Quinta Cervantes), on spike of ea mays, 9-XI-1987, C, 
f/lana, 4,AH 10346, [bidem, 17-X1-1987, RL AR 10948, 


Enteridium lycoperdon Bull. 
GRANADA; Sierra Nevada, on a stem of Salix sp,, i-VI-1982, @, Moreno, WN, Petnado $ 
JM, Losa-@uintana, #.AR 3992) 


Fuligo septica ‘L.) Wigg. 

MADRID; Alcala de Henares, (Finca La Orugad, on a stump of Populus a/ba, 14-V-1988 
N,0, Hedo, E, Galer2 $ C, Ilana, BL Ad 11037, 

AVILA: Casavieja, on a Stem and needies of Pinus pinaster, 4-VI-1988, & Fransesch & 
€, fllana, A.A 11026, 11027 and 11029, 

SEGOVIA: Mountain pass of la Guesera, on dead wood of Fagus sylvatica, 16-1X-1982, 6 
Morena, H,AK 11025, 


Leocarpus fragilis (Dickson) Rostaf. 
MALLORCA: Road from LLuch to Saller, on humus of @uercus flay, 25-1-1383, /2, 
Srguter, H,AH 10925, 


Licea. minima Fr. 
ASTURIAS: Cangas de Narcea (Reserva Sioldgica de Muntellos), on the fruit-body 
hymanophore of Piptoporus betulinus, 5-1-1953, 4.7, Martinez, KLAN 10311, 


Lycogala flavofuscum ‘Ehrenb.) Rostaf. 
MADRIO: Alcala de Henares (Finca La Oruga}, on a stumo of Populus 2/da, among leaves, 
3i-X-1987, ¢, Ji/ana, H,AH 10944, 


Lycogala epidendrum L. 

AVILA; Casavieja, on a : branch of Finaws pinaster, a-VI-1988, &, Fransasch, F, 
Sdochez $C. fllana, RAN 11035, 

MADRID: aera on sis wood of Corvlus avellana, Z4-VI-1566, ¢, Jilana, 6, 
Morena & F, Esteave-Aaventds, BAR i024, 


Mucilago crustacea Wiggs. 

GUADALAJARA: Tamajén fe 
Moreno, 4,AKW 10936, Chilios 
Rielo, HAR 10943, 


}, on Thymus vulgaris, \-X-1984, @, 
28 under Fopulus alba, 25~X1-1987, €, 


eanebra 


los es 
on pests of a re 


= 
a oo 


MADRIO: Alcala de Henares (Cuesta Zulema}, on needies of Pinus falepensis, 22-XI- 
1987, ¢, filana, H,AH 10941, Somosierra, on dead wood of a Sroad-leaved tree, 24-VI- 
1 o a 


938, ¢€. Illana, 8, Moreno § F, Estave-Raventds, WLAN 11028, 


Physarum cinereum (Batsch) Pers. 
MADRID: Alcala de Henares (Quinta Cervantes), on foaceae, /7-81-/887, C, L1fana, Wax 
1103) 


Physarum compressum Aib. & Schwein. 

CASTELLON: Oropesa del Mar, on rests of dried leaves (still on the stem) of Phoenix 
canariensis, 28-X11-51, @, Merenoa, A,An 10920, 

MADRID: Alcala da Henares, on a dead leaf af broag-leaved trea, 15-V-i585, C, Liliana, 
HAH 1101} 


634 


Physarum leucophaeum Fr. 

MADRID: Mountain pass of Canencia, on a stump of Aetu/a alba, 8-X11-1983, @, Morano & 
F. Esteve-Raventds, H,ARK 10914, 10917, 10918 and 10927, Ibidem, on dried rests of 
herbaceous plants and on leaves and fruits of Cystus Jawrifolrus, 4,AHN 10928, 


Physarum nutans Pers. 

AVILA: Casavieja, on Folyporus sp, on Quercus suber, 4-VI-1988, B&B Fransesch § C, 
filana, H,AH 11039, Montejo de la Sierra, on wood of @wercus pyrenaica, 24-V1-1388, ¢, 
Iilana, @, Moreno § F, Esteve-Raventds, HAR 11022, Mountain pass of Somosierra, on 
bark of a dead branch of feftwla alba, 24-VI-1988, ¢, Jilana, G, Moreno & F, Esteve- 
Raventds, HAH 11021, 


Physarum pezizoideum (Jungh.) Pav. & Lag. var. microsporum Farr 
GUADALAJARA: Valfarmoso de Tajufa, on carpophores of dAuricusaria masentarica, |0-XI- 
1985, 7, Martin, H,AK 10910, 
Only known in Spain from one other collection from the province of 
Madrid, (NANNENGA-BREMEKAMP & LADO, 1985). 


Physarum pusillum (Berk. & Curt.) G. Lister 
MADRID; Alcala de Henares (Quinta Cervantes), on Poaceae, /7-#1-1987, C, illana, 4, 4A 
10950, Ibidem, on a dead leaf of Jrrs germantca, H,AK 10951, 


Physarum straminipes Lister 

CASTELLON: Qropesa del mar, on rests of Phoenty canariensis, 3-1-1984, G, Moreno, 
H,AH 11014 and 17015, 

MADRID: Alcala de Henares (Quinta Cervantes), on a stem of Ai/anthus altissima, 2\-1I- 
1988, ¢, JiJana, HAR 11010, 


Physarum vernum Sommert. 

CASTELLON: Oropesa del Mar, on a dried stem of Fhoenty canartensis, 2-1V-1986, &, 
Morano, H,AH 10923, 

MADRID: Alcala de Hanares (Quinta Cervantes), on branches of Arlanthus alfissina, \7- 
X1-1987, ¢, Jidana, H,AK 10939, 


Stemonitis fusca Roth 
CIUDAD REAL: Sierra de Riofrio, on leaves of Betula parvibracteata, \1-VI-1982, 6, 
Moreno, M, Pernado & A, Velasco, H,AK 10924, , 


Stemonitis virginiensis Rex 

GUADALAJARA; Embalse de El Vado, on a branch of Pinus pinaster, 4-1V-1981, @, Moreno 
& JL, Manson, H,AK 10338 and 11001, ; 

CIUDAD REAL: Sierra de Riofrio, on trunk wood of Aetula parvifracteata, 5-VI1I-1982, 
G, Moreno, &, Rivas-Nartinez $ Petnado, 4,AK 10912 and 30913, 


Symphytocarpus flaccidus (Lister) Ing & Nann.-Bren. 
AVILA; Casavieja, on piled branches of Pinus pinaster, 4-V1-1988, FP, Sanchez, B, 
Fransesch & €, Lldana, H,AK 11043, 


Trichia varia (Pers.) Pers. 

MADRID: Mountain pass of Canencia, on a stump of Betula alba, 10-X-1984, 6, Moreno $ 
F, Esteve-Raventés, H,AH 10940, Montejo de la Sierra, on woody rests of Fagus 
sylvatica, 15-IV-1983, G, Moreno, HAH 11034, 


Tubifera ferruginosa (Batsch) J.F. Gmelin 


635 


SEGOVIA; Nacimiento del rio Moros, on stump and needles of Praus sylvestris, 1-VII- 
1987, @ Moreno & F, Esteve-Raventds, HAM 10424, 10425, 10426 and 10426, 


ACKNOWLEDGEMENTS 


We wish to express our gratitude to Mrs. Nannenga-Bremekamp for 
revising part of the material examined and for carefuly going through 
the manuscript. We are also greatly indebted to Prof. Granada-Godoy 
for the translation of the diagnosis into latin. 


REFERENCES 


-Heykoop, M., Illana, C. & Moreno, G. (1988). Nueva aportacién al 
estudio de los Myxomycetes de Alcala de Henares (Madrid). Bol. Soc. 
Micol. Madrid 12:3-8 

~Kowalski, D.T. (1975). The genus Diacheopsis. Mycologia 67: 616-628. 
-Moreno, G. Heykoop, M. & Illana, C. (1987). Interesting Myxomycetes 
Tmemenentcate de Henares (Madrid). Bol. Soc, Micol. Madrid 11:213- 
216 

~Nannenga-Bremekamp, N.E, (1974), De Nederlandse Myxomyceten. 
Koninklijke Nederlandse Natuurhistorische Vereniging. 460 p. 
-Nannenga-Bremekamp, N.E. & Lado, C. (1985). Notes on some Myxomycetes 
from Central Spain. Froc. Kon. Ned. Akad. Wetensch. Sér. C. 88: 219- 
esl. 


MY COTAXON 


Vol. XXXIV, No. 2, pp. 637-642 January-March 1989 


A NEW OPHIOSTOMA ON POLYPORES 


O. Constantinescu & S. Ryman 


Uppsala University, The Herbarium 
Pee Ue boxo04 ts, S-751 21) Uppsala, Sweden 


Abstract. Ophiostoma polyporicola n. sp., 


is described from several collections on 
polypores from Sweden and Finland. Both a 
Sporothrix anamorph and the  teleomorph 
were obtained in cultures initiated from 
ascospores. 


Two polypores, Fomitopsis pinicola and lyromyces stipticus, 
colonized by an Ophiostoma, were collected several times in 


Sweden and once in Finland over the years. Cultures obtained 
from the ascospores of two recently collected specimens 
produced a Sporothrix anamorph. This fungus differs from the 
known species of Ophiostoma by a number of characters and is 
here described as new. 


Cultures were initiated by inoculating ascospores collected 
in droplets at the tip of the ascoma neck on malt extract 
‘agar (2% malt extract and 1.5% agar per litre) on a modified 
malt agar according to Upadhyay (1981) and on oatmeal agar. 
Cultures were incubated at 20-22°C in darkness for 5 days 
followed by up to 30 days incubation at the same tempera- 
ture under near ultraviolet and cool daylight on a regime of 
Age hi’. 


Ophiostoma polyporicola 0. Const. & Ryman, sp. nov. - Figs 
late 


Ascomata solitaria vel caespitosa; basis nigra, globosa, 
feevis.. 200-300 um diam.; rostrum nigrum, 750-2250 (im 
longum, ad basim 45-75 wm, ad apicem 17-30 ym latum; hyphae 
ostiolares absentes. Asci non visi. Ascosporae hyalinae, 
aseptatae, lunatae, utrinque rotundatae, 3-3.5 x 1-1.5 [m; 
vagina gelatinosa egentes. 


638 


a 


Fig. 1. Ophiostoma polyporicola. a. Ascomata in culture; b. 
ascomata on Tyromyces  stipticus (S. Ryman 6102, photo I. 
Holmasen) sic. ascomalinecks)) do sectioned) acenne wall; e. 


conidiogenous cells and conidia; conidia: ascospores 
from culture. (a, c-g from UPSC 1704). 


639 


Anamorphosis ad genus Sporothrix pertinens. Cellulae 
conidiogenae hyalinae, continuae, 10-40 x 1-2 um; conidia 
hyalina, laevia, clavata, plus minusve curvata, ad basim 
truncata, 6-14 x 2-2.5 um. 


Holotype: on Fomitopsis _pinicola, Sweden, Uppland: Vange 
par., Fiby urskog, 13 Sept. 1985, S. Ryman 8069 (UPS). 
Living culture ex type: UPSC 1704 = CBS 669.88. 


Mycelium not apparent. Ascomata single or in groups on 
the basidiocarps of the hosts, mostly immersed in the tubes 
feesaittang at ‘the pore surface, but also’ on the: tissues 
surrounding the pores; base black, globose, smooth, 200-4500 
um diam; wall 15-25 um thick, composed of an outer layer of 
2-3, brown, thick-walled, irregularly shaped cells, 4-10 um 
diam, and an inner layer of 1-2, paler and more thin-walled 
cells; neck black, 750-2250 um long, 45-75 um wide at the 
eee weeiradially tapering) to 17-30 pum at the tip, 
occasionally with 1(-2) swellings and often with a subapical 
constricted ring; neck composed of parallel, 2-5 wm wide 


Fig. 2. Ophiostoma_ polyporicola. Conidiogenous cells, 
conidia and ascospores from culture. a. from UPSC 1/04; b. 
Peom UPSC 2555. 


640 


hyphae, brown or dark-brown and with various thickenings of 
the walls at the base of the neck but paler and smoother 
toward the tip; ostiolar hyphae absent. Asci evanescent, 
not seen. Ascospores’' hyaline, 1-celled, Jlunate with 
rounded ends in side view, 3-3.5 x 1-1.3 um, without 
gelatinous sheath, collecting in a milky droplet at the tip 
of the neck. Scattered conidia and a few conidiophores and 
conidia similar to those obtained in culture (described 
below) were noticed in three specimens. 


Colonies attaining a diameter of 1/-18 mm after 12 days 
and 36-45 mm after 30 days, at first smooth but later 
covered with radially oriented hyphal strands or synnema- 
like structures, ochreous; margin submerged, fimbriate; 
exudate and diffusible pigment absent; reverse with a 
yellowish tinge. Submerged advancing hyphae 1-1.5 um wide, 
older hyphae up to 2.5 um. Conidiogenous cells arising 
from undifferentiated hyphae, continuous, 10-40 um long, 1-2 
um wide, slightly tapering towards the tip, with several 
terminal and intercalary, 1-2 um long and O.5-1 wm wide 
conidium-bearing denticles; conidiiferous part sometimes 
Slightly inflated. Conidia hyaline, smooth and thin- 
walled, clavate, more or less curved, 6-14 x 2-2.5 um, base 
truncate, c. 1 um wide. Ascomata and ascospores morphologi- 
cally similar to those occurring on the natural substratum 
are formed after 25-30 days on 2% malt extract agar, and to 
a lesser extent on oatmeal agar. 


OTHER MATERIAL EXAMINED (all in UPS). 


On Fomitopsis pinicola. FINLAND: Kuusamo: Kuusamo par., 5 km 
NE of Ruka, 26 Aug. 1978, N. Lundqvist 11 7S6.q5 Swain: 
Smaland: Varnamo, Kavsj6, 13 July 1987, 0. Constantinescu & 
W. Gams (UPSC 2355 = (CBS 670.88, living ee 
Vastmanland: AngsG par., close to the castle of Angs6, 21 
May 1969, R. Santesson 21171b; Gastrikland: Gavle, Valls- 
hage, 11 Oct. 1969, J. A. Nannfeldt 20712b; Angermanland: 
Nordingra par., Villmyran Nature Reserve, 17 June 1982, N. 
Lundqvist 13782. 

On Tyromyces stipticus. SWEDEN: Uppland: Stockholm, 4 Nov. 
1894, L. Romell; Uppland: Vange par., Fiby urskog, 3 Oct. 
1780," o.eRhymaneG 102% 


DISCUSSION 


Having short, lunate ascospores with rounded ends, deprived 


of a gelatinous sheath, Ophiostoma polyporicola should be 
placed in the Pilifera group of Ceratocystis (Olchowecki & 


641 


Reid 1974), or in the section Ophiostoma of the same genus 
(Upadhyay 1981). However, by consistently forming a 
Sporothrix anamorph in culture, its place is in Ophiostoma 
as defined by de Hoog 1974, Weijman & de Hoog 1975, Samuels 
& Muller 1979 and de Hoog & Scheffer 1984. 

There are three species of Ophiostoma having’ some 
resemblance with 0. polyporicola: 0. epigloeum (Guerrero) de 
Hoog differs by smaller ascomata with shorter neck and 
Beraight and shorter . conidia, 0.-.angusticollis Wright & 
Griffin has a faster growth rate, smaller ascomata with 
narrower neck, reniform ascospores and much smaller conidia, 
and 0. nigrocarpum (Davis) de Hoog has a faster growth rate, 
much smaller ascomata and smaller, straight conidia. 

The anamorph of OQ. polyporicola should be compared with 
other Sporothrix species having clavate, more or less curved 
conidia with truncate base. Only Sporothrix curviconia de 
Hoog has some resemblance with our fungus, but it has a 
Slower growth and smaller conidia. In addition, 5. 
curviconia has a second type of Jlateral, guttuliform 
conidia, lacks a teleomorph and is of tropical origin. 

Most species of Ophiostoma are saprophytic on wood, a few 
are parasitic, and only occasionally they are found on other 
Substrata (Upadhyay 1981). Only three species have been 
described from fungi: O. epigloeum (Guerrero) de Hoog (1974) 
on tTremella fuciformis, and O. roraimense and O. grande, 
these last described by Samuels & Miller (1979) from 
stromata of Diatrype. All three differ from 0. polyporicola 
in the characters of both the telomorph and the anamorph. 
Griffin (1968) found an Ophiostoma on Fomitopsis pinicola in 
Canada and considered it identical with Ceratocystis 
perparvispora Hunt, now known as Ophiostoma_microsporum 
(Davids.) v. Arx. The examination of this specimen (SSMF 
#25-6126-01 =.MFB 7405). showed that it fits O. microsporum 
indeed, but the conidial state on the accompanying dried 
culture, described by Griffin l.c. as ‘conidia abundant, 
produced directly on the mycelium’ is Acremonium olidum W. 
Gams (1971). Two living cultures of Ophiostoma microsporum 
(CBS 315.77 and 440.69) were also compared with our fungus. 
These strains showed a superficial resemblence with O. 
polyporicola regarding the culture characteristics, but they 
produce broader ascospores and typically reniform conidia, 
as depicted by De Hoog (1974) and Maekawa et al. (1987). 

Ophiostoma polyporicola us often associated with 
Acremonium olidum W. Gams and Hypocrea pulvinata Fuckel. 
Gams (1971) mentioned this association when he described 
Acremonium olidum from a polyporaceous fruitbody. Moreover, 
he noticed the presence of an Ophiostoma (Ceratocystis) in 
fie ctype specimen. 


642 


Acknowledgements. We thank the Curator of CBS _ for 
providing two strains of Ophiostoma microsporum, Dr. D. |. 
Myren for the loan of Ceratocystis perparvispora, Mrs Ulla- 
Britt Sahlstrom for photographic assistance, and Mr l. 
Holmasen for providing the photograph used as fig. 1 b. Drs 
W. Gams ‘and G. ‘S.°-de ‘Hoog critically “reévieweceienies 
manuscript. The senior author gratefully acknowledges the 
financial support of the Swedish Natural Science Research 
Council and: the » Swedish Council for -(\Forestryeaeecs 
Agricultural Research. 


LITERATURE CITED 


Gams, W. 1971. Cephalosporium-artige Schimmelpilze (Hypho- 
mycetes). Stuttgart: Gustav Fischer. 

Griffin, H. D. 1968. The genus Ceratocystis in Ontario. Can. 
J BO.) 462 609-7 16. 

Hoog, G.:S. de, 1974. The genera BlastobotryisiSpororncuss 
Calcarisporium and Calcarisporiella gen. nov. _ Stud. 
Mycol., Baarn 7: 1-84. 

Hoog, G. 5S. de & Scheffer, R: J. 1984. Ceratocyettemveren= 
Ophiostoma: A Reappraisal. Mycologia 76: 292-299. 

Maekawa, N., Tsuneda, A. & Arita, I. 1987. Ceratocystis 
species occurring on the Lentinus edodes bedlogs. Rep. 
lotteriy mycol -osinstw 25: 6-147 

Olchowecki, A. & Reid, J. 1974. Taxonomy ‘Gfthewigenges 
Ceratocystis in Manitoba. Can. J. Bot.°523 16/5= i 

samuels, G.. T. & MUller,.E. 1979. Life-historyeee. cee 
Brazilian Ascomycetes 5. Sydowia 31: 169-179. 

Upadhyay, H. P. 1981. A Monograph of Ceratocystis and 
Ceratocystiopsis. Athens: University of Georgia Press. 
Weijman, A. C. M. & Hoog, G.S. de 1975. On thewenhaaeee 
of the genus Ceratocystis. Antonie van Leeuwenhoek 41: 

353-360. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 643-645 January-March 1989 


LITHOGRAPHA, A LICHEN GENUS NEW TO 
CONTINENTAL NORTH AMERICA 


JoAnn W. Flock 


Herbarium, University of Colorado, Boulder, CO 80309 


During the summers of 1978, 1979, and 1980 the author made 
extensive collections of the lichens of Cape Prince of Wales on the tip of 
the Seward Peninsula at the Bering Strait in Alaska. Among the crustose 
lichens collected on dark grey, fine-grained schist inland from the coast 
was an interesting lichen with lirellate apothecia and a grey areolate thallus. 
The lichen has been identified as Lithographa tesserata which has been 
reported for Sweden, England, Wales, Germany, Austria, Italy, and 
Rumania by Redinger (1937); in the Arctic and Caucasus of the U.S.S.R. 
by Kopaczevskaja et al. (1977); and the Narssaq District of Greenland by 
Alstrup (1979). Neither the genus or the species has been reported for 
Continental United States or Canada in the Fifth Checklist of Lichens 
(Egan, 1987). Dr. W.A. Weber of the University of Colorado Herbarium 
agrees with the identification, and Dr. John W. Thomson of the University 
of Wisconsin confirms that the genus and species have not previously 
been reported for the Alaskan-Canadian Arctic. 


The species is very distinctive and easily recognized but may not be 
spotted without the aid of a hand lens. The Alaskan specimen covered 
only a small area and did not stand out against the dark substrate. The 
species may be restricted to extreme western Alaska, but it may also have a 
wider distribution over the Continental United States and Canada and been 
overlooked. The following descriptions of the genus and species are 
presented with the hope that other lichenologists will look for the species _ 
and enhance the possibility that it might be found elsewhere in the 
Continental United States and Canada. 


The following descriptions have been provided by Dr. Thompson: 
Lithographa Ny]. 


Acta Soc. Linn. Bordeaux 21:393. 1856. Placographa Th. Fr. Gen. 
Heterolich. 95. 1861. 


644 


Thallus crustose, finely granulose to warty or areolate. Apothecia 
round to elongate with open slit-like disk or widening; exciple black; 
paraphyses branched, often gelatinizing; asci clavate; spores 8 or many, 
simple, hyaline, ellipsoid. Algae Pleurococcus. 


Lithographa tesserata (DC.) Nyl. 


Acta Soc. Linn. Bordeaux 21:441. 1856. Opegrapha tesserata DC., in 
Lam. & DC., Flor. Franc. 3:313. 1805. Placographa tesserata Th. Fr., 
Lich. Scand. 1:636. 1874. 


Thallus thick, granular, warty or smooth to chinky areolate with 
angular 1 mm broad areoles, or with rounded raised portions with a black 
hypothallus, the areoles pale to dark brown or whitish gray, dull, K+ 
yellow becoming red, C-, the hypothallus present in var. petraea (Ach.) 
Redinger, absent or nearly so in var. nivalis (Th. Fr.) Zahlbr. Apothecia 
dispersed, slightly projecting over the areoles; with the base narrowed and 
the upper edges sickle-like and slit-like turned inwards making the mouth 
nearly closed; the exciple thick, black; a thin dark brown epithecium is 
granulose; the hymenium hyaline to brownish, not inspersed, I+ blue; 
paraphyses slender, non-capitate, nearly unbranched; asci clavate; spores 
8, uniseriate to biseriate, ellipsoid to ovate, simple, hyaline, 9-15 x 5-8 
micrometers. 


Fig. 1. Lithographa tesserata (DC.) Nyl. 
(COLO L-83471). Bar = 1 mm. 


645 


Acknowledgments 


I wish to thank Dr. W. A. Weber for confirmation of the 
identification of Lithographa tesserata and Dr. John W. Thomson for 
confirmation of the new record for the genus and species for the North 
American Arctic and for the English translation of the genus and species 
descriptions. Photographic work was supported in part by BRSG Grant 
RRO7013-16 awarded to the Department of Molecular, Cellular, and 
Developmental Biology, University of Colorado, Boulder by the 
Biomedical Research Grant Program, Division of Research Resources, 
National Institutes of Health. The Bering Strait study was sponsored by 
the Museum, University of Colorado, Boulder. 


Literature Cited 


Alstrup, V. 1979. Notes on selected Greenlandic Lichens. Bot. Tidsskr. 
74:155-163. 

Egan, R. S. 1987. A Fifth Checklist of the Lichen-Forming, 
Lichenicolous and Allied Fungi of the Continental United States and 
Canada. The Bryologist 90:77-173. 

Redinger, K, 1937. Graphidaceae in Rabenh. Kryptogamen-Flora 9,2(1): 
181-404, Lithographa 187-198. 

Kopaczevskaja, E. G,, M. F, Makarevicz, and A. N. Oxner. 1977. 
Handbook of the Lichens of the U.S.S.R. 4. Verrucariaceae- 
Pilocarpaceae, Lithographa tesserata 216-217. 


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MY COTAXON 


Vol. XXXIV, No. 2, pp. 647-653 January-March 1989 
EES, ERs BREE Tet 2S RENE PS ee oe canes I MT reese oP) Eee Oe Sa OS OT aS Disa 


NOTES ON ONE LICHENICOLOUS 
AND ONE FUNGICOLOUS DISCOMYCETE 


WEN-YING ZHUANG and RICHARD P. KORF* 


Systematic Mycology & Lichenology Laboratory 
Institute of Microbiology, Academia Sinica, Beijing, China 


1. A New Combination for a Skyttea on Lecanora effusa. 


In connection with the study on Unguiculariopsis thallophila (Karst.) 
Zhuang (1988), which is on Lecanora carpinea, the description of 
another interesting lichenicolous discomycete, Mollisia lesdainii (Vouaux 
in Bouly de Lesd.) Vouaux (1914), drew our attention; this was also 
reported on a Lecanora, L. effusa. The morphological characteristics 
described by the original author are similar to those of U. thallophila. 
To discover whether both names refer to the same fungus was our 
original purpose. 

First published as a lichen, Nesolechia lesdainii Vouaux in Bouly de 
Lesdain (1910), the fungus was later transferred to Mollisia as M. 
lesdainii (Vouaux, 1914). It was collected by Bouly de Lesdain on 
Lecanora effusa on Salix sp. in Dunkerque, France. The name later 
appeared in a list of Vouaux’s fungal parasites of lichens prepared by 
Prof. Y. Rondon (1970). In order to locate the type specimen of this 
fungus the junior author contacted Dr. D. L. Hawksworth in England and 
Dr. J. Astier in Marseilles, France, for Prof. Rondon’s office and home 
addresses. We were then able to write Prof. Rondon, but unfortunately 
received no answer. Through the continuing help of Dr. J. Astier we 
finally received the type specimen two years later. Although the senior 
author has published her monographic study on the genus 
Unguiculariopsis we think that it is worthwhile to prepare a note on the 
taxonomic position of this rare fungus because it was very difficult to 
locate the type specimen and because perhaps no attention will be paid to 
the fungus otherwise. 

Our type examinations of M. lesdainii and of U. thallophila indicate 
that both fungi are similar in the shape of asci and of ascospores and in 
possessing brown ectal excipular cells forming an ectal excipulum of tex- 


* Present address and reprint requests: Plant Pathology Herbarium, Comell University, Ithaca, NY 14853, USA. 


648 


tura angularis. But M. lesdainii is different from U. thallophila in 
having urceolate apothecia submerged in the lichen thalli instead of 
having cupulate apothecia which are superficial at maturity; in possessing 
very few hair-like cells which are short, straight, slightly tapered towards 
the apex, and distributed only on the inner face of the margin 
(“summit”), instead of possessing well-developed hairs with a fine, 
hooked apex and a swollen base distributed both at the margin and on the 
flanks; in possessing ectal excipular cells not loosely interconnected and 
with ectal excipular tissues slightly gelatinized, instead of loosely 
interconnected cells and non-gelatinized tissues. 

Sherwood et al. (1980) published the genus Skyttea to accommodate 
a small group of lichenicolous discomycetes which have small, urceolate, 
immersed, dark brown or olive green, non-carbonized apothecia, small- 
celled, pseudoparenchymatic ectal excipulum, with hyaline to brown, 
thin-walled, smooth hairs on the inner face of the margin, filiform 
paraphyses, thick-walled, J— asci with the apex thickened, and 0-3-septate 
ascospores. Mollisia lesdainii possesses many features of Skyttea except 
for the morphology of the ascus apex which is not obviously thickened. 
But the ascus morphology is similar to that of Skyttea cruciata Sherwood 
et al. We think that M. lesdainii fits Skyttea better than any other genus 
and transfer this fungus to Skyttea. In passing, it should be noted that 
Sherwood et al. (1980) had referred Unguiculariopsis thallophila to 
Skyttea, which the senior author later transferred to Unguiculariopsis 
(Zhuang, 1988). 

Mollisia lesdainii differs from the somewhat similar S. cruciata in 
the shorter asci, smaller ascospores, shape and length of marginal “hairs,” 
larger ectal excipular cells, and occurrence on the thalli of Lecanora 
effusa and not on Diploicia canescens. 


Skyttea lesdainii (Vouaux in Bouly de Lesd.) W.-y. Zhuang & Korf, 
comb. nov. (Fig. 1) 
= Nesolechia lesdainii Vouaux in Bouly de Lesd. (‘lesdaini’), Rech. 
Mich Dunks pee bo 10) 
= Mollisia lesdainii (Vouaux in Bouly de Lesd.) Vouaux (‘lesdaini’), 
Bull. Soc. Mycol. France 30: 180, 1914. 


Apothecia urceolate when young and cupulate at maturity, submerged 
in lichen thalli, sessile, solitary or a few in a group, dark brown, 0.1-0.2 
mm in diam when dry. Ectal excipulum of textura angularis, 10-20 um 
thick, cells brown, subglobose, 5-8(-10) sm in diam, tissue slightly 
gelatinized, cells on inner face of margin (“summit”) elongating to form 
short hairs or hair-like cells, ca. 10-15 tum long, straight, tapered towards 
the apex, wall slightly thickened. Medullary excipulum of textura 
intricata, thin, hyphae subhyaline. Asci inoperculate, 8-spored, clavate, J— 


649 


Fig. 1. Skyttea lesdainii (holotype): a. anatomy of an immersed 
apothecium on lichen thallus, x 370; b. hairs, x 1760; c. two asci, an 
ascus apex and two paraphysis apices, x 1760. 


650 


in Melzer's Reagent, walls slightly thickened at apex, 23-35 x 5.7—7 um, 
arising from croziers. Ascospores irregularly biseriate, ellipsoid, hyaline, 
non-septate, biguttulate, S-7.5 x 1.8—2.2 tm. Paraphyses filiform, septate, 
not exceeding asci. 

On thalli of Lecanora effusa. 

Holotype examined: on thalli of Lecanora effusa on Salix sp., 
Dunkerque, France, Bouly de Lesdain, 23 IX 1903, Vouaux Herbarium, 
from Prof. Y. Rondon (personal herbarium?), courtesy of Dr. J.-Astier. 


2. A New Species of Unguiculella on Diaporthe sp. 


An interesting discomycete was discovered on the fine beaks of old 
fruit bodies of a Diaporthe species in Jamaica. The apothecia of this 
fungus are extremely small and not easy to detect with a hand lens in the 
field. Our laboratory observations show that the receptacle surface of this 
fungus is covered with short, glassy hairs which are characterized by a 
swollen base and a fine, hooked apex, that the ectal excipulum is of 
textura prismatica to textura angularis, and that two kinds of paraphyses 
are present in the same apothecium, one with a normal apex and the other 
with a hair-like, elongate, hooked apex. 

Judging from the morphology of the Jamaican fungicolous 
discomycete, Unguiculariopsis and Unguiculella are genera in which the 
fungus might possibly be accommodated. The habit on fungi and the 
hooked hairs fit the characteristics of Unguiculariopsis, whereas the 
excipular structure of textura prismatica to textura angularis and the 
presence of hair-like, hooked paraphyses are not consistent with the 
generic concept of Unguiculariopsis in which the ectal excipulum is of 
textura globulosa to textura angularis and the paraphyses are filiform 
with a normal apex (Raschle, 1977; Korf & Kohn, 1980; Zhuang, 1988). 
In early drafts of the senior author's monograph of Unguiculariopsis 
(Zhuang, 1988) this species was tentatively placed in that genus. Our later 
studies indicate that it is better treated as an Unguiculella. By error the 
name Unguiculariopsis jamaicensis appeared once in the thesis 
publication (Zhuang, 1988). 

Presence of both hooked hairs and tapering, hooked paraphyses is 
clearly a diagnostic feature of Hohnel's original concept of Unguiculella 
(Hohnel, 1906). Moreover, a few species of Unguiculella, U. aggregata 
(Feltg.) Hohn., U. meliolicola Dennis, and U. oregonensis (Kanouse) 
Dennis, are also reported on fungi (Kanouse, 1941; Dennis 1955; S. 


Fig. 2. Unguiculella jamaicensis (holotype): a. shape of apothecium, from top to 
bottom showing hymenium, medullary excipulum, and ectal excipulum; b. structure of 
ectal excipulum at flanks; c. hairs; d. ascospores; e. two asci, an ascus apex, and two 
kinds of paraphyses; a x 100, b-e x 1300. 


652 


Huhtinen, personal communication). The ectal excipular structure of the 
Jamaican fungus is very similar to these three Unguiculella species. 
Although two kinds of paraphyses have not previously been reported in 
any species of Unguiculella, we think that most features of the Jamaican 
collection are like Unguiculella. 

U. oregonensis is possibly the closest species to the Jamaican 
discomycete. But the Jamaican species can be distinguished from U. 
oregonensis by the much smaller apothecia, which are faintly vinaceous 
instead of white and cupulate instead of flat to convex when rehydrated, 
by wider asci and longer ascospores, and by the presence of normal 
paraphyses in addition to tapered, hooked paraphyses. We treat the fungus 
as anew species of Unguiculella. 


Unguiculella jamaicensis W.-y. Zhuang & Korf, sp. nov. (Fig. 2) 


Ab Unguiculella oregonensi apotheciis minoribus, ascis latioribus, 
ascosporis longioribus, et paraphysibus normalibus filiformibus 
praesentibus una cum paraphysibus gradatim angustatis uncinatis apice, 
etiam presentia in Diaporthae specie differens. 


Apothecia cupulate, sessile to subsessile, 0.1-0.15 mm in diam when 
fresh; hymenium faintly vinaceous when fresh, receptacle surface 
furfuraceous, somewhat whitish. Hairs covering the entire receptacle 
surface, thin- and smooth- walled, with a swollen base and a fine curved 
apex, pale brown at the base and subhyaline at the apex, not becoming 
purple in Melzer's Reagent with or without 10% KOH pretreatment, 
nonseptate, with lumen extending to the apex, 13-23 um long. Ectal 
excipulum of textura prismatica to textura angularis, 11-22 Um thick, 
cells brown, thin-walled. Medullary excipulum of textura intricata, 
hyphae subhyaline to pale brown. Subhymenium indistinguishable. Asci 
inoperculate, 8-spored, clavate, J— in Melzer's Reagent, 35-45 x 6.6-8.8 
um. Ascospores biseriate, ellipsoid, hyaline, nonseptate, mostly 
biguttulate, some multiguttulate, 7.8-10.2 x 3.0—-3.5 tm. Paraphyses 
filiform, septate, some with a normal apex and some with an elongated, 
tapering, hooked apex, normal paraphyses not exceeding asci and hooked 
paraphyses exceeding asci, normal paraphyses and the lower part of 
hooked paraphyses ca. 1 um wide. 

On fruit bodies of an old Diaporthe sp. 

Holotype: on beak and other superficial parts of fruit bodies of a 
Diaporthe species on a branch, near entrance to Wallingford Coffee 
Industry Development Co., above St. Peters, St. Andrew Parish, Jamaica, 
3750 ft. elev., R. P. Korf, T. Iturriaga, and W.-y. Zhuang, 8. XII. 1986, 
CUP-MJ 1013. 


653 


ACKNOWLEDGEMENTS 


We deeply thank Dr. J. Astier, Marseilles, France for locating the 
type specimen of the lichenicolous discomycete, Dr. W. J. Dress of the 
Bailey Hortorium, Cornell University for providing the Latin diagnosis, 
and Dr. G. J. Samuels of the New York Botanical Garden for identifying 
the host fungus of Unguiculella jamaicensis. The junior author gratefully 
acknowledges financial assistance from the Systematic Mycology and 
Lichenology Laboratory, Academia Sinica, that made his stay and studies 
in China in the autumn of 1988 possible. 


REFERENCES 


Bouly de Lesdain, M. 1910. Recherches sur les Lichens des Environs 
de Dunkerque. Société Dunkerquoise. Dunkerque. 301 pp. 

Dennis, R. W. G. 1955. Fungi from Sierra Leone: Pezizales and 
Helotiales. Kew Bull. 1955: 363-368. 

Hohnel, F. von. 1906. Revision von 292 der von J. Feltgen 
aufgestellten Ascomycetenformen auf Grund der Originalexemplare. 
Sitzungsber Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl., Abt. 1, 
115: 1189-1327. 

Kanouse, B. B. 1941. New and unusual species of Discomycetes. 
Mycologia 33: 461-467. 

Korf, R. P. & L. M. Kohn. 1980. Revisionary studies in the 
Hyaloscypheae. I. On genera with “glassy” hairs. Mycotaxon 10: 
503-512. 

Raschle, P. 1977. Taxonomische Untersuchungen an Ascomyceten aus 
der Familie der Hyaloscyphaceae Nannfeldt. Sydowia 29: 170-236. 

Rondon, Y. 1970. L’herbier des champignons parasites des lichens de 
Pabbé Vouaux. Rev. Bryol. Lichénol. 36: 737-745. (1969). 

Sherwood, M. A., D. L. Hawksworth, and B. J. Coppins. 1980. 
Skyttea, a new genus of odontotremoid lichenicolous fungi. Trans. 
Brit. Mycol. Soc. 75: 479-490. 

Vouaux, L. 1914. Synopsis des champignons parasites de lichens. Bull. 
Soc. Mycol. France 30: 135-198. 

Zhuang, W.-y. 1988. A monograph of the genus Unguiculariopsis 
(Leotiaceae, Encoelioideae). Mycotaxon 32: 1-83. 


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MYCOTAXON 


Vol. XXXIV, No. 2, pp. 655-665 January-March 1989 


NOTEWORTHY CORTICOLOUS LICHENS IN NOTHOFAGUS 
FORESTS, NORTH-WESTERN PATAGONIA 


SUSANA CALVELO and LAURA LORENZO 


Centro Regional Universitario Bariloche 
Universidad Nacional del Comahue 
C.C. 1336,Bariloche, 8400 RN, Argentina 


SUMMARY 


Six corticolous lichens are reported 
fomeone tirst time from Argentina: Cali— 
cium leucochlorum Tuck.var. paraguayen- 
se Sant., Haematomma hilare Zahlbr., 
Hypogymnia pulverata (Nyl.ex Crombie) 
Elix, Leioderma amphibolum (Knight) Ga- 
lloway & J¢érge., Leptogium laceroides B. 
de Lesd.and Parmeliella thysanota (Stir- 
ton) Zahlbr. Four of these species show 
Geremarkable!’ disjunct, distribution: 
Southern South America—Australasia 


INTRODUCTION 


wie bachen flora of Argentina’ in general: and 
that of the Nahuel Huapi National Park in partic 
ular has not been well investigated and the avail 
able information is very scarce (Grassi,1950a). 
fieveware studies on the foliose ‘and Eruticose 
lichens from Tucuman Province (Grassi 1950b, 
1953). Specimens collected in the Nahuel Huapi 
and Los Alerces National Parks were reported by 
Lamb(1958). Recent contributions to the lichen 
flora are concerned mainly. with the central and 
NE parts of Argentina (Ferraro, 1986; Osorio, 


MWogeeadler ,° 1987). 


656 


This paper is part of a more extensive study 
that aims at improving the knowledge of the 1li- 
chen flora in the area of NW) Patagonian 


MATERIALS AND METHODS 


The lichens for this study were collected by 
the authors. Collection-sites, their gedgrapna= 
cal position and characterization are specified 
below the description of each species. All speci 
mens are deposited’ in the Herbarium of Ghee 
tro Regional Universitario Bariloche (BCRU) ,Uni- 
versidad Nacional del Comahue, Argentina. 


The material was examined under a dissecting 
microscope for’ habit amd, morphology .euqe anatomi 
cal studies, sections 10-25 pm thick were prepad: 
ed using a freezing microtome and examined by 
light microscope after mounting in water or lace 
tophenol cotton-blue. 


STUDY AREA 


Collecting sites: The lichens describedmin 
this study were collected from the Nahuel Huapi 
National Park and surroundings, in anWareawoceae 
ed between 41°02! and 41°14! S latitude amide gem 
tween 71°19! and 71°50' W longitude )(Figaeeee 
The area has typical Andean. relief, with e@lacear 
lakes at ca.800 m altitude, surrounded by moun- 
Cains OL up! to 24005 meabovercoeam eves 


Climate: It.is characterized as perhumud@tosnue 
mid and microthermal, according to the Classi ta= 
cation of Thornthwaite (1948). The avenageman= 
nual temperature is 8°C. Average temperature for 
the warmest month (January) is 17°C and for the 
coldest (July) atsis 3°C., Lowest, tempemami ae 
vary from -15 to -25°C. These temperatures were 
measured at 760m alt. Annual average precipita- 
tion decreases abruptly from W to E, from 3000mn 
year in Puerto Blest to 1400mm/year in Cerro Car 
bon (Munozaret) ade Losses x 


657 


Fig.1: Southern South America 
(Nahuel Huapi area (*) 
indicated) 


UY 


ej;Bariloche 


41910’ 


Fig.2: Collecting sites: 1-Puerto Blest, 2- Cerro Catedral, 3- Cerro 
1dpez, 4- Cerro Carbon. 
—.-.-—.— Intemational border 
Scale 1: 500.000 


658 


Vegetation: Puerto Blest 1s includegiims: 
the Valdivian,District,. with temperate, siaeas 
open rainforest, dominated by Nothofagus dombe 
yi, typically with a dense bamboo understory of 
Chusquea culeou. Epiphytes and lianas are common. 


The slopes of Cerro Lopez and Cerro Cate- 
dral present an altitudinal vegetation gradient, 
with Nothofagus dombeyi open woodlands, frequent 
ly in association with Lomatia hirsuta and Aus- 
trocedrus chilensis, from 760 to ca. 1100 m 
above sea level. At higher altitudes, there are 
subandine woodlands of deciduous Nothofagus pu- 
milio up to the timberline, ca. 1600m above sea 
level. However, from 1500 m Nothofagus pumilio 
appears as a shrub. Cerro Carbon presents No- 
thofagus pumilio from 1000 to 1500 m alt. 


THE EL ELCHEN SPECIES 


CALICIUM LEUCOCHLORUM Tuck. var. PARAGUAYENSE 
Sant.(Fig*. 3A)'.. Santesson,SR. Arkiv) formmpowees 
NPL QO C14) see Hyves LOA 


Thallus episubstratic, upper surface verrucose, 
yellow-grey when dry, yellow-green when wet. 
Verruca 0.3 to 0.8 mm. Forming discontinuous 
patches -Without® soredia. 

Apothécia black to «brownish-black) 2) Vo mje 
high, with very well defined limit between 
stipe and capitulum. Sometimes with’ ¢greyaise 
pruina.on the Lower ,side of the.capitulug 
Stipe of constant diam. (to 0.2 :mm,). (Canasgmeeeee 
0.4 to 0.8 mm diams, 0.5 /to 0.S°mm hi eh 
plane for young apothecia and convex for ma- 
ture apothecia.. Mazaedium black. 

Asci_ cylindrical to clavate with uniseriate 
spores. Ascospores 1 or 2-celled, dark brown, 


et Ste a VAT ITN 


Habitat: Found on Nothofagus pumilio near 

the base of the tree in a disturbed forest. 
Cerro Carbon at 1250.m above sea «level. NOGwiae 
quent. 


Specimen examined:Argentina. Parque Nacional 


659 


Neewednapi ,) Cerro, Carbon, SW Sslope,col.' (Calve— 
lo, 2-V-1987 (BCRU 00022). 


Distribution: Paraguay and Argentina. 


Mises tne Trirst record from Argentina. 


HAEMATOMMA HILARE Zahlbr. (Fig. 3B). 
Zatubrmuckner,A. Denkschr. Akad. Wiss. Wein Math. 
Peteewaes . KS 10423508 1941. 


Thallus crustose, whitish, slightly shiny,spread 
over the substrate with a black line delimiting 
it. Elongated up to 8 cm. Without soredia. 
Bpothnecia Lecanorine,, sessile; circular, up to 
1.5 mm diameter. Disc orange-red, without pruina, 
surrounded by a thin margin concolorous with the 
thallus. Epithecium K+, magenta. Ascospores aci- 
cular 11-13(-15)-transversely septate,62-70 x 
4-5 pm. 

The colored apothecia are very conspicuous. 


Habitat: Found on young, smooth-barked Nothofa- 
gus dombeyi, forming circular bands round the 
trunk, usually in very humid but exposed areas. 
Puerto Blest at 800 m above sea level. 


Specimen examined: Argentina. Parque Nacional Na 
Niewmnaapr.. Puerto Blest’, path to Cascada Los 
Senger oas col. Calvelo, 14-VII-1985 (BCRU. 00044). 


Distribution: New Zealand and Argentina. 


tive ise the farst record outside New Zealand. 


HYPOGYMNIA PULVERATA (Nyl.ex Crombie)Elix (Fig. 
pour iaex, JA. Brunonia 273217. 1979. 
Parmelia mudata var. pulverata Nyl.ex Crombie, 
Oeermesoc. Lond. “Bot..15: 395, 1879. 


Pei O11 OSe  heteromerous, ‘46to'10 cm ‘diam. , 
Pierwiuacet to 5 mm wide, dichotomically branch— 
ed close together in center and more. dispersed 
peripherally. Upper surface whitish-grey, cover- 
ed with powdery soredia, especially the older 


660 


branches. Medulla white, solid. Lower surface 
black in the center, whitish-brown at the edges. 


Apothecia not seen. 


Habitat: Found on Nothofagus dombeyi near 
streams in very humid microhabitats. Cerro Cate- 
dral at 850 m above’ sea level. 


Specimen examined: Argentina. Parque National 
Nahuel Huapi, Cerro Catedral, SE slope ;c¢o Raa. 
Velo Ww 10-X>100 OR CBCRUMO004 7 


Distribution; Australasia and Argentinas 


This 1s the first record outside Austrateaswee 


LELODERMA AMPHIBOLUM (Knight) Galloway & J¢grg. 
(Fig. 3D). 

Galloway,D.J..& P.M. Jérgensen in Kantva lane 
James, Lichenologist 19:28.1987.- Pannaria amphi 
bola Knight,. Trans.’ N.Z. Inst.12: 36029 eee 
Parmeliella amphibola (Knight) Miill. Arg., Bull. 
Herb. Boissier 2 (App:.~1): 44 . 1894 )>Panmagrd 
amphibela Nyl':, Lich. Nov “Zel.: 50.916 Ga 
Pannularia amphibela (Nyl.) Hue, Nouv. Archs Mus 
Hist? nat? Paris) sen. a5) 0s eee oe 


Thallus foliose, heteromerous, up to 4 emeiiaae 
lobes 2 ‘to 3°>mm wide, iurregulary and) note, eae 
frequently branched, disposed very closely to= 
gether, sometimes imbricated. Margins undulate, 
covered with verruciform, granular Of (COma epee 
soredia. Upper surface brownish - green when 


Fig. 3: Specimens investigated 


Calicium leucochlorum Tuck.var.paraguayense Sant.. (x5) 
Haematomma hilare Zahlbr. (x7) 

Hypogymnia pulverata (Nyl.ex Crombie) Elix (x4) 
Leioderma amphibolum (Knight)Galloway & Jérg. (x4) 
Leptogium laceroides B.de lesd. (x4) 

Parmeliella thysanota (Stirton) Zahlbr. (x15) 


et st) -@) Gs > 


661 


662 


wet, brown when dry, mat, fissured, sometimes 
with laminar soredia. Cyanobiont Nostoc, packed 
in clusters, individual cells 6-10 zm diam.-Lower 
surface without cortex, \whitish at thevapwees 
and with brown tomentum, formed by short-celled 
hyphae, centrally; with white or brown rhizines 
Upson Om or 


Apothecia not seen. 


Habitat: Found on Nothofagus pumilio, near the 
base of the trees, on shaded areas. Growing on 
trees covered with mosses or diréctly(omeoeeee 
Cerro Lopez at 1200 m above sea level. 


Specimen examined: Argentina, Provincia de Rio 
Negro, Dpto. Bariloche, Cerro Lépez, E slope, 
col’. Calvelo,-4=1V-1987.. (BGRUVO004B IE 


Distribution: New Zealand, Tasmania, Argentina. 


This is the first record outside. Australasia 


LEPTOGIUM LACEROIDES B. de Lesd. (Fig. 3E). 
Bouly de Lesdain,M.., Annals /Cryptog. (Exouusse 
CGY) KenL ATED Rami LO 


Thallus foliose, homoiomerous, lobes rounded up 
to 8 mm wide, extending over the substrate, 
except for the margins that may be ascendent; 
margins incised. Gelatinous when wet, papery 
when dry. Upper surface almost black when wet, 
blue-grey when dry, smooth, with patches covered 
with numerous cylindrical to lobulate isidia.Low 
er surface pale grey, tomentose. Apotheciawlamms 
nate, 1.5%to 2.5 mm diam., disc reddish=bpewee 
Ascospores ellipsoid, muriform, 30-33 x 11-16mm. 


Habitat: Found on Nothofagus pumilio in shaded 
humid habitats, frequently near streams, on mos- 
ses. Cerro Carboén at 1050 m above sea level;Ce- 
rro Catedral at 1370 m.above sea level and Germ 
Lopez at 1320 m above sea level. 


Specimen examined: Argentina. Parque Nacional Na 
huel *Huapi, Cerro Carboén, SW slope,col.)Calweuass 
2-V-1907" (BCRU 00032). 


663 


Distribution: Cosmopolitan. 


The first record for Argentina. 


PARMELIELLA THYSANOTA (Stirton) Zahlbr. (Fig.3F). 
eteiier. A. Cat. Lich. Unive 39225. 1025.— 
Pannaria thysanota Stirton, Proc. Phil. Soc. 
ier 1059293. 1877. 


Thallus squamulose, heteromerous, squamules 
Comedie GO Clongated» 41 to (2. mm)jin) diam. ») with 
Mareins incised, créenated, toothed, loosely at- 
tachet to the substrate. Upper surface yellow- 
grey when dry to brownish-grey when wet. Lower 
surface brown centrally to white at the margins, 
with scattered rhizines. Marginal prothallus pro 
minent, blackish-blue. i 
Bpolrecitaysessile to,subpedicelLlate,’ 0.2 to, 0.3 
(-0.4) mm diam., disc red-brown, with a thick 
Mat eineeconcolorous whith. the ‘thallus. 
Aseaspores ellipsoid, 16-17 x 6-—7.mm. 


Habitat: Found on specimens of Nothofagus dombe- 
yi covered with mosses, in the more humid and 
shadiest habitats of the area. Puerto Blest at 
800 m above sea level. 


Specimen examined: Argentina. Parque Nacional Na 
huel Huapi, Puerto Blest, between Puerto Blest 
fiommearouririas, icol.Lorenzo, 14=X1-1986 | (BCRU 
OOO 35) 


Geographical distribution: New Zealand and Ar- 
gentina. 


First record outside New Zealand. 


DISCUSSION 


Peeeepeciecs, of corticolous Jichens aré’ report 
Pome@oietie tirst tame from Argentina. 

Leptogium laceroides, which is cosmopolitan 
UGxrtioway, 1965), is’ now added “to the’ known flo— 
ra of Argentina. For Calicium leucochlorum var. 
paraguayense, this find constitutes the south- 
Preamesvelocality Of ats distribution, since’ it 
MeoexnOown, up to now, from Gran Chaco, Paraguay 
(Santesson, 1943). Parmeliella thysanota and 
Haematomma hilare were known as endemic for New 


664 


Zealand Nothofagus forests (Galloway, 1985; 
Rogers and Bartlett, 1986), Hypogymnia pulverata, 
recorded previously for Australasia (Galloway, 
1985), and Leioderma amphibolum known from New 
Zealand and Tasmania cool temperate rainforest 
(Galloway and J@érgensen, 1987), are now added to 
the known flora of Nothofagus forest in Argentina . 


The last four, species mentaoned presen 
widely disjunct«circumantarctic“distripuvaeus 
Australasia - Southern South America. They have 
been found in habatats with ‘similar eeokosteasL 
characteristics, and three of them (Haematomma 
hilare, Leioderma amphibolum and Parmeliella 
thysanota) even on phorophytes of the same genus 
(Nothofagus). This distribution pattern can be 
explained by the: late. relationship BeGweemme 
areas as parts of 'Gondwanaland “and7ormmoy 
more recent’ long’ distance “dispéersaiot epee 
pagules by the West Wind Drift (Galloway, 1987) 
The observations made are further evidence of 
the resemblance of the cool temperate Nothofagus 
forest flora in cir¢eumantarctic terri ae 


ACKNOWLEDGEMENTS 


We would like’ to thank Dre, Gernot” Vobioemana 
Dr. Thorsten Lumbsch, Philipps University. 
Marburg, Germany for critical commentseonmanuae 
paper and’ Dr. Mason E. Hale, Smithsontangeine wee 
tion, Washington D.C.; U.SsA. for readingame mee 
nuscript. Atso to Mrs. Yolanda Qudtiteanaeee ot 
kindly typed) the manuscript. 


REFERENCES 


Adler,M. 1987.A New species of the genus Canopar 
melia from Argentina. Mycotaxon 28:251-254 

Ferraro,L.L. 1986.Contribution, the std yaoi 
gentina Parmeliaceae. The genus Punctelia 
Krog and Flavopunctelia (Krog) Hale.Phyto- 
Goa Oil ante tee ae Oe Dae . 

Galloway,D.J. 1985. Flora. of New Zealand igen 
New Zealand Government Printer.pp. 662 


665 


Galloway,D.J. 1987. Austral lichen genera:some 
bueeecorraphi cal problems. [nveocress 
and Problems-in Dlichenology inthe Bast 
Res a Dibl ws echenols 1252, 865-200; 7 

Calloway,D.J. and P.M. Jé¢rgensen. 1987. Studies 
one coew lichens tamily  Pannarvaceac Li. 
The genus Leioderma Nyl. Lichenologist 
19(4): 345-400. 

eee 19504. Contriubucion al Catalogo de \li- 
quenes argentinos. Lilloa XXIV: 5-296. 

Graget.M.)1950b. Los liquenes foliosos y fruticu 
ieeos de Tucuman. (ad loa xX LV: 207-295. 

SieesieM. 1953.leloschistaceae de Tucuman.Lilloa 
AVE Sana sO) 815 2, 

Pembet-M. 1958. La vegetacion liquénica de los 
Parques Nacionales Patagonicos. Anales 
de Parques Nacionales VII.pp.1388. 

Mange oy A.Garay. 1955.Caracterizacion clima- 
mrcade aba ce roy . derkio, Negro. LUNTA. 
EERA. Bariloche.pp 56. 

Pearce. LOS. Contribution to the ltchen 
meOroanOlwArSeN Uy ia \ Velie pbiblbnosrapay 
covering 1950-1985. Comunicaciones Bota 
Wwieas’Mus. de Hist .Nat. Montevideo, 5 
Go 2 ltd” 

Rogers, R.W. and J.K.Bartlett. 1986. The lichen 
genus Haematomma in New Zealand. Liche- 
Molec ste bo(93)): 2477 2 55°. 

Peaimesaon., Rk. LOAS. South American, Calicia. Arkiv 
for shotanwky 3014 jas pl-a12:- 

Thornthwaite,C.W. 1948. An approach toward a 
Pemvonawe cased. ical Lon Ol) Culmaue. 


Geor eRev. esol): §55=—04:; 


Mt 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 667-677 January-March 1989 


GIGASPORA RAMISPOROPHORA: A NEW SPECIES WITH 
NOVEL SPOROPHORES FROM BRAZIL 


JOYCE LANCE SPAIN 


Se NiO bel oi Guat Gas an 1S 
Pasi aeolian eer aglvGr Pes 1500 


EWALD SIEVERDING 
Institut ftir Pflanzenbau in den Tropen 
Universitaet Gottingen, Grisebachstrasse 6 
D-3400 Gottingen, West Germany 
NORMAN C. SCHENCK 
Plant Pathology Department 


tives weylof Florida; Gainesville,’ Florida ’32611 


ABSTRACT 


Gigaspora ramisporophora develops sporophores 
(term suggested ae HAG eh ee bearing structure 


in the Endogonaceae) with morphological variation 


from other described species in the genus. The 
sporophore Mey DeeaSaamipreree ih arveliycn aseLeriin oe 
sporogenous cell (term recommended to replace 
Momeni ot bike cell lj), or branched,» forming 
multiple (2-3) sporogenous cells. A protrusion 
On a sporogenous cel may give rise to a branch 
with a spore. The spore ws ‘characterized hy va 
hyaline to sub-hyaline wall over two pigmented 
walls. pnevmiddle wall is wlaminate . (Germ tubes 


emerge from the germinal wall (term introduced to 
describe a wall distinctive in both texture and 
function) and directly penetrate the outer walls. 


SUMARIO 


Gigaspora ramisporophora desenvolve esporéforos 
(termo Sugerido para estruturas que sustentam o 
€sporo nas Endogoneceae) com caracteristicas 
morfolégicas que variam de outras espécies 
descritas no género O esporéforo pode ser 


simples, tendo uma Gnica célula esporogénica 


668 


(termo recomendado para substituir célula de 
sustentacao), ou ramificado,  formando Gerurae 
esporogénicas maltiplas, (2-3). Uma protuberancia 
na célula esporogénica pode desenvolver uma 
rane) fic ac aro. Com ie elSsOrooon 0 (e'sip omonee 
caracterizado pela presenga de uma parede hialina 
a sub-hialina sobre duas paredes pigmentadas. A 
parede mediana’€@ laminada. Tubos de @pgemminacag 
emergem da parede germinal (termo introduzido 
para descrever uma parede distinta em textura e 
fungao) e penetram directamente nas paredes 
externas. : 


INTRODUCTION 


Spores of an undescribed speciés” Of Gt eae ue. 
were recovered by wet sieving soil and root 
samples from ‘plots ‘of Pueraria phaseovoumwees The 
golden yellow to yellowish-brown spores varied 
greatly in size and some were borne on branched 


S/p,0'rs0' Di Ouse: Six The, novel, features eo weere 
sporophore are described in this papers The 
suggested terms “"“sporophore’’, “Wsie Ween 
Endogonaceae by Tandy (1975). and Perea.) 
Herrera (1980).,-.-and <"sporogenows /§@ eu eoeee 
discussed. The spore wall description follows 


the terminology of Walker (1983 )=y ieee 
exception of “germinal wall”. 


GIGASPORA RAMISPOROPHORA Spain, Sievengipewe. 
Schenck Sp Rinoveoriroures 14 


Spora formatur plerumque in apicem@ee ae 
sporogenae. Sporae lutae vel fulvae, globosae, 
Oe a pore oC! um diam... vel. subgiopusace 

143-)150-400 “X, 200-450 (501) (ise edd Totus 
paries sporae 9+31 Um Crassus, (C Ot) See 
stratis tribus: stratum exterior, 1) 4-490) 
tenue; mediranum, lamellatum, 4-28 “wg 


interior 1.3-2.6 jm tenue cum papillisiieagneeea 
ad stratum secundum vel separatum in regione quo 
spora verminat. Sporophori plerumque singulari 
Vel ee Ditpart tt vel Sure art 1 ¢ isa noSie Sporophori 
cum cellula inflata. terminant; cel Puasa 
quae cellula sporogena est, (32-)40-60(-72) um 
lata et (50-)60-80(-83) um longa; unaewrarewaee 
cellulae inflatae sub cellula $ pO 
Sporophori novi non rare ex cellula sporogena 
progerminant. Cellulae auxiliares usque ad 20 in 
fasiculo humi vel in cortice” radiciarumveeeeee 
stercore herbarum formantur. Cellulae auxiliares 
primo leves deinde cum projectionibus digitoidis, 
UU SAy es es OU A IN uae O tio od fae Spora <direchetoe. 
parietem germinans. Endomycorrhizam cum 
arbusculis Vil wore Ciooe bit Sr. 


669 


Figure 1. Gigaspora ramisporophora. A. Young 
spore (YS) on sSporophore SaaS SET emanating 
from sporogenous cel SC) attached to immature 
spore (IS) 160X. B. Colonized organic matter 63X 
C. Spore walls: unit (U); laminated (L); germinal 
132 with papillae (P) 400X. D. Sporogenous cell 
SC) with contiguous bulbous cell (CBC) 400X. E. 
Auxiliary cells: ornamented (0); young (Y) 400X. 
F. Germ tube (GT) 400K. G. Branched sporophore 
with two sporogenous cells: young (Y); detached 
from spore (D) 250X. 


670 


SPORE, usually formed at apex of bulbous 
sporogenous cell, smooth, golden yellow to 
yellowish-brown, globose, (96-)200-450(567) wm to 
sub-globose, (143-)150-400 X 200-450(—-501)- pm; 
total wall thickness 9-31 wm. 


SPORE) WALL STRUCTURE, (Figs..11-C (4) 330 feecn ee 
walls in a single group. Spores measured in 
water or fixed in FAA and mounted in lactophenol. 
Walid’ 1, ‘unit wall, hyaline to subshveaire. 
brittle; 1.4-4.0(-—5.7). wm thick, continugeneawao 
outer wall of sporogenous cell and usually 
adherent to wall 2; wall 2 laminate, yellow to 
yellowish-brown, 4.0-28 wm thick, adherent to 
wall 3; outer part of wall 2 continuous with 
middle wall of sporogenous cell; wall 3, germinal 
wall, yellow to yellowish-brown, 1.3-2.6 wm thick 
excluding papillae, 1.4-8.3 pm diam. and up to 
5.6 pm high, which develop prior to germination. 
Walls 2 and 3 are reactive to Melzer's reagent 
turning reddish-purple. Spore contents finely 
granular. 


BULBOUS SPOROGENOUS CELL (Fig. 2), apical or sub- 
apical cell of sporophore, (32-—) 40-60 (— 72a 
(50-)60-83 pm usually with three walls totalling 
6-11(-14) wpm thick: outer wall hyaline; laminate 


middle wall and innermost wall brown. Wall 
thickness variable, usually greater at spore 
initial site. Sporogenous cells may develop a 


branch (sporogenous cell with connecting hypha) 
which gives rise to another spore (Fig. 1-A). 
The sporogenous cell may have thick-walled, 3-4 
pm, hyphal protrusions (pegs), 8-10 pm diam. A 
thin-walled hypha, 0.7-3 pm, with 6-8 pm dianm., 
may extend from a thick-walled protrusion or 
develop directly from the sporogenous cell wall. 


SPOROPHORE (Figs. 1-G, 3) light brown, simple or 
branched with 1-3 sporogenous cells, formed of 
specialized septate hypha, 9.3-13.9 ym diam., 
wall 1.4-2.8 wm thick (excluding bulbous cell 
measurements); distance between septae variable 
but septation more frequent near sporogenous 
cell. One or rarely two bulbous cells, some with 
hyphal protrusions, may be contiguous with apical 
sporogenous cell (Figs. 1-D, 2 I-L). 


671 


s 


a 
ie 


ak 
ri¥ 


Rogie 2 *. Sporogenous cells of Gigasporophora 
ramisporophora: A-F) apical sporogenous cells; 
weenesporogenous cells with thick hyphal 
protrusions (pegs) or slender hyphae; G-H) sub- 
apical sporogenous celis; I-L) sporogenous cells 
with contiguous bulbous cells. Camera lucida 
drawings. 


U 
io HP -_ 
ie 
ce es 


672 


Figure 3. Sporophores of Gigaspora 
ramisporophora: A) young spore forming from 
sporogenous cell on simple sporophore; B) 
branched sporophore with three sporogenous cells, 
two with spores detached (qd); C) branched 
sporophore with two sporogenous cells, one with 
spore detached (d). Camera lucida drawings. 


673 


Figure 4. Murograph: Gigaspora 


ramisporophora. One group, A; 
three walls: unit, laminated & 


germinal. 


VUYUUYUY) @ 


\auncemnrangaapactrcesense 
A 


Se tierARYeCELLS (Fig../1sE)-tan,? round to clavate, 
Sa4eedenum diam.; smooth initially becoming 
ornamented with digitate projections (rarely bi- 
Poeeatejmup, to, 10 im. high; borne in clusters of 
less than 20 on coiled hyphae, 6-8 pm diam., with 
Ehime walls, 0.5-1.3 ym; coils develop on. short 
hyphal stalks or along the coenocytic hyphae. 
Miem@eietarnvecelis found in the mineral soit, 
decomposing organic matter and in the cortex of 
roots. 


DIRECT GERMINATION (Fig.1-F): one to several germ 
tubes (1-8; x 4) emerge from the germinal wall 
where the papillae are largest. The germ tube 
within the spore wall is 5.6-9.7 pm diam. and at 
egress 14-17(-22) wm diam. 


ENDOMYCORRHIZAE formed with arbuscules. 


TYPE obtained from a pot culture with Sorghum 
halipense, previously seeded with P. phaseoloides 
Benth, culture CP 75, at the Centro de Pesquisas 
Rome ama. (CEPEC.)., .lrtabuna,. Bahia, .Brasil. 
Holotype FLAS; Isotypes OSC, GOET and IBt. 


ETYMOLOGY: Latin, rami (branch), referring to the 
sporophore configuration. 


Tiorr BUTTON, HABITAT AND MYCORRHIZAL 


ASSOCIATIONS: Gigaspora ramisporophora is known 
from a single site in southern Bahia. Spores 


associated with P. phaseoloides were collected 


674 


from a CEPEC forage seed production field 2 km 


east of Itapebi. Decomposing plant residues were 
frequently colonized .and abundant sspoge 
DroOdvetron .occurred: Chemical analysis of 


surface soil: .pH 6.3; P’ 4 mg/kg soil; exenyeae 
0.07% Caviael, MG bes) “Key 0.26" meq/ LO0g saga: Por 
cultures were established in CEPEC, Itabuna, 
Bahia; the fungus formed a symbiosis with P. 
phaseoloides and S$. halipense producing 
arbuscules and auxiliary cells in roots and 
auxiliary cells and spores in decomposing organic 
matter and mineral soil. 


DISCUSSION 


Gigaspora ramisporophora spores closely resemble 
yellow spores of G. decipiens, however, spores of 


the latter species can also be colorless, pale 
greenish-yellow or light brown at germination. 
Total wall thickness parameters overlap but the 
wall configurations “are distin CG. 
ramisporophora spores have an outer unit wall, a 
laminated middle wall and an inner germinal wall; 
Hall & Abbott (1984) describe a spore wall which 
changes as the spores age; older spores of G. 
decipiens have two.laminated walls (the outer 
wall being replaced by 2-3 laminations) over the 


germinal wall. The refringency of the germinal 
wall differs from the adhering laminated wall in 
both species. Large papillae are more numerous 


and more widely distributed on the germinal wall 
of G. ramisporophora; germ tubes develop in the 
area where the papillae are largest, near the 
sporogenous cell or elsewhere. Germ tubes of G. 
decipiens emerge near the sporogenous cell 
usually from clusters of large papillae. 
Although small papillae may be generalized in 
both spores they are more often widespread on the 
germinal wall of G. ramisporophora and confined 
to the area around the sporogenous cell of G. 
decipiens. The germinal wall of both species 
adheres tightly to the middle laminated wall, 
however, the papillose area can wrinkle when 
Spores are mounted in lactophenol. The 
sporophore of G. ramisporophora may be simple 
with a single sporogenous cell or branched having 


675 


2-3 sporogenous cells. Branched sporophores have 
not been reported for other species in the genus. 


Dr. R.E. Koske (pers. comm.) interprets wall 3 as 
the innermost layer of laminated wall 2. We 
think the germinal wall is a separate wall. This 


wall was first described by Sward (1981a, b, c) 
studying the spore wall structure of G. margarita 
Becker & Hall. The germinal wall (Fig. 4) is 
represented by a depiction of papillae which form 
on the inside prior to germination. 


One or rarely two bulbous cells contiguous with 
an apical sporogenous cell are sometimes present 

on sporophores of G. ramisporophora (Figs.1-D, 
2 I-L). Similar structures have been observed on 
sporophores of Scutellospora pellucida (Nicol. & 
Schenck) Walker & Sanders, S. heterogama (Nicol. 
& Gerd.) Walker & Sanders and an undescribed 
scutellospora species. Nicolson and Gerdemann 
(1968) may have reported the same anomaly when 
they noted that Endogone heterogama (S. 
heterogama) had "usually one, occasionally two, 
bulbous suspensors 21-26 wm diam. with a slender 
hypha extending from the suspensor to the base of 
the spore...". 


Wie iorngs suggest that, thick hyphal 
protrusions, and perhaps the slender hyphae, 
extending from the sporogenous cells of Gigaspora 
and Scutellospora species, once thought to be 
antheridium-like (Gerdemann, 1955) or small 
suspensors (Nicolson and Gerdemann, 1968) may be 
vestigal sporophore branches. A branch 
(Sporogenous cell and connecting hypha) 
emanating from an apical, bulbous sporogenous 
Seamemeias been, a frequent, finding, in. Gc. 
ramisporophora; three observations have been made 
by the senior author of immature spores forming 
on these branches (Fig. 1-A). 


The colonization of decomposing plant residues by 
G. ramisporophora (Fig. 1-B) results in abundant 
spore production. This phenomenon has also been 
observed with some other Gigaspora and 
scutellospora species. Ferrer and Herrera (1980) 
reported finding S. heterogama spores inside seed 


676 


coats and Arvantes and Taber (1985) reported 
abundant Gigaspora spores in weed seeds. 


TERMINOLOGY 


Becker and Hall (1976) use the term "subtending 
hypha" for the connecting hypha and the structure 
variously referred to as a bulbous swelling 
(Gerdemann, 1955), a-~suspensor (Nicolson and 
Gerdemann, 1968), an attachment of a bulbous 
nature interpreted as a suspensor (Old et al., 
1973), a suspensor-like cell (Gerdemann and 
Trappe, 1974) and bulbous and hyphal suspensor- 
like**cells® (Brown and: ‘King y*19820— Prigr Fuo0 
spore formation "subtending hypha" would be a 
misnomer; another term is needed. Ferrer and 
Herrera (1980), describing spores in the genus 
Gigaspora, apparently limit the term "sporophore" 
to the "sporogenous cell", however, it can be 
interpreted broadly (Snell and Dickeweto7. 
Hawksworth et al, 1983) ° and “is /thev cern 
suggested for the spore bearing hypha(e) in the 
Endogonaceae. Tandy (1975) used the term for a 
Glomus species. The term "sporogenous cell" 
(spore-producing cell(s) of the sporophore) 
recognizes the specialized function of the cell. 


ACKNOWLEDGEMENTS 


Special thanks are due Dr. Paulo de Tarso Alvin, 
Dr. Percy Cabala Rosand and Marilete Wolff of 
CEPEC, the Mycorrhiza Project at’ themeencurc 
Internacional de Agricultura Tropical (GIAT on 
Cali, Colombia and Dr. James M. Spain, a very 
willing collaborator. , Dr. Li KY Abpea 
viable spores of G. decipiens from the collection 
in Western Australia and Dr. Jeanne Claessen de 
Miranda prepared the Portuguese translation. We 
would also like to thank Dr. R.E. = Kostemror 
perusing slides, reviewing the manuscript and 
making helpful suggestions. 


j 

: 4; 

| 677 

| LITERATURE CITED 

Arvantes, E.M. & R.A. Taber, 1985. Observations of 

VAM spores inhabiting weed seed in Northeast 
fexas  SOlls. In: Proceedings’ of: the 6th North 
American Conference on Mycorrhizae, R. Molina 
pee) Oregon State University, Corvallis, OR. 


Seeeere WN. & I.R. Hall, 1976. Gigaspora 
margarita, a new species in the Endogonaceae. 
yeootaxon 4:155-160. 

Gee Mer. & E.J. King, 1982. pened shiek and 

| histolo ee vesicular-arbuscular mycorrhizae. 
BieeeMethods and Principals of yoorrhd zal 
Research, N:C. Schenck (Ed.). The American 
Brmovath.) SSOC., ot. Paul) MN. a hoes 

merrer, Piece reer re ras ssl S 0; Exe gGenero 
Gigaspora Gerdemann et Trappe (Endogonaceae) en 
nee REV oie DOU. NACE 145-66. 

Gerdemann, J.W., 1955. Relation of a large soil- 
borne spore to Phytomycetous mycorrhizal 
infections. Mycologia 47:619-632. 

_emeoe tan, JuaWw. & J.M. Trappe, i974. ve 
meaeomacecac ini the Pacific: Northwest. 
Mycouogia Mem. 5:1-76. 

merr, L.R. & L.K. Abbott, 1984. Some Endogonaceae 
from South Western Australia. Trans /2 Brit. 
MueeiresOc. GC3:203—208. 

Mawksworth, D.L., B.C. Sutton & G.C. Ainsworth, 
1983 Ainsworth & Bisby's Dictionary of the 
Fouuncy ds. Seventh edition. Commonwealth 
Mueowodgical Institute, Kew. P. SOse 

Nicolson, T.H. & J.W. Gerdemann, 1968. Mycorrhizal 
Endogone species. Mycologia 60:313-325. 


Old, EM 3G, w.He Nicolson) & J.F.' Redhead, 1973. A 
new species of mycorrhizal Endogone from 
Nigeria with a distinctive spore wall. New 
Peo i2°2S17-823). 

Seer We Hs 6 6& «6UEWA. Dick, 1971. A Glossary of 
Mycology. Revised ed. Harvard Univ. Press, 


Cambridge, MA. 
ard, KReJ., 1981a. The structure of the spores of 
Gigaspora margarita. I. The dormant spore. New 


. Onn ys = : 
sward, ae 1981b. The structure of the spores of 
Gigaspora margarita. II. Changes accompanying 
germination. New Phytol. 88:661-666. 
eet... , 19Sic. The structure of the spores of 


Gigaspora margarita. III. Germ-tube emergence 
ad epeteelt Non PUNASSE BSn.O Os Orit 


Re re was Y ep orocarpic species of 
ndogonaceae in Australia. AGS ee. .Ost 
23:849-66. ; 
meer, C., 1983. Taxonomic concepts, in’ the 


Endogonaceae: spore wall characteristics in 
species descriptions. Mycotaxon 18:443-455. 


ve 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 679-696 January-March 1989 


A NUMERICAL TAXONOMIC STUDY OF SOME PHIALIDIC GENERA 
OF HYPHOMYCETES: CLUSTER ANALYSIS 


Angélica M. Arambarri! and Marta N. Cabello 


feeoreuco, "Spepazzini'™,53 N2 477.1900, La Plata, Argentina. 
SUMMARY 


Cluster analysis by the unweighted pair group method 
using arithmetic averages (UPGMA) was performed on morpho- 
Boercatedava from 28 characters of 114 species of some 
dematiaceous, phialidic genera of Hyphomycetes. 

As a result of this analysis some species were found 
Mervercwosely related isuch as the species of the genera 
Penueuvrapeeja, Gonytrichum and Ménisporopsis, and the 
Bevowivy ore the “species of Dictyochaeta and Menispora, 
Dut Others need a revision namely: Codinaea * apicalis, 
mete ce Ol, Chactosphacria dingleyac, ©. ‘elegantissima, 
bw reemenvosa,® C. glauco-nigra, CC.’ ,intermedia,; C. maha- 
rashtrensis and Dictyochaeta menisporoides, which probably 
need a different treatment. The same occurs with Menispora 
Brivannica and M: uncinata. The genera Chaetopsina, Chae- 
topsis and Kionochaeta probably must be reconsidered. 


INTRODUCTION 


The anamorph genera have always been a great problem 
nouewortrers On ‘taxonomy. This is due to the characters 
used to define them, that are often inadequate, poorly 
defined or purely monothetic. As pointed out by Kendrick 
fice roosmo (1979), the final solution to this problem 
Bitteepessrrivyed at only when we can find the relationships 
with the teleomorphic-fornm. 

Pevacugh the problems involved? are »known vo every 
aveonogier, only .a few attempts: have been made, to find 
Penerat rules for linking the two systems. On the other 
hand the amount of anamorph genera described in the litera- 
fipem@on inereasing day’ by day. ‘This. fact has led’ ‘us to 
Pvoaue  ceonera whose concepts are superposed: and for, this 
reason some species share characters with more than one 
genus. 


1) Researcher of the Consejo Nacional. de Investigaciones 
Cientificas y Técnicas (CONICET), Argentina. 

2) Researcher of the Comisién de Investigaciones Cientifi- 
cas de la Provincia de Buenos Aires (CIC), Argentina. 


680 


Kendrick (1980) considered the generic concepts in 
Hyphomycetes and proposed the amalgamation of several 
genera, because their delimitation was confusing and inade- 
quate. As a first approach DiCosmo et al. (1983) hypothesi- 
zed that there was not enough justification for maintaining 
Cylindrotrichum Bonorden and Chaetopsis Greville as separa- 
te anamorph genera. For this reason they disregarded them. 
Nevertheless, Cabello and Arambarri (1988) revised the 
genus Cylindrotrichum and found out it was valid and diffe- 
rent from Chaetopsis. ¥ 

In this paper we are including 114 form-genera having 
in common macronematous, ‘dematiaceous, unbranched or 
oceasionaly branched ‘conidiophores, with or withouu setae 
that..may be independent structures or become) (he we tens 
or fertile prolongations of conidiophores. “In tnemiaties 
case, the conidiogenous cells are arranged (at Gaeuema 
level of the conidiophore..and may be arise Sessile wor 
otherwise on branches of varing length. In all the species 
considered, here .the ontogenic originy of | URGee Ome 
phialidic with or without percurrent and/or sympodial 
proliferation. The conidia are grouped in slime sneaga. 

Gams and Holubova-Jechova (1976) and DiCosmo et al. 
(1983) diagramed and illustrated the habit of some genera 
considered in the present paper. 

One of the purpose of this paper is to analyse phene- 
tic similarities among the species of the genera considered 
here, and:to establish if “all the species 0] sae 
are clustering together or if they appear mixed Wwithe one 
Species of other genera. -The latter situatlOmeywieee 
analysed in. another , paper with the purpose mgt ge. me 
out the delimitation of the genera that appear contuae 
and inadequate according with the idea of Kendrick 


SPECIES ANALYSED 


Bahusutrabeeja angularis Vasant Rao & de Hoog; 
Bahusutrabeeja dwaya Subram. & Bhat; 


Chaetopsina auburnensis Morgan Jones; 
Chaetopsina catenulata Samuels; 


Chaetopsina wflulval Rambelia.; 
Chaetopsina penicilliata Samuels; 
Chaetopsina polyblastia Samuels; 


Chaetopsina splendida Sutton & Hodges; 
Chaetopsina unilatéralis Kirk; 


Chaetopsis grisea (Ehrenberg) Saccardo; 
Chaetopsis romantica (Rambelli & Lunghini) DiCosmo, Berch 
& Kendrick; 


Chloridium atanagildae Holubovaé-Jechové apud Holubov4-Je- 
chova & Mercado; 

Chloridium botryoideum (Corda) Hughes; 

Chloridium caudigerum (Hohnel ) Hughes; 

Chloridium chlamydosporis (van Beyma) Hughes; 

Chloridium clavaeforme (Preuss) Gams & Holubova-Jechovéa; 


681 


Culeridium codinaeoides Pirozynski; 

Chloridium cubense Holubovaé-Jechovéa4; 

Chloridium cylindrosporum Gams & Holubovdé-Jechovéd; 
Chloridium lignicola (Mangenot) Gams & Holubovaé-Jechov4; 
Chloridium matsushimae Gams & Holubové-Jechova;3 

Chloridium pachytrachelum Gams & Holubovaé-Jechov43 
Chloridium paucisporum Wang & Wilcox; 

Chloridium phaeosporum Gams & Holubova-Jechova3 

Chloridium preussii Gams & Holubovdaé-Jechové; 

Chloridium reniforme Matsushima; 

Gnltorrdaium smithiae Sinclair & Eicker; 

Chloridium transvaalense Morgan Jones; 

GOhloridium virescens (Pers. ex Pers.) Gams & Holubova-Je- 
chova$ 

Cylindrotrichum clavatum Gams apud Gams & Holubova-Jechovéa; 


eviindrotprichum curvatum Morgan Jones; 


Syeenatorri chun ellisii Morgan Jones; 
ey wanouroerrichum, @xcentricum Chirat) TEEUSUbtom I Cabeldio 2 


Arambarri; 
fvaagnurotricnum fasciculatum Mercado Sierra; 
Cylindrotrichum hennebertii Gams & Holubova-Jechovéd; 


Cylindrotrichum oblongisporum Morgan Jones; 

Cylindrotrichum oligospermum Bonorden; 

Gyeindrotrichum probosciophorum (DiCosmo, Berch & Kendrick) 
Arambarri & Cabello; 
Cyveundrotrichum proliferum Matsushima; 

Cylindrotrichum triseptatum Matsushima; 


Cylindrotrichum zygnoellae (Hohnel) Gams & Holubov4é-Jecho- 
va; 


Dictyochaeta abnormis Holubovdé-Jechovad; 
Dictyochaeta -apiculata (Matsushima) Arambarri & Cabello 
comb’. nov. 

Basionym: Codinaea apiculata Matsushima; 
Meee yoocuaets aristata (Maire) Arambarri & Cabello comb. 
nov. 

Peatonvyn: Codinaea aristata Maire}; 
Dee wonectla assamnica (Agnihothrudu) Arambarri;, Cabello 
& Mengascini; 
Dictyochaeta botulispora (Hughes & Kendrterk)” Gabel lor re 
Arambarri comb. nov. 

bastonym: Codinaea botulispora Hughes '& Kendrick; 
Peer oopacta  bDrevisetula (Hughes  & ‘Kendrick) Cabelio & 
Areambarri comb. nov. 

bestouym: Codinaea bDrevisetula Hughes i¢é Kendrick 
Drcryvocnaeta state of “Chaetosphaeria.callimorpha (Mont.’) 
Deco. 
Dictyochaeta clavulata (Holubovd-Jechova) Holubovd-Jechova; 
Dictyochaeta coffeae (Maggi & Persiani) Cabello & Arambarri 
comb. nov. 

pesgenynm:: Codinaea coffeae Maggi &.Pérsianis 
Dictyochaeta cylindrospora (Morgan Jones & Ingram) Aramba- 


Boe Cabello TeomD . lOV. 


bastouym: Codinaeatycylindrospora Morgan” Jones “& In- 


gram; 
Dictyochaeta dimorpha (Toyazaki & Udagawa) Arambarri & 


682 


Gabe liow comb. TONG. 

Basionym: Codinaea dimorpha Toyazaki & Udagawa; 
Dictyochaeta eucalypti (Sutton & Hodges) Cabello & Aramba- 
rrInCOMmDusnov: 


Basionym: Codinaea eucalypti Sutton & Hodges; 
Dictyochaeta fertilis (Hughes & Kendrick) Holubovad-Jechovd; 


Dictyochaeta fuegiana wopegaZzZini $ 
Dictyochaeta gamundii Arambarri & Cabello apud Arambarri, 
Cabello & Mengascini; 
Dictyochaeta heteroderae (Morgan Jones) Cabello & Arambarri 
comb.nov. 1 
Basionym: Codinaea heteroderae Morgan Joness 
Dictyochaeta hughesii — (Ellis) Arambarri & Cabelitcomecen 
nov. 
Basionym: Codinaea hughesii Elis: 
Dictyochaeta illinoensis: (Hewings & Grane), Arambareiee 
Cabello comb. (nev. 
Basionym: 'Codinaea illinoensis Hewings & Crane: 
Dictyochaeta longispora (Hughes & Kendrick) Cabello & 


Arambparri COMD LR nOM. 

Basionym: Codinaea longispora Hughes & Kendrick; 
Dictyochaeta lunata (Matsushima) Arambarri & Cabello comb. 
nov. 

Boston: Codinaea lJunata Matsushima; 

Dictyochaeta lunulospora (Hewings & Crane) Cabello & Aram- 


Dar Law COM .a NOM. 

Basionym: Codinaea lunulospora Hewings & Creer 
Dictyochaeta matsushimae (Hewings & Crane) Arambarri’ & 
Cabello comb. nov. 

Basionym: Codinaea matsushimae Hewings & Crane; 


Dictyochaeta menisporoides Holubova-Jechov4a; 

Dictyochaeta novae-guineensis (Matsushima) Romero; 
Dictyochaeta .obesispora (Hughes & Kendrick) |) Capesiausa 
Arambarrid vcomb'. nov. 

Basionym: Codinaea obesispora Hughes & Kendrick; 
Dictyochaeta parva (Hughes & Kendrick) Arambarri & Cabello 
COMDe UNO. 

Basionym: Codinaea parva Hughes & Kendrick}; 


Dictyochaeta state of Chaétosphaeria pulchrise@pamea eee 
Kendrick 4 Shoemaker; 


Dictyochaeta ;querna Kirk; 
Dictyochaeta septata (Sutton & Hodges) Arambarri & Cabello 
comb. nov. 1 

Basionym: Codinaea’septata, Sutton «& Hodgese 
Dictyochaeta setosa Cig hess & Kendrick) Cabello & Arambarri 
CGOMDE NOV. 

Basionym: Codinaea setosa Hughes & Kendrick; 
Dictyochaeta simplex (Hughes & Kendrick) Holubova4-Jechovd; 
Dictyochaeta state of Chaetosphaeria . talboti@ = iaeneas 
Kendrick & Shoemaker}; 

Dict yochaetaphatakt res Bhat. Miioupeons 

Dictyochaeta triseptata (Matsushima) Castaneda; 
Dictyochaeta unisetula (Morgan Jones & Ingram) Cabello 
& Arambarri comb. nov. 

Basionym: Codinaea unisetula Morgan Jones & Ingram; 
Dictyochaeta vulgaris ( (Hughes & Kendrick) Cabello & Aramba- 


RECON umil Olivine 


mw kendrick. 


Gonytrichum 
Gonytrichum 
Gonytrichum 
Gonytrichum 
~Jechova; 


Gonytrichum 


Kionochaeta 
Kionochaeta 
Kionochaeta 
Kionochaeta 
Kionochaeta 


683 


Basionym: Codinaea vulgaris Hughes 


caesium G.G.& Nees ex Leman; 

chlamydosporium Barron & ohat: 

macrocladum (Sacc.) Hughes; 

mirabile Holubovd4-Jechovad apud Gams & Holubova- 


ypsilosporum Holubov4-Jechové; 


erasteta Kirk apt'd (Kirk: é& sutton} 

svoriensis. (Rambelilzi-& Lunghini) Kirk & Sutton; 
Peniensis /CKirk)V Kirk .&€ sutton} 

malaysiana Kirk apud Kirk & Sutton; 

nanophora Kuthubutheen & Nawawi$ 


Kionochaeta pughii Kuthubutheen & Nawawi; 

Kionochaeta ramifera (Matsushima) Kirk & Sutton; 
Kionochaeta spissa Kirk & Sutton; 

Kionochaeta virtuosa (Rambelli & Lunghini) Kirk & Sutton; 


Menispora britannica (el aS ek Dr kas 
Menispora ciliata Corda3 


Menispora glauca Pers. 


Menispora manitobaensis Sutton}; 
Menispora state of Chaetosphaeria pulviscula (Curr.) Booth; 
Menispora tortuosa Corda; 

Menispora uncinata Hughes & Kendrick; 

ludoviciana (Grane \f&> <Schoknécht). “Kirk 


Menisporopsis & 


Sutton; 
Menisporopsis novae-zelandiae Hughes & Kendrick; 


Menisporopsis pleiosetosa Rao & de Hoog; 
Menisporopsis profusa Pirozynski & Hodges; 
Menisporopsis theobromae Hughes; 


Zakatoshia hirschiopori Sutton; 


Zanclospora austroamericana Sutton & Hodges; 


Zanclospora brevispora Hughes & Kendrick; 
Zanclospora novae-~zelandiae Hughes & Kendrick. 


**Codinaea 
**Codinaea 
**Codinaea 
SP Ono. Tt 3 

**Codinaea 
& Onofri; 

**Codinaea 
**Codinaea 
a Onofri; 

**Codinaea 


epicaias! Berk té (TNE S 
state of Chaetosphaeria dingleyae; 
elegantissima Lunghini apud Lunghini, Rambelli 


filamentosa | Onofri apud. Lunghint, Rambelli 


glauco-nigra Cooke & Ellis 


intermedia “Rambelli apud > Lunghini» Rambelli 


maharashtrensis Pirozynski & Patil. 


**These species are not recombinated because 
be reubicated in alatter paper. 


they will 


OBSERVATIONS: 
epecies: 


considered 
Castaneda; 


We have not 
Chaetopsis cubensis 


the following 
Cylindrotrichum 


684 


gorii Lunghini; Codinaea pakhalensis S.M. & S.S. Reddy; 
Codinaea australensis Sutton; Codinaea tortuosa Sutton; 
Menisporopsis pirozynskii Varghese & Rao; Gonytrichum 
indicum Sharma & Munjal and Zanclospora mystica ZdZucconi 
& Rambelli; since the original descriptions on), them type 
material were not available. On the other hand Cylindrotri- 
chum helisciforme Marvanova, as DiCosmo et al. 1983) 
have suggested, could be a species of  Heliscus eaaces, 
because its characters do not match the genus Cylindrotri-= 
chum. 


MATERIALS AND METHODS 


The two species of the genus Bahusutrabeeja Subran. 
& Bhat; 7 species of Chaetopsina Rambelli;,2 species sor 
Chaetopsis Greville; 18 species of  Chloridiuieiin ea, 
7 species’ of Godinaea Maire; 12 species of Cylinarovrienu: 
Bonorden; 36 spécies of Dictyochacta Spegazzinis especies 
of Gonytrichum.C. J. & Fs Nees ex Leman: ) Ouse eee 
Kionochaeta Kirk & Sutton; 7 species of, Menispora@rereogn, 
5 species of Menisporopsis Hughes; one species "of aZakacco- 
shia Sutton and 3 species of Zanclospora Hughes Rengraen. 
constitute the 114 Operational . TaxonomicaljyUniGcemeue 
(Sneath & Sokal, 1973). 

The characters. used have been obtained after studying 
ori vpinalwasescriptcLons ol thetepeciess 

The data consisted*‘of 28 characters scored eionmeeacn 
of the 114 OTU. All the characters were qualitative wee 
of the. characters were scored as double state dateawepreccn= 
ce (+t),. absence (0). The double state data, exeliudang 
state, were scored with (1) or (2), indistinctly pee a 
characters such as: conidiogenous cells sessile. on the 
conidiophore and conidiogenous cells growing "onepraneneas 
we found, that .some species ineluded both statea aaa 
clear that the best ‘scoring (to cover the  p0eeqg eee 
could be as follows: .conidiogenous’ cells Seseigeae ee 
conidiophore ae conidiogenous cells ‘on branches ee 
and, conidiogenous cells sessile or/on branches (125). 
A similar situation -oectirs with “characters #1) ue 
ete 

The characters were scored for each-O1U age) aie 
1) Conidiophores: mononematous (Fig. 1B & C) (1); synnema- 
Tous iie . heehee & 
2) Conidiophores synnematous with central setae :present 
(Rigen el) CW @ormebeenint 0: a 
3) Conidiophore branched: present (1) or absent (O)% 
4) Conidiogenous cells growing on the conidiophore (not 
as a cells of the conidiophore ): present (Fig. 1 D-E-I-J-K 
& Li) horkabesecnany (On. 
5) Conidiogenous cells terminal (Fig. 1. G-H) (1), terminal 
and intercalary (Fig. 1°F) (2), terminal on) brancheesnaes 
6)Conidiophore extending far from the conidiogenous cells: 
present (Fig. 1 D-I-K &°L) (1), or wabsent (fig saemeee 
SF TCO 
7) Extension of (‘the conidiophore fertile: presemuman 
orpapsents (0); 


) Extension of the conidiophore: straight (1) or flexuose 
2} e 

) Conidiophore without setae (0), with several setae 
Fig. 1 Bee C)° (1), with only *one’'seta (Pig. cae 

@ 


if 
) Fertile setae: present (Fig. 1 B) (1) or absent (Fig 
GPC. 
) 
) 
) 


NS ae es te St ear se. 


1 Setae: Straight (Fig. 1 B) (1) ,flexuosev Bipwa es 

mike 

RQ Conidiogenous cells on the conidiophore ( as cells 

f the conidiophiore) only at the apex: present (Fig. 
G & H) (1), or absent: (0). 

3) Conidiogenous cells sessile on the conidiophore (Fig. 
K) (1), on branches: (Fig. 1 D-E-I & J)(2), sessile or/on 


branches: (Fie. 1 U1 eS). 

14) Conidiogenous cells on branches ( not collar hyphae): 

apical dispossed. (Fig. 1 ‘J) (1), ain thes migeipareees 
1 Deol) {203 .epicalvand tn the midi part ome 

15) Conidiophore bearing short curved lateral branches 
(collar hyphae): present (Fig. 1 E) (1) or absent (0). 

16) Conidiogenous cells unilaterally arranged (Fig. 1 
1)(1))° anphigenous (verticillate) (Fig. 1 7° D=Ri ekg 

and unilaterally and anphigenous (1.5) 

17) Conidiogenous cells without proliferation (0), with 
sympodial proliferation (Fig. 1G) (1), pereurrent prolife- 
ration (Fig. 1 H) (2) and sympodial and percurrent prolife- 
PavUonr yl a). 

18) Proliferations sympodial and/or percurrent scanty 
(1) or many) (2). 

19) Conidiogenous cells straight (1) or strongly curved 
CPA g ders a 2) 

20) Conidiogenous cells with conspicuous collaretve: 

present (1) or absent (0). 

21) Conidiogenous loci terminal (Fig. 1 °H) (99 emene 

andvintercadary =(Fie., dau ee.) 

an Conidiogenous cells lageniform: present (1) or absent 
O). 

23) Conidia O-septate (1), septate (2). 

24). Conidia with setulae: present (1) or absent (0)% 

25) Setulae as a prolongation of the wall of | thewean use 
(1), mucous setulae (2). 

26) Conidia straight: present (1) tor’ absent (0 
27) Conidia symmetric: present (1) or absent ( 
28) Ratio L/W 1:1: present (1)-or absent (0). 
Data processing: a)’ A Basic Data Matrix (B.DJMWjmepe ee 
1) of 28 characters by 114 OTU was developed.) ie Sa ire 
no comparable (NC), an characters’ 2, 7,’ 8) (O@R em 
14, 155.16, 18, 25 are-a~product ‘of the kindof Meares 
tion. 

The computer work was done on a IBM 4361 using the 
NT-SYS programs developed by Rohlf et al. (1971). It was 
carried -out at the Centro de Estudios Superioreagwpare 
el Procesamiento de la Informacién (CESPI) of the Universi- 
EY Ot Wai Pla pal. 

b) Achievement of Similarity Matrix using the Average 
Euclidean Distance (Taxonomic Distance). 
ec) Cluster analysis: this analysis uses techniques) that 


) 
QO). 


TABLE 1. BASIC DATA MATRIX 


Characters 


Species 


Fe Dod es os < fet = OMS eh, 1 12.13 it eh Old 1G 1: 20 alee eo 2h 2) eo AT ee 


1 


1 
1 


4 
cal 


2 NC 
2 NC 


1 
1 


oa, te 
OE Ce 
1 


1 
1 


1 
1 
1 
1 
1 
1 


1 NC NC NC NC 1.5 1 


1 NC 0010 NC NC O NC NC 
1 NC 1010 NC NC O NC NC 


Bahusutrabeeja angularis 
Bahusutrabeeja dwaya 


NC NC NC NC 1.5 1 


1 


0 


1 
1 
1 


1 ONC O 
0 


@) 
1 
1 
1 
1 
1 
1 
1 
1 


Tm NG ONG Ogee ete ar gO NG 


0 


1 SNE OST 3-71 


Chaetopsina auburnensis 


1 
1 
1 
1 
1 


OT 2a) NG 


CF ee 0 NG 1 


Te Ne NGS Oat ee ke 2) 


1 


TONG YT 1-30 NG NGO NC NC =O" 2 


TeNGlOe Loe) 


Chaetopsina catenulata 


Chaetopsina fulva 


O NC 
O NC 
O NC 
O NC 


1 
1 
1 
1 
1 


1 
1 
1 
1 
1 


nr U2 OANG 


0 


0 


Oe = NG 
i Pv eS 


1 
1 


1ONG Us i320 NCONG “ONG NC 20 = 2 
TNC | Sie 3 ONG NG-OUNG NC 2 Oc 2 


Chaetopsina penicilliata 
Chaetopsina polyblastia 


Chaetopsina splendida 


1 
1 


O NG 0. 0,8 


1 


1 
1 
1 
1 


fe NCO NG ae): te eer we as Ore 


0 

0 

0 

1 
1 NC 001 0NC NC O NC NC 
i NC-OS0= 120° NG NC O“NC NG 
4 NGO 20s 120 NG NGO ONG= NC 
1 NC 001 0NC NC O NC NC 
1 NC 0010 NC NC O NC NC 
1 NC 001 0NC NC O NC NC 
1 NC 0010 NC NC O NC NC 


A NCeO -O4150 NC NG 4 


ta Geote loud 


TNC SO = ONE 
TO NG ONG 02s ae eC eg Ot 


LO NG-NG 0 


to NGA le jad 


Chaetopsina unilateralis 


Chaetopsis grisea 


1 
1 


1 
1 
0 UNG 05 40.2 O 


1 


1 NGOS Lael 


O NC 


1 
1 
1 
d 
1 
1 
1 
1 
1 
1 


lO NOE NGO ee (2-6) 1 5. ONG 1 


A NGHOR 13s 


Chaetopsis romantica 


0 


Z 


1 


1 
1 


1 NC NC NC NC 
1 NC NC NC NC 


ildae 


Chloridium botryoideum 


8 
Chloridium caudigerum 


Ghloridium atana 


1 


1 
1 
1 
1 
1 
1 


2 


1 
1 
1 


@) 


1 
1 


1 
1 


1 NC NC NC NC O NC 
1 NC NC NC NC O NC 
1 NC NC NC NC 2 
1 NC NC NC NC 2 


Chloridium chlamydosporis 


Chloridium clavaeforme 


1 
1 


0 


Chloridium codinaeoides 


Chloridium cubense 


1 
1 
1 


1 


cimey® 


1 
1 


1 NC NC NC NC 1.5 1 


1 


1 NC NC NC NC 1.5 1 


1 


1 


Chloridium cylindrosporum 


Chloridium lignicola 


1 


1 
1 
1 
1 


1 NC NC NC NC 2 
1 NC NC NC NC 2 


1 NC 0010NC NC O NC NC 
1 NC 0010 NC NC O NC NC 
1 NC 001 0NC NC ONC NC 
1 NC 0010 NC NC O NC NC 
1 NGS) 202 t7O NE NCeO" NG NG 
1 NC 0010 NC NC O NC NC 
1 NC 0010NC NC O NC NC 
1 NC 001 0NC NC O NC NC 
1 NC O01 0NC NC O NC NC 
je NG O07 O2t2U°NG NCLOONG: NG 


Chloridium matsushimae 


1 


0: NCQ OL =) 


1 


1 
1 
1 
1 
1 
1 
1 
1 


R 


1 
1 


te NGoNG NG NG 10574 


1 NC NC NC NC 
1 NC NC NC NC 


Chloridium pachytrachelum 
Chloridium paucisporum 
Chloridium phaeosporum 


Chloridium preussii 


t n4 
eey 
je cl 
: a 
1 
1 
Go a So NG seus a! 


1 
1 


0 


1 


1 


1 NC NC NC NC 2 
1 NC NC NC NC O NC 
1 NC NC NC NC 1.52 


O 
‘ 
1 


1 
1 
1 


Chloridium reniforme 


O NC 
QO NC 


0 


1 


2 


Chloridium smithiae 


1 NC NC NC NC ONC 


Chloridium transvaalense 


Chloridium virescens 
Codinaea apicalis 


1 


1 NC NC NC NC 2 
2-0 NG NGS O21 52a 0-1-5550 NG 


1 


1 
1 
1 


1 
1 
1 


1 


Ga bens se ea 


1 


1 


Get. 5. 2 Oe es oe 


ta NGO NG. OO re ee 


1 


@) 


1 


@) 


1 


Na el 


Codinaea st. C. dingleyae 
Codinaea elegantissima 


O NC 


1 


Pe NOaT Si-33 <4 


688 


OEY aU CON Oe ake. Ee a BS ON 
a ele eet ee Ad eee be abe ek IN 
ied Pa A eae) oo OR eg ae ak ep iy SS 
ih. ee be tg obs Uelou te ta Gy LON 
ig tee Gee Dor ON te oe be DING a wen 
eo Velveeta, Ton me rN 
OL FUE Use hokey ot gah oN OE aN 
el eel eek ee) ae, wey aoc to ON. 
CSN eee UR re Oe GN 
OO, Oe IN eG i eels Than be SON 
eer ee mC ee beg Oh La IN 
eee rN cee to rer ee Set a GON 
ee ey ee ee ee ba OE ee ON 
es BP cata ey) epi FL rb Le ON 
Oe rer mth cn aCe bade Ost ON 
Po C2) 6 na i 0 eo ee See ee a 
A tn eee Clee: 2G) Peer ON 
Cire IN Ur ce Ol tt LON “Os ON 
Ua we rere Ne hee G a bes (LNG Ge IM. 
et a ee Cor rca eel dela LON 
ae ee mC mre) a eL ah Sr To ON 
Oe age a) Se es O ac een ba te CaN 
et, sem ee ae) = ble 2 le ON 
GO eee aN A) a Gh ame et = ONO ON 
Cee ar ate bee Pe oe cae ON 
ee am eG el Cie ie CN 
Die ee ONO eC a OQ cor baw Sl» eh ON 
ee eee aol Cee 0m Les oc | ON 
Oe er UN Sir, GR 1 bes 

We ee boee ieel) ore ede Lo ONG 

OP UieeO tN Gae) “tie | ot NaS al 
a elem) ete be) eaten fee La ON One C 

CeCe Ome, curl awe Ol 8 lal le OP 


ON 


ON 


ON 


bt 


ON 


ON 
ON 
ON 
ON 
OL 


Ox: OO '@ O OOOO OO OO OOO Cy, CO - — Oo — © 6 oleae 


ON 


ON 


PES ENDING) 


SO at eh el et SOO OS SNS OMe tO ete ia ele eieKee Gis GS GOO Oo 


a 


Wis = 3 O10 OOO OO @0:O 10 0 0 COC ©: 0 OC OC.C SOloretetserere) 
CVs y- s— 3 OO OOO © OOOO © OC: OC1O | C'@, OOOO OC Oe eee ene 


oO OB 0.0 OV ee ee ee el eee eee ee vee eel ee ee ee ee el ed ee oe oe EO 0 


HTavi 


SN i NN NN a 


eelepoisez,sey 
Ttpunues 

euetsony 

STL T4495 

tydATeone 

eydioutp 
erodsoaputT fo 

seat joo 

eSYLeVTNASTO 
eydiout[ [eo °*9 °48 
e[N7esTAsasq 
ezodst[nz0q 
BeoOTUesSse 

ey eystse 
eyeTnotde 
sTu.zouqe 


ey yoeeyookyoTG 
ey yeeyoosyoTG 
eyeeyookyoTq 
eyoeeyoosy0TG 
eyoeyoosy0TG 
eyeeyoos 0TG 
eyoeyoosyoTG 
eyoeeyoosyoTg 
eyeeyookyoTG 
eyoeyookyoTg 
eyoeyoosyoTG 
eyoeyooskqyoTg 
eyoeyoosyoTg 
eyeeyoosqyoTq 
eyoeyoosyoTg 
eyosyookyoTg 


SVT [SOUusTZ 
unyeydestz4 
unteyttoard 
unzoydotdssoqoid 
unuztedsostTo 
unztodstsuoTqo 
TT4.19q suuSYy 
uUNPE[TNOTOSey 
wnoTtryUeoxSE 
TEST[Te 
unz}eAINo 
UN}BABTO 


STSUSTVUSeIeyeU 
eTpoeuteyUut 
eistu-ooneTa 
esoyUusUelT ty 


unyotr,OIput TA) 
unyoTry,oIputT Ay 
unyotszyorputTAg 
wnyotazyorputtA9 
unyotrzorputpTAy9 
unyoTryoarputTAg 
wnyotrzOIputtAy 
unyotsrzoOrputtAg 
wnyotrzyzoarputpt Ag 
wnyots,orputTég 
UNYOTIZOIPUTTAY 
unyotryOIputTTA) 
eOeUT pO) 
BeeUTpo) 
BEBUTPOD 
BO9eUTpoO) 
setoeds 
S19eqyoereyg 


CON acess 


TABLE 1. 


Characters 


Species 


2 wo SiGe Ss Oe Gah OeII sts yn lo Oe fe | Glo Une) keer ee eo eee. 2o 


TONG: O-O S120: NO ONG? 1 


1 


Ga 2 Ue er NOM fo) 


1 
1 


1 NC NC NC NC 1.5 1 


1 


O 


Dictyochaeta hughesii 


1 
1 


1 


1 


1 NC NC NC NC O NC 


1 NC NC NC NC 


1 NC 0010NC NC ONC NC 
1 NC 0010NC NC O NC NC 
1 NC 0010 NC NC O NC NC 
1 NC 001 0NC NC O NC NG 


1 NC 0010 NC NC 1 


Dictyochaeta illinoensis 


Chg R be eae, 


; 


1 


1 


Dictyochaeta longispora 
Dictyochaeta lunata 


1 
1 


1 1 
1 
BUR 


1 


1 NC NC NC NC 1.5 1 


1 NC NC NC NC 
1 NC NC NC NC 


0 


2 


1 
1 
1 
1 


1 
1 
1 
1 


Dictyochaeta lunulospora 


1 


1 
1 
1 


1 
1 
1 
1 


1 
1 
1 
1 


1 


0 


Dictyochaeta matsushimae 


a hs OS NGRNG LOGI 2 One 
1T-NG2O O41 (0 NG{NC 1 


tat eit Sel 


Dictyochaeta menisporoides 


pe WO ie is =O 


e 


1 NC NC NC NC 


1 


1 


Dictyochaeta novae-guineensis 
Dictyochaeta obesispora 
Dictyochaeta parva 


1 NC NC NC NC 1.5 1 


2 0: 010. {eo NGe NC OoNG ONG 
t°NG O20 (R-OSNCANG TOANCANG 


1 


1 
1 
1 
if 


1 NC NC NC NC 


1 
1 
1 
1 


1 NC NC NC NC 
1 NC NC NC NC 
1 NC NC NC NC 
1 NC NC NC NC 


1 
1 
1 
1 


0 
1 
1 


Dictyochaeta st. C. pulchriseta 1 NC 0010 NC NC 1 


;. SORNGs F035 40 a0) 
a EUR 


a 


12 NG<0 40-1) -O-NCENG 4 
1aNG*0: 501-0 NCANC =] 
TENCSO TO FO NG ONG 


Dictyochaeta querna 


Oss" =O 


1 


1 
1 
1 


1 
1 
1 


Dictyochaeta septata 


0 


Oe SOPNGe eg 


Pa 
2 


1 


1 NC NC NC NC 1.5 1 


0 


Dictyochaeta setosa 


TENG O OQ 1 YO NCENG*OANCANG 
TANG-O20 51 -OSNCENG ONC NG 
1° NC-0°O-1*O°NC-NC-O. NC NC 
1 NC 001 0NC NC ONC NC 
1 NC 0010 NC NC O NC NC 
1 NC 001 0NC NC O NC NG 


Dictyochaeta simplex 


ONGe ROS SO TEU 


1 


1 
1 


1 

1 

1 

1 

1 
2 ONC 
2 ONC 
2 ONC 
CEOENG 
2 ONC 


1 


1 NC NC NC NC 
A OSNOG ENG: Ole mee ma eee TOC, 


1 NC NC NC NC 
1 NC NC NC NC 
1 NC NC NC NC 
1 NC NC NC NC 


Dictyochaeta st. C. talbotii 
Dictyochaeta tilikfrei 


If SOshOS SU 


1 
1 


OL Oreo 


1 


Fa AAS os 


1 
1 
1 


1 
1 
1 
1 
1 
1 
1 
1 
1 


1 
1 
1 


Dictyochaeta triseptata 


1 
1 
1 
1 


Dictyochaeta unisetula 
Dictyochaeta vulgaris 
Gonytrichum caesium 


Oe O> 0 
1 
1 
1 


1 


1 


par ie 


1 
1 
1 


1 
1 
1 
1 
1 


Pe POSNC ONG? BO eee ee HX) 


INGA Sens ae ONC ONG - tCae 2 NC 


ANOS) #1 Se | 


To] O=NCENG 80) 2) NC 


1 
1 


Gonytrichum chlamydosporium 


Gonytrichum macrocladum 
Gonytrichum mirabile 


1 
1 
1 


pL UENG NG Os =2 NC 


leNG Sines 8G 072-0 ENOSNG 80" fee NG 
1 NC+0 443-0 =O. -2-O=NCANG. “O82. NC 


teNG Omi 83 ©O 
TENG SO S133? I 


ied 
ONC NC 1 
| 


O NC 


1 
1 


@) 


1 


OE me 8) 


Gonytrichum ypsilosporum 


Kionochaeta aristata 


1 


1 


O NC 


@) 


1 


ie 
02-0780 
OMCs AO ato 


O NC 


0 am | 


0) 
@) 


1 
1 
1 


LENCSO <1 2374 


Kionochaeta ivoriensis 


1 


1 


120 eNCENG “09 B2= Se Oe tz. 0) SNC 


0 
1 
1 


ANG lee P23: = 


Kionochaeta keniensis 


ARO ANCSNC 60s ees 2) ee OENG 


1 
1 
1 
1 


1 
1 


VONGaO ah 3:24 


Kionochaeta malaysiana 
Kionochaeta nanophora 
Kionochaeta pughii 


OSNG OO 203" 0 


2 
Cee 
0 


2 


0 


(OS CaN Gr Omer St o> sme 


1 


t=) aNC SNC ae 22 


TONG SO Sia! 


1 ONG OS ra) 


1 


1 
1 


1 


1 


1 


eG ral shore 


Oe NGr*-O. ka) 


i OUNCSNOG 2 Osete. Ae nee ONG 


0 


Ae NiGess teeth as ol 


Kionochaeta ramifera 


690 


CON iierere c 


TABLE 1. 


Characters 


Species 


a Se ht 6 PS BO OA 2a 321 1617-1 B19 t 2Uee eee seo eo Gee en 


Te NGe OF s= Se1 


1 


(SG NGte Gs Oo 


1 
1 


0 


1 


1 


i. O* NGaNCs COL Sea 22 = 2 08 NG 


12 02 NEANCS Ors ece Oe 0 


1 


0 


1] NGO? 01> O= NGSNG™ 1 


teNG-Osiea3 4 


Kionochaeta spissa 


AN Oe eerie) 


1 


1 


O NC 
1 


; 


TENG. Oe 32-7 


Kionochaeta virtuosa 


1 


1 NC NC NC NC 


8 ee 
Os 2 eNO NG Oi 2 Oe) 


Menispora britannica 
Menispora ciliata 
Menispora glauca 


0 
0 


: 
1 
1 
1 


ONC 2 O 


1 


2 


Oia 27 OANG NOs Uee2s <2eOe 1. 5 Cents ooecO 


i Noe OF 34 


0 


1 


Ore ee Nowe 
Oo NG OF 30 


@) 


OR e80: NO=NGa Oe se ea. be OE NG a2 0 


On 2nU NG NOs Ores e250) 


TENG CORT S4 


Menispora manitobaensis 


1 


Om NG ome) 


1 


eNC- 04-351 


Menispora st. C. pulviscula 


Menispora tortuosa 


@) 


Cate NG- NG Oa 2 OS 0 CNG aa 


1eNe- 0°32 


1 


1 
1 


eee see 
1 
1 


1 


OG 2e— ING NCANGENGae 


12 NGO el =GANC NG] 


Menispora uncinata 


1 


1 


O NC 


1 


1 


Pod oa os 


1 
1 


1 NC NC NC NC O NC 


1 NC NC NC NC 


1 
1 
1 
1 
1 


1 NC O020NC NC ONC NC ONC NC NC NC ONC 


1 


1000-1 =O NG-NGe2 

teOFO PO NG=NC< 22 =0 
Te OsGais0 NC: NGacee 
41> OA D212 OeNC- NG 22—0 
1 O~ UST SOSNCeNGues =O 


2 
2 
Z 
2 
R 


Menisporopsis ludoviciana 


M 


CaO 


1 


1 
1 


1 


-zelandiae 


leiosetosa 


enisporopsis novae 


1 NC NC NC NC 2 
1 NC NC NC NC 


. 


isporopsis p 


. 
. 


Men 
Men 


Zz 


1 


fusa 


isporopsis pro 


1 


1 
1 
1 
1 
1 


1 NC NC NC NC O NC 


Menisporopsis theobromae 
Zakatoshia hirschiopori 


1 


TOS NOP ee] 
1* 202 NC--Oe5 O20 
O< NG Ue Ce a0 


1 
1 


NOG 


1 
1 
1 


1 
1 


pe Nive Uae SONG 
TaNGsn0 —-2s202NC 
Ve NGO S222 05NC 


TOF NG-NCe GC 


1 NC=06 723 -0a-0 


Zanclospora austroamericana 


Zanclospora brevispora 


re ebeOr NGo NG £0 


1° NGeOs 550 
Te NG- Os he O40 


0 


1 


ONC O 


; 


0 


1 O0NC NC O 


Zanclospora novae-zelandiae 


691 


make up groups of OTU related by their grade of similarity. 

A phenogram of 114 OTU was obtained by the aplication 
of the unweighted pair group method. In this method the 
cophenetic correlation coefficient (r) was computed as 
a measurement of distortion (Sokal & Rohlf, 1962). 


RESULTS AND DISCUSSION 


Becune. 2 shows the 114 OTU- phenogram. The similarity 
values are shown in the upper part. 

iM unis phenogram there. are two. groups: “the speciés 
Merrercomtaiorpenous, cells as part of - the conidiophore are 
Peete ine sone | first group." This’ group <i6) made (up ‘of '2 
Species, Of Bahusutrabeeja; 18 species of Chloridium, 12 
Speges) OL *Cylindrotrichum,, 2 -species (of \Menispora,) 5 
Species, Of Mernisporopsis, 35 species of Dictyochaeta and 
the monotypic genus Zakatoshia. 

im stne second group jwe ican see the species with coni- 
diogenous cells growing laterally or anphigenous on the 
bomiu@opnore «that may be sessile or on) branches. In this 
Pecwewer con tind the 7 species) of Chaetopsina, 2 species 
previaevepsis, S species of Codinaea, 5 species; of Gonytri- 
Pie especies: of Kionochaeta;, 5 species’ of. Menispora 
and 3 species of Zanclospora. 

Peciewstirst }cluster (twowispecies of  Bahusutrabeec ja 
and. some species of Chloridium and Cylindrotrichum appear 
closely related. 

ingeune characters we have gone. through in this ana- 
ieee Olly ‘daiferences are given by’ the: shape and 
the ratio length/width Ot sone contvdne ) iin eGy lLindrotricaun 
fPaeeecentoia are cylindrical and septate, the ratio L/W 
omer eer etnan- |, instead of the species of.) the’ genus 
Chloridium where the conidia are O-septate and the ratio 
L/W is near 1. The remaining characters, such as conidio- 
Pioteceend) .conidiogenous cells, -are so Similar, between 
Mietediabe unere “is a correlation of the species of both 
genera. Bahusutrabeeja angularis and B. dwaya appear in 
thisereroup- because’ their “conidiophores and ‘conidiogenous 
cells are similar to those of Chloridium-and Gylindrotri- 
Smuneetitering from them only by the (conidial morphology. 
Theecne “senus. Bahusutrabeeja the conidia are rounded cubi- 
Poy i One ) of \'the*/ angles being ‘the basal. sscar, and 
each corner provided with a thin hyaline appendage. 

Piss OL ecLyocnaeta ‘cluster ws related by “the presence 
of the conidiophores with sympodial and/or percurrent 
proliferation, conspicuous collarette, eyvyisniacircal or 
ellipsoidal, curved and asymmetric conidia. In this same 
group we can also find Chloridium atanagildae, C. paucispo- 
rum, Cylindrotrichum curvatum, Menispora britannica and 
Meeeuncinalayrrelated'’ to then.’ These presence of the above 
mentioned species is due to the characters which they 
show that relates them more to Dictyochaeta than to their 
own genera. 

According to the author of the species (Holubova-Je- 
chova apud Holubova-Jechova & Mercado, 1984), Chloridium 
atanagildae is distinguised from the rest of the Chloridium 


ep[nqastun: 


xaTdcurts 

Barer 

ezodsopnunt 
eICGST[TN}0q 

ejeunt 

eqernorde 
eJestayotnd *9 *4s 
aewtysnsj ew 
BTNJaestaaiqg 

esoqas 


eZaeyIok TG 
BJaeYyIOAIITG 
eQVaeeyookqITg 
BJaeYIOARITG 
eQaeyIoARITG 
BJaeYyIOARITG 
eJaeyIoARITITG 
eqaeydoAqoTG 
eJeeyIoARITG 
eJaeysokqatg 
B}yaeYIOARITG 


WNZEAINI wnydTIJOApuTTAyg 


eqyeqdastaq 
eqejdes 
STSusautns8-aeaou 
eriodstsaqo 
stTiesq—na 
FoIFATTT? 
eiodst3uoT 
aeayjoo 

STTTI19F5 
Ttpunwes 
eotuesse 
eqeqstie 
Trseysny 
eqyerTnaepto 

euranb 

euetsony 
eydiomr{T Tes °-5) -IAs 
stwiouqe 


BeJaeyIoARITG 
eJaIeYIOARITG 
eJzaeyIoAjITG 
eJoeyookq TG 
eJaeyIoAQITG 
eJVaeyookqITG 
BJaeyIoARITG 
eJVeeyIohRITG 
eJeeysoAjITG 
eJaeyookqITGg 
eJaeyooAqITG 
eJaeyIoARAITG 
eJeeyso0k{qITG 
eJoeysokq TG 
BJaeYyIOARITG 
eJZaeydoARITG 
eJaeYyIoARITG 
eJaeyookqITG 


wniodstoned wntptas0py9 


were not 
identified 
=Codinaea 


( 


polyphialides 
considered that 


(1974) 


et: weal 
(1983) 


proliferation, 


eeTaporsqzay eqaeyIokAITG 


692 


TTIOGTeI *D) *4S BJaRYIOA4AITG 
eeprTrseueje wntpraso0ptyy) 


wndtTiquaoxe 
wnioudotssoqoid 
wniodst8uoTqo 
TTJiIaqauusey 
uniasytptoid 
unuwiadsostto 
FESTITS 
wN}ZeAeTO 


wnyotizyorpurytiAy 
unyotTr,orputytAg 
wnydItTiorTputrpAy 
unyotiqzorpuryzag 
wnydtizOIputytAg 
wnyItizoIpurypAg 
wunyodtizyoriputrypAg 
wnyoItizorpurpiAy 


wuniodsoseyd wntpraoptyg 


SeTTeouskz 


wnyotizOaputtzag 


Saptoseurpod wnrpraopty) 
wniodsoiputytAds wntptiopy9 
eewrtysnsjzew wntpraiopTyg 
BTOOTUSTT wnNtpri0cTyg 
SusoSeiTA WNTptIOTYD 
esuaqnd wNtTptsz0Tyg 
Ttssnaid wntptiz0pTy) 
SWUIOFOPARTI wNTpTIOTY) 
eeTyyTus wntprsz0oTy) 
wnToeyserzAyoed wntptr0Ty9 
efemp efaoqerqnsnyeg 
stiep~nsue efsaqeriqnsnyeg 


© 
° 
) 


“NLO 7LL 40 WVYDONHHd 


£20) ORS os} wey i & & 
‘a. Ol So )-@ Pil> oO ®O 
= aes OW+r a, = rt = ~P 
“d rr Yn oO 2) 
Dl 1 ® o a]Qa0 « 
re) i) Ais @ 12) 4 
* OF Gs =| Olq, O @ 
ST's a oS 5 
ojaa © 
“d (e) in. Ss 
gene bs ee a Ss 
BS nd Ukedy SF ® ,2 
Oe) 45 o a 5 2 P oP) 
oO el Es (.) GS _ Sees = 
ra} M ocd ase 
ape Ome) rca 
3 ph.q4 2 ow Cog 
a ota PAO 
BS oe a 
Cl eee et gece at 
(2 iS OD oO 
Ome AGH 3 
Ss Og p 4.4] oO ® 
Sle mca al seer a 
SAlg o Fqdn Oo GO 
S) “dy re ee 
og CO ag CHISS oo 
Ke On =O] GS 
Oe oe Olea 
OOO Ss ef aati Ol 
dq “a Qe ed[ad 6 
Oe etetoe a ames! S| tae 
e 44 Ps o's 45] | = 
Smee: A ope aes 
B64 o. G4 OOS eat 
O36 Sao oa. 
= 6 @ CO} S\c —) 
tay C= = oir Ne os 
oO ri GH oI/4 = 8 2 
soto © Od} GTS) 
Ope > det Al Ol a ts 
4 & Oo 4 oO 
es Ole) OMe | ree 
®o Te ite Goa ow Ps “d 
“d Ora Ora c= 
Onn te ON et ere 
Ore eerie Ou Hd dG 
Ow ewe Se ated > 
=) 2) DO QO 2,4 


Wilcox 
and«erthey tentatively 


Chloridium. 
Of mebha's 
curvatum is also found in Chis ermoms 
sympodial 
DiCosmo et al. 
species. 


has 
it would be necessary to propose a new eenue 


genus 
sure 
dats 
and 
"intermediate!" 


Cyl Pndrourrehum 


because 


and “curmed conidia. 
curvatum is reminiscent of Dictyochaeta Speg. 


Liles VepeGie sot ron Lord aan 


completely 


of the 
Lor y2uCchH 


C. 


Maire 


693 


untziodsopAwe[Tyos wnyostaqzAuoy 
aTtqeiatw wnyotaqAuoy 
wniodsottsdA wnydt1izfu0y 
unpezTIOIIeW WHYyITIAAUODN 
untseseds wnyotaqyAuo0y 

easti3s stsdoqjaeyy) 
erqsertqttod eutsdojoeyy 
eJe[Nnusjed eutsdoqoeyy) 
e[nostatnd *9 *4s eirodstuay 
Stsuoeqgoqjtuew eiodstuop 
esonji0q ePr0dstuop 

eoneTs eiodstuoq 

BJeTTLIO eirodstusp 

Ttysnd eqjoey .ouory 
erioydoueu ejJaeyo.ouoTy 
septoiodstuew eqaeydsoAqITG 
BipewiajurT eaeutpog 
BWISSTJURSaTO eBaeUTpO) 
eBSOjJUsWeTTJ eaeuTpo) 
aekeTsutp *9 *4s eaeuUTpo) 
STSusIYysSeieyeuw eaeuTpo) 
Bi8stu-ooneyTs eaeurpoyg 
styTeotde eaeurpoy 
aetTpuertoz—-9eaou eisodsozrosue7 
erodstaaiq eiodsoTpo.uez 
euedtTisweoOrysne eiodsoz,ouezZ 
eqet{[[rotued eursdoqjeeyy) 


with 
mono 


that have 


cells 


JIA noSenSe iS 
other with 


D. 


Dictyochaeta 


conidrorscenous 
and jthe 


of 


BSON VITA 
eqyeqstie 
st[Teieqyertun 
STSUSTIOAT 


BQVoaeyIouOoTy 
eqVoey.ou0ry 
eutsdojaeyy) 
eJVoeyIouoTy 


eoT}JueWwoOI stsdoqsaeyy 


eptpuotds 
BA[N} 
elojytwed 
stsuotuoy 
epuetskeTeuw 
esstds 
stsuouinqne 


eutsdoqjaeyy) 
eutsdojeeyy) 
eqVoaeyIouoTy 
eJyaeYyIoUOTY 
eVOePYIOUOTY 
eVoeyIouOTy 
eutsdoqeeyy) 


esoqjesotetd stsdoiodstuap 
aeworqoayq stsdoiodstuop 
esnjoid stsdozodstuap 
aetpueztoz—-aeaou stsdoiodstuay 
BuetotaAopnyt stsdoiodstuap 
eiodsoirputpTAd eqaeyoso0hqITG 
wnjeqdestiq wnydtaqyorputpiAg 
wn}zeTNotoOsezy wnyItzA,VOApuTTAg 
awioytuetl wntptszopyyg 
Tiodotyssity erysoqzeyxezZ 
@SUSTRERBASURI WNTIPTIOT YN 
stiodsopAwe[yo wntpti0T yg 
unias8tpneds wnNtptTroT yg 
wnaptok130q wWNTptsz0oT yy 
STSUDOUTTTI BJIeYIOARITG 
tadfkyTeono eqaey rok ITG 
eydioutp eqyoaeyoof TG 
eBJeutounN esodstusp 


epotTuueqjtaq erodstuay 


species 
wauocrpha hasytwov different kinds .of conidlophores), 


eoyociaets dimorpha, D. ceucalypti and 


Oeeuieomewith polyphialidic 
sympodial proliferations 


merery ’particular 


Boh ESE 


D 
D. 


igri t sal 
® BAe 
pm 
a0 wo aS 
ey a0 0 
fd OH oO 

> oor 
snrn® 
AS [| ae = 

oP i 
Yn 

oD) 
Bg? = ys 
Ox o) 
< OG 45 
Goa ela 

4 SS 

ae) 

‘a sy 
G4 Cee ay 
OB g 7S 

gy @ “ey 

o 

fo. 
SH og a @® 
) qos 
Sg err 
Oa we 

See 

yore 

bas oO) 
1) on ee 
reg | O 
dH Ogee 
o Oo a + 
OH 6 >) 

O- Ore @ 

pales ee 
nO, 10 
3h p w 
oOoa So 
oo 900.4 
o) ge - 
Qo ia] 
CF Meet 
Bi wm Oo 
oa iS Ol. 
Soa (a) 
06°C ged 

w “4 

qf 

es © O 
de et Oe) © 
OoOr BB 
AN ores 
don Qo 
Cet 3 0 > 
AGP OM Y& 
cle Os Mees 
OL Dey - 


no setae accompanying the conidiophore. 


are 
one 
many 


Pmarshe. polypnialidicpiormy wach sepaa te 


Species 
and setulate conidia. 


dered the 


oii | 
Bo ay} 
aa pa 
3 O) 
ep 
4 oO 
i) = 
$5) Oo 
n Sy 
oO 0 wh 
CaS 
ay} 
2% 
yo fe) 
1 cS 
ao 
© a “4 
oO c= 
n 
3 ® 
© re 
S ) 
© cn 
Q0 OQ, 
Oo ) 
“cd Pe, 
se O 
op = 
= qf 
o O 
S) 
3) 
2 Yn 
Ae, 
ae) ray) 
Sey 


ti 


eucal 
pice dincercalary without proliferations, 


Le, 


illinoensis 


Pas)’. 
D. 


Gurved and septate conidia. 
tions, 


slightly 


1-septate, 


accasionally 


convdwaw ace 
symmetric and bearing rudimentary setulae. 


the 


curved, 


694 


For such reasons. these species of Dictyochaeéta sams 
less related with the other species of the genus. 

Another group is made up by Chloridium botryoideun, 
C. caudigerum, C. transvaalense, Zakatoshia hirschiopori, 
Chloridium reniforme, Cylindrotrichum  fasciculavtumeaae 


triseptatum and Dictyochaeta cylindrospora. In these spe- 
cies the absence of proliferations either sympodial and/or 


percurrent is what relates them. The relationships between 
D. cylbindrospora and the species of Chloridium and Cylin- 
scone mentioned above is not strange, since Morgan 
Jones (1976) stated: 'Codinaea cylindrospora is not enti- 
relly typical of Codinaea as defined by Hughes and Kendrick 
(1968) on account of the absence of phialidic prolifera- 
tions, to produce polyphialides'. The absence of prolifera 
tion, \and the non septate  ; “straight. and Gy Bio 
conidia appear to relate D. cylindrospora with the species 
of the genera above mentioned. Zakatoshia hirschiopors 
also appear in this group by the absence of proliferavign 
and the conidia similar to those of the Chloridium 

The species of the genus Menisporopsis appeéarwrelaver 
by the presence of synnematous conidiophores, the synnemave 
have central thick-walled setae. 

The second group is featured by the conidiogendue 
cells growing sessile or in branches on the conia@gepgiiee. 
Chaetopsina auburnensis, Kionochaeta spissa, K. malaysiana, 
K. keniensis, K. ramifera make up '(a 'groupy" rela vecwaae 
another made up by “Chaetopsina fulva, CG. splendida, GChae= 
topsis romantica and Kionochaeta ivoriensis. Less related 
with these two. groups* we found Chaetopsina unilateraiias 
Kionochaeta aristata and K. virtuosa. K. nanophora and 


K. pughii which constitute a cluster, andseGhéegop sae 
catenulata, G. polyblastia and Chaetopsis grisea whic 
constitute another. Both are far from the other Chaetopsing 
and Kionochaeta, because they have sympodial or percurrent 
proliterat pom. 

Chaetopsina® and .Kionochaeta include species ewig 
setiform conidiophore ‘and - conidiogenous., celta eee 
on branches arranged at-the-apex, at the mid» pargegne ne 
conidiophore or with both dispositions, as "in hee 
Kirk & Sutton (1985) coneurred with Samuels (1985) 
restricted Chaetopsina to the anamorph of the Nectriaceae 
with reddish-brown setose conidiophores that turn yellow 
with 100% lactic acid. When chose the characters Wwowais 
not include, neither the presence or absence of téleomnorpr 
nor the reaction with: lactic acid because they aremenaraces 
ters that were not studied'in the majority of ther speciee 
included in this .analysis.: However “we wonder) jee 

two. characters used by Kirk and Sutton to separate boum 
genera have sufficient taxonomical value to bemused 
such a way. On the other hand, what will happen with some 
species of Kionochaeta if new studies of the anamorph-te= 
leomorph relationships result in them being ~anamorpne 
of a» Vectriaceae?.. 

Chaetopsina penicilliata, Zanclospora novae-zelandiae, 
Z. brevispora and Z. austroamericana form another group. 
They share the following characters: conidiophores exten- 
ding but not forming setae, a lagéeniform coOnidieperars 


695 


cells without proliferations. The presence of ‘conidiogenous 
eglls growing on apical branches is the difference’ between 
Chaetopsina penicilliata and the three species of Zanclos- 
mora. these species have sessile conidiogenous,.cells at 
the mid level of the conidiophore. 

Codinaea apicalis, C. glauco-nigra, C. maharashtren- 
sis,C. state of Chaetosphaeria dingleyae, C. filamentosa, 
C. elegantissima, C. intermedia and Dictyochaeta menispo- 
roides make a group different from the other species of 
the genus Dictyochaeta because they have branched conidio- 
phores and conidiogenous cells growing on them, they have 
Conspicuous collarettes and the extension of the conidio- 
phores may be straight or flexuose in Codinaea filamentosa. 
Morgan Jones etal.(19%) considered that C. apicalis and C. 
glauco-nigra could be separated in a new genus due to 
the characters last described. Holubova-Jechova (1984) 
consider that C. elegantissima, G. intermedia, ©. filamen- 
tosa, Dictyochaeta menisporoides, GC. state of Chaetos- 
phaeria dingleyae, and C. maharashtrensis are in similar 
Situation. 

Menispora ciliata, M. glauca, M. tortuosa, M. manito- 
baensis and M. state of Chaetosphaeria pulviscula share 
the following characters: conidiogenous cells uncinate 
without proliferations and growing on the conidiophores. 
M. britannica and M. uncinata differ from the above mentio- 
ned Menispora because they have the conidiogenous cells 
integrated to the conidiophore; they have sympodial proli- 
ferations and independent setiform . structures, due to 
iiveyemessons they are closely related to Dictyochaeta 
in the first group. Nevertheless they have conidiogenous 
cells abruptly and strongly recurved towards the conidio- 
phorous axis so that a poorly differentiated tubular colla- 
rette is directed towards the base of the phialide (uncina- 
te). For this characters Hughes & Kendrick (1968) and 
Kirk (1985) considered them in Menispora. 

The species of the genus Gonytrichum form a cluster. 
The presence of conidiophores bearing one or several short, 
curved, lateral branches (collar hyphar) below the septa 
from which cluster of phialides arise appearing verti- 
cilliate and the conidiophore extending forming or not 
a setae, are the characters that we only find in the spe- 
Clesrorytnis genus. 


CONCLUSIONS 


From the analysis of the results we can conclude: 

The genus Dictyochaeta is well defined. Codinaea 
apicalis, C. state of Chaetosphaeria dingleyae, C. elegan- 
tissima, C. filamentosa, C. lauco-nigra, G. intermedia, 
C. maharashtrensis and Dictyochaeta menisporoides' need 
a different treatment. For this reason we remaing them 
in Codinaea. 

Bahusutrabeeja, Chloridium, Cylindrotrichum and Zaka- 
toshia are genera very related and the differences between 
them,are based only on’ the; morphology of the conidia, 
and sometimes these differences are so minute that it 
is difficult to find then. 


696 


Menispora britannica and M. uncinata must be reconsi- 
dered their systematic position in Menispora or Dictyochae- 
ta. 

Menisporopsis is a good genus with well defined cha- 
racters. The same happens with Gonytrichum and Zanclospora. 

The considerations about Chaetopsina and Kionochaeta 
are in the results. Probably both genera must be reconside- 
rece 

The two species of Ghaetopsis considered in this 
analysis appear clustered with different species of Chae- 
topsina and Kionochaeta and need a revision. 


ACKNOWLEDGEMENTS 


We thank Dr. Irma Gamundi, Dr. Jorge Wright and Dr. Maria 
F. Lopez -Armengol for their critical review of this manus— 
Cd Pils 


REFERENCES 


CABELLO, M. & A. ARAMBARRI. 1988. Considerations about the validity 
of the genus Cylindrotrichum Bonorden. Mycotaxon 31(2): 435-438. 

DICOSMO, F.; S. BERCH & B. KENDRICK. 1983. Cylindrotrichum, Chae- 
topsis and two new genera of Hyphomycetes, Kylindria and Xenoky- 
lindria. Mycologia 75(6): 949-973. 

GAMS W. & V. HOLUBOVA-JECHOVA. 1976. Chloridium and some other dema- 
tiaceous Hyphomycetes growing on decaying wood. Stud. Mycol. 
13: 1-99. 

HOLUBOVA-JECHOVA V. 1984. Lignicolous Hyphomycetes. ‘7. Folia Geobota- 
nica et Phytotaxonomica 19: 421-438. 

HOLUBOVA-JECHOVA, V. & A. MERCADO SIERRA. 1984. Hyphomycetes from 
Cuba II. Ceska Mycologie 38(2): 103-104. 

HUGHES S.J. & B. KENDRICK. 1968. New Zealand Fungi. 12: Menispora, 
Codinaea, Menisporopsis. N. Z. Jl. Bot. 6: 323-375. 

KENDRICK, B. 1980. The generic concept in Hyphomycetes a reappraisal. 
Mycotaxon 11: 339-364. 

KENDRICK, B. & F. DICOSMO. 1979. Teleomorph-anamorph connections 
in Ascomycetes pp. 283-410. In: The whole fungus Ed.,B. Kendrick. 
National Museum of Natural Sciences, National Museums of Canada 
and Kananaskis Foundation, Ottawa. 

KIRK, P.M. 1985.- New or interesting microfungi XIV. Dematiaeeous 
Hyphomycetes from Kenya. Mycotaxon 23: 305-352. 

KIRK, P.M. & B.C. SUTTON. 1985. A reassessment of the anamorph genus 
Chaetopsina (Hyphomycetes). Trans. Br. Mycol. Soc. 85(4):709-718. 

MORGAN JONES G. & E. G. INGRAM. 1976. Notes on Hyphomycetes XV. Two 
new species of Codinaea. Mycotaxon 4(2) 502-509. 

ROHLF, F.J.; J. KISHPAUGH & D. KIRK. 1971. NT-SYS. Numerical Taxonomy 
System of Multivariate Statistical Programs. Tech. Rep. State 
University of New York at Stony Books, New York, N.Y., 87 pp. 

SAMUELS, G.J. 1985. Four new species of Nectria and their Chaetopsina 
anamorphs. Mycotaxon 22: 13-32. 

SNEATH, P.H.A. & RR. SOKAL. 1973. Numerical Taxonomy. The principles 
and practice of numerical classification. Freeman, San Francisco, 
UOeg AV st OV oUp. 

SOKAL, RP. & F.J. ROHLF. 1962. The comparison of dendrograms by 
objetive methods. Taxon 11:33. 

WILCOX H.E.; R. GAMMORE-NEUMANN & C.J.K. WANG. 1974. Characteristics 
of two fungi, producing ectendomycorrhizae in Pinus resinosa. 
Caren ws Bots Desnen (92290. 


MYCOTAXON 


Vol. XXXIV, No. 2, pp. 697-712 January-March 1989 


THE GENUS CLADIDIUM (LICHENIZED ASCOMYCOTINA) 
BRUCE D. RYAN 


Department of Botany, Arizona State University 
Tempe, AZ 85287 USA 


ABSTRACT. The dwarf fruticose lichens of Lecanora sensu lato 
described by Tuckerman from coastal California are revised, with 
emphasis on the genus Cladidium Hafellner. The new combination 
Cladidium bolanderi (Tuck.) Ryan is made, with C. thamnitis (Tuck.) 
Hafeliner treated as a synonym. These taxa and Lecanora phryganitis 
Tuck. are lectotypified. 


INTRODUCTION 


The dwarf fruticose species of Lecanora Ach. sensu lato described by 
Tuckerman (1864, 1866) from coastal California have not been investigated in much 
detail previously. The purposes of the present paper are: 1) to clarify the 
supraspecific classification of Lecanora bolanderi Tuck. and Lecanora thamnitis Tuck.; 
2) to provide detailed descriptions of these taxa; 3) to discuss the relationships of 
these taxa to each other and to another species, Lecanora phryganitis Tuck. (to be 
discussed in more detail in a separate article) in relation to the lectotypifications 
made in this study. 


METHODS AND TERMINOLOGY 


In the descriptions of the species below, the following items should be noted: 
numbers given in parentheses after colors refer to Kelly (1965); colors of the thallus 
and apothecia are as seen under a dissecting microscope (American Optical Spencer 
with fluorescent lighting, at 40x). More critical observations of external morphology 
were made with a Zeiss dissecting microscope with fiber optic lighting. Unless stated 
otherwise, observations and measurements of spores, hyphae, etc. were made on 
wet mounts, using an Olympus model BHB microscope. Crystalline deposits were 
observed using polarized light (crossed Nichol filters). Most sections were made by 
hand and mounted in water or lactophenol-cotton blue for semi-permanent slides 
(coverslip attached on one side by plastic cement). Photomicrographs of selected 
hand-sectioned, gold-coated specimens were made with a _ scanning electron 
ie pecope with the help of Mason Hale and a technician at the Smithsonian 
nstitution. 


The chemical analyses reported here were made by the standard thin-layer 
chromatographic method using the three main solvent systems of Culberson (1972), 
with substitution of methyl tertiary-butyl ether for diethyl ether in system B (Culberson, 
1982) (= B’) and substitution of toluene for benzene in system A (Culberson, et al., 
1981). The chemistry of several specimens was also analyzed by C. Leuckert, using 
lichen mass spectrometry. 


Spot tests and anatomical observations were made using the following 
reagents: 10% aqueous potassium hydroxide solution (K), freshly prepared dilute 
alcoholic solution of paraphenylenediamine (Pd), sodium hypochlorite solution in the 
form of household bleach (C), concentrated nitric acid (N), and iodine-potassium 
iodine solution (I,Kl). Chlor-zinc-iodine solution, which stains for cellulose, was used 
for detecting empty algal cells, after pre-treatment in KOH and flushing with water. 


698 


Voucher specimens are deposited in the Arizona State University herbarium 
(ASU) unless otherwise cited. Herbarium abbreviations follow those in Holmgren & 
Keuken (1974): ASU, CANL, COLO, DAV, FH, G, GZU, H, IRVC, LAM, M, SFSU, 
UC, UPS, US, WU. Nomenclature of lichen taxa mentioned in this article, other than 
the fruticose Lecanoras, follows Egan (1987). 


Most of the terminology employed in the descriptions follows the usage of 
Eigler (1969). A few of the terms used in the present article need further 
explanation: 1) "spermogonia" and their "spermatia” are used here because it is 
likely that the very slender "pycnospores" which contain very little cytoplasm function 
in sexual reproduction rather than as asexual "conidia"; 2) the "hypothecium” is 
regarded here as the lower part of the excipulum, and "subhymenium" is used for the 
ascogenous layer between the hypothecium and the hymenium; and 3) “epithecium", 
which properly applies only to a plectenchyma, is replaced here by “epihymenium” for 
the upper part of the hymenium and "epipsamma" for granular material in or on top 
of the hymenium. 


SUPRASPECIFIC CLASSIFICATION 


Tuckerman’s concept of Lecanora sect. Cladodium (which he mentioned in 
1872 and defined in 1882) included all three of his coastal Californian fruticose taxa, 
L. bolanderi, L. phryganitis and L. thamnitis, and was based only on the "fruticulose"” 
form of the thallus, sessile apothecia and simple spores. Zahlbruckner (1907; 1926) 
gave a slightly more detailed description of the section, including the misleading 
statement that the thallus is “unberindet" (ecorticate). In his classification system, 
Rasanen (1943) included "Cladodium Tuck.” as a genus, but he cited neither 
individual species nor the basionym (Lecanora sect. Cladodium Tuck.) and therefore 
did not validly publish a new combination for it as a genus. 


Hue (1909a) described the genus Polycauliona, based mainly on growth form, 
and he gave as the synonym "Placodium sect. Thamnonoma Tuck." [= Caloplaca 
sect. Thamnoma (Tuck.) Zahibr.]. Hue (1909b) transferred Lecanora bolanderi, L. 
phryganitis and L. thamnitis (along with other species) to Polycauliona, and this time 
also included Lecanora sect. Cladodium among the synonyms. Both Zahlbruckner 
(1901-1914) and Poelt & Pelleter (1984) argued that Polycauliona is a heterogenous 
and unnatural group. Although Polycauliona needs to lectoty ified, the name was 
originally based on a group of species now in Caloplaca and therefore cannot be 
used as a genus for Tuckerman’s fruticose Lecanoras. 


Gyelnik (1934) raised Lecanora sect. Cladodium Tuck. to the level of genus, 
citing Tuckerman (1872). However, he explicitly mentioned only Cladodium 
Hosseanum Gyelnik (isotype at M! = Rhizoplaca cf. melanophthalma), which was not 
included in the protologue of the basionym and therefore cannot be the lectotype 
species of Lecanora sect. Cladodium. According to the Code (Voss, et_al., 1983; 
Articles 64.1 and 45.3) Gyelnik’s genus Cladodium must be rejected as a later 
homonym for the moss genus Cladodium Bridel (1826). 


Eigler (1969), in his treatment of Lecanora sensu lato, suggested that (based 
on various anatomical features) the "L. thamnitis group" should be a separate genus, 
but he did not formally propose a new combination. He explicitly cited L. thamnitis 
as type species of Lecanora sect. Cladodium, but did not cite particular specimens, 
nor did he mention L. bolanderi or L. phryganitis. 


Hafeliner (1984) used the genus Cladidium as nomen novum for Cladodium 
Gyelnik (1934), cited Lecanora sect. Cladodium Tuck. (1866) as a synonym, and 
changed the "o" in Cladodium to an "i" to legitimize it as a genus. He explicitly cited 
L. thamnitis as the type species of the genus Cladidium and as Eigler’s lectotype of 
Lecanora sect. Cladodium. Hafellner stated that in contrast to L. thamnitis, in which 
the ascus tips have a very wide axial mass (similar to that found in Candelariaceae), 
L. bolander! has a typical Lecanora-type ascus and should be excluded from the 
genus Cladidium. e specimen of L. thamnitis he cited was Relig. Tuck. 116 
(GZU!), which is an isolectotype of that species (see below). Hafellner based his 
concept of "L. bolanderi” on a misidentification of L. phryganitis. However, based on 


699 


the lectotypes chosen below, true L. bolanderi belongs in the genus Cladidium, and 
L. thamnitis is treated here as a synonym of L. bolanderi. 


Relationship of Cladidium Hafellner to Other Taxa 


At present the genus Cladidium appears to have no obvious close relatives. 
The distinctive thallus anatomy of L. bolanderi and L. thamnitis (alternation of dense 
fungal tissue with loose algal tissue--Fig. 1) helps to separate Cladidium from 
Lecanora. However, a similar structure occurs in most fruticose species of Caloplaca 
(e.g., C. coralloides), is common in various otherwise dissimilar lichens of coastal and 
xeric habitats, and probably has an adaptive function connected with water relations 
(Poelt & Pelleter, 1984). A less extreme form of this anatomical type (confined to the 
upper layers of the thallus) occurs in some lobate crustose Lecanora species. 


Lecanora phryganitis (which often occurs side by side with Cladidium and with 
Caloplaca coralloides), is however quite different in thallus anatomy (Fig. 2) as well 
as other features (Table 2) and belongs in a natural group with L. pinguis Tuck. (type 
species of Lecanora subg. Placodium sect. Endochloris, described by Poelt [1958)). 
Lecanora phryganitis is excluded from Cladidium, and is treated in similar detail in a 
separate paper as: 


Lecanora pbryganitis Tuck., Lich. Calif. p. 19 (1866). 

Type (from the protologue): "On sandstone rocks, coast; Mr. Bolander." 
Lectotype (selected here): "...on metamorphic sandstone, near the Mission Dolores 
... 1/12/64", Bolander 63, (FH!) (Figs. 3-4). 


Polycauliona phryganitis (Tuck.) Hue, Compt. rend. congrés Soc. Sav. en 1908: 
153 (1909b). 


DESCRIPTION OF THE TAXA 


Cladidium Hafellner, Beih. f. Nova Hedw. 79: 296 (1984). Nomen novum for 
Cladedium (Tuck.) Gyelnik. Lectotype species (Hafellner, 1984): L. thamnitis Tuck. 


Lecanora sect. Cladodium Tuck., Synopsis N. Am. Lich. 1: 181 (1882). 
Lectotype species (Eigler, Diss. bot. 4: 169 [1969]): L. thamnitis Tuck. (1866). 


io SS (Tuck.) Gyelnik , Repert. Spec. Nov. Veg. 33: 305 (1934), non Bridel 


Cladidium bolanderi (Tuck.) Ryan, comb. nov. (Figs. 1, 5-12) 


Lecanora (Squamaria) Bolanderi Tuck., Proc. Amer. Acad. Arts Sci. 6: 266 
(1864). Type (from the Proeieeuay: "On rocks (‘metamorphic sandstone’) in Marin 
County, California, Mr. H. N. Bolander." Lectotype (selected here): "On 
metamorphic sandstone, Marin Co.", undated, Bolander 29 [specimen on card] (FH!). 


Pepe canna: Bolander!) Hue, Compt-rend. congrie, Soci Sev. en. 1208: 153 


Lecanora thamnitis Tuck., Lich. Calif.: 20 (1866). 
Type (from the protologue): "Sandstone rocks...Mr. Bolander." Lectotype (selected 
here): Oakland Hills, undated, Bolander, s.n., Relig. Tuck. 116 (FHI); 
Isolectotypes: (COLO!: 2 packets, GZU!, UC)!). 


Polycauliona_thamnitis (Tuck.) Hue, Compt.-rend. congrés Soc. Sav. en 1908: 
153 (1909b). . 


Cladidium thamnitis (Tuck.) Hafellner, Beih. f. Nova Hedw. 79: 296 (1984). 


Description of Cladidium bolanderi: 


700 


THALLUS: Dwarf fruticose, caespitose. Starting as small granules or 
verrucae. In the typical and most widespread form, becoming loosely tufted, much 
branched + dichotomously, the branches erect (or decumbent towards thallus margin), 
forming irregularly rounded "sods" (to 2-10 cm across) which are continuous to 
fissured and have an uneven surface (Figs. 7-8). In "L. thamnitis" form, branches 
densely crowded, mostly erect and similar in height, giving patches a more even 
surface; forming either extensive crustlike mats without definite margin (Fig. 10), or 
discrete hemispherical to irregular peltate clumps 5-15 mm diam. (Fig. 11), with 
branches radiating from a central main trunk, attached to rock by a single irregular 
area 1-4 mm wide. Branches: Terete, to irregularly compressed near thallus margin 
(or terete throughout in "L. thamnitis"), 5-10(-15) mm, irregularly thickened (0.5-1 
mm), tips papilliform (usually scarcely widened, 25-50 per cm’; narrower and up to 
100-200 per cm’ in "L. thamnitis"). Surface: Matt; continuous and + even, not 
verrucose or granular, but pale spots often slightly raised and roughened; without 
vegetative propagules. Color: Basic color mostly light to dark grayish olive (109- 
111) or grayish greenish yellow (105), with irregularly shaped pale whitish or 
yellowish areas (pseudocyphellae or maculae in the broad sense), especially toward 
lobe tips (in "L. thamnitis", mainly the strongly whitened branch tips are visible from 
above). Little changed when wet. Undersides and basal parts mostly pale yellowish 
green (121), pale to moderate yellow (87, 89) to moderate orange-yellow (71) or 
moderate yellowish brown (77), without pale spots. In old herbarium specimens, 
turning moderate orange (53) to various shades of brown (55-56) or yellowish brown 
(74-75), paler below, darker and more grayish towards base. Consistency: 
Cartilaginous, brittle but not crumbly. 


APOTHECIA: Usually common; 0.5-4 mm diam., terminal or subterminal, 
sessile to somewhat constricted at base (in "L. thamnitis" appearing to be pany 
immersed within thallus patch). Disc: Concave to plane; usually yellowish to whitis 
or grayish Pluie. (at least when young); mostly various shades of yellow (86-87, 
101) or yellowish brown (75, 77) under pruina, little changed when wet. Sometimes 
turning blackish or reddish due to parasites. Margin: Usually swollen (to 0.5 mm 
thick); entire or (especially in L. thamnitis) becoming crenate and irregularly lobed; 
even with disc or somewhat raised and inflexed when young; persistent. Concolorous 
with upper parts of thallus, usually strongly pale spotted and somewhat roughened 
above. No proper margin visible. 


SPERMOGONIA: Present on or near tips of branches; completely immersed; 
ostiole pale. Cavity 150-200 um diam. in section; wall hyaline. Spermatia: 
Threadlike, curved, (12-)15-25 ym (based on Jordan 3447 [GZU], Bolander, s.n. [H- 
Nyl. 28323], Setzer 697, Ryan 21968). 


THALLUS ANATOMY:  Thalius composed of irregular areas of dense 
medullary tissues, alternating with loose tissues mostly containing algae; with network 
of irregular elongated areas appearing opaque due to yellowish granules (K- 
insoluble), partly associated with algae. Cortex: True cortex (without empty algal 
Cells); Continuous; ca. 25-45 wm thick; inspersed toward outside with yellowish 
granules (partly soluble in K). Hyphae mostly anticlinal, otherwise similar to those of 
medulla. With 2-3 tim thick epinecral layer. Algae: Algal cells Trebouxioid, 7.5-10(- 
15) ym diam.; arranged in irregular, often radially elongated clumps in areas of 
optimal light, occurring within areas of loosely arranged tissue with gaps in it 
(aerenchyma), which consist of hyaline, thin-walled hyphae (ca. 2 wm diam.) and 
penetrate to the surface as pseudocyphellae. Medulla: Prosoplectenchymatous 
(scleroplectenchymatous); hyphae densely interwoven and agglutinated, irregularly 
thick-walled, ca. 2 um diam., with indistinct outer boundaries, non-granular yellow- 
orange wall pigmentation, and distinct lumina. 


APOTHECIAL ANATOMY:  "“Thamnitis-Typ" of Eigler (1969--Plate 25: 3). 
Hymenium: "Thamnitis-Typ" of Eigler (1969--not illustrated), (50-)70-90(-100) um 
high, colorless below, not inspersed. Epihymenium pale; epipsamma 10-20 um thick, 
mostly superficial, composed of yellowish granules (partly soluble in K). Spores: 
Ellipsoid to oblong-ellipsoid or ovoid-ellipsoid (L:W = 1.5-2.5:1), mostly (8-)10-14(-16) 
x 5-6(-8) um; moderately thin-walled; mostly uniseriate, 8 per ascus; simple, usually 
with one or two oil drops. Asci: Clavate to subcylindrical, ca. 50 x 8-12 ym; outer 
wall layer and gelatin strongly amyloid, tips with amyloid ring, with axial mass very 


701 


Pann iar varsarae ere 


Figs. 1-2. Scanning electron micrographs of iongitudinal sections of branches. Scale 
= 200 pm. -1. Cladidium bolanderi (Tuck.) Ryan: typical morphotype (Krypt. exs. 
Vindob. 1870, US). -2. Lecanora phryganitis Tuck. (COLO Exs. 475, US). -3,4. 
Morphology of L. phryganitis. Parts of lectotype (Bolander 63, FH). Scale = mm. 


702 


broad toward the outside and appearing almost non-amyloid (based on Relig. Tuck. 
116 [FH], Bolander, s.n. [H-Nyl. 28323], Setzer 697). Paraphyses: "Atrynea-Typ" of 
Eigler (1969--Plate 17: 33); simple to furcate; septate; 1-1.5 ym thick below; apical 
cells only slightly thickened (to 2-2.5 ym), colorless, thin-walled; somewhat loosely 
coherent, not strongly gelatinized. Subhymenium: About 50 um thick, colorless. 
Excipulum: Hypothecium: To 500 ym thick, of agglutinated hyphae with threadlike 
lumina, randomly oriented to mostly anticlinal, colorless to yellow-orange, not 
inspersed. Parathecium: 25-90 um thick, of parallel hyphae. Amphithecium: 
Structure similar to that of thallus. Cortex: About 50 um thick. Algal Layer: 
Present in margin and below hypothecium; to 65 ym thick, divided into clumps of 
algal cells. 


SPOT TESTS: Cortex: C-; partly K+ yellow; KC+ yellowish; Pd-; N-. 
Medulla: C-; K-; KC-; Pd-; N-; I,Kl-. Discs: C-; K-; Pd-. 


CHEMISTRY: Usnic acid or isousnic acid (occasionally both), usually with 
several unknown (probably phenolic) substances found by thin-layer chromatography 
(Table 1), in almost all possible combinations ("L._thamnitis" morphotype usually 
contains usnic acid and unknown CBL-1, but "L. bolanderi” morphotype shows more 
variability, often containing isousnic acid and unknown CBL-2). Report of usnic acid, 
arthothelin and thiophanic acid from L. bolanderi (Santesson, 1969) is based on L. 
phryganitis (specimen annotated by Santesson not seen). 


Table 1. Partial list of secondary metabolic substances found in Cladidium bolanderi 
sensu lato (including "Lecanora thamnitis"). Data are from thin-layer chromatography, 
based on the method of Culberson (1972), with substitution of toluene for benzene in 
solvent system A and methyl-tert.-butyl ether for diethyl ether in solvent system B 
(=B’). Numbers after colors in "Color after acid" column refer to Kelly (1965). 


R,-Classes Substance UV (LW) Color UV (LW) 
after acid after 
A B' C charring charring 
2-3 2 2 CBL-1 light violet yellowish gray dark greenish 
[210] (faint)[93] gray [156] 
4 2-3 3 CBL-2 grayish brown light yellow grayish brown 
[61] [86] [61] 
5-6 6 6 CBL-3 very light 
green [143] 
5 5 5 CBL-4 light orange 
yellow [70] 
6 6 6 Usnic quench light to dark quench 
acid greenish gray 
[154-155] 
Prades: ‘é Isousnic quench grayish olive quench 
acid [109-110] 


ECOLOGY: On siliceous or ultramafic rocks (sandstone, chert, granite, serpentine), 
on horizontal to steep but not overhanging surfaces, often on large, exposed boulders 
on bluffs near the seashore, 20-550 m elev. ("L. thamnitis" may be more common at 
upper end of this range). Probably strongly nitrophilous; sometimes on rocks covered 
by guano. Associated organisms: Associated vascular plant communities, include 
Umbellularia-Quercus chaparral and mixed grassland. Associated lichens include: 
Lecanora_phryganitis, L. pinguis, L. muralis, Arthonia_phaeobaea, Buellia spp., 
Caloplaca rosei, C. coralloides, Lecidella subincongrua, Niebla spp., Phaeophyscia 


703 


PPPUECPTEE LTPP UEP ee 


TET 


[ad 
Cael 
bail 
boa 
reset 
Res 
- 
_ 
ee. 
boat 


Figs. 5-8. Cladidium bolanderi (Tuck.) Bye typical morphotype. -5. Lectotype of 
Lecanora bolanderi Tuck. (Bolander 29, FH: specimen attached to card). Scale = 
mm. -6. Box labelled “original specimens! (Bolander, s.n., FH). -7,8. 
Representative thalli attached to rock (Ryan 21968, ASU). Scale = 5 mm. 


704 


spp., Physcia spp., Punctelia_stictica, Xanthoria candelaria, and sometimes Collema 
spp. or Verrucaria cf. maura. Hymenium and apothecial margins sometimes eaten by 
snails. Often with numerous black mites (Acarina) within the thallus clumps. Often 
with black ascocarps (Arthonia sp.) and red areas on the thallus and discs 
(unidentified parasite). 


DISTRIBUTION: USA: Central California, north to southern Oregon. Report of 
L. thamnitis from Alaska by Calkins (1899--specimen not seen) is probably erroneous. 


ADDITIONAL LITERATURE REPORTS: See Zahlbruckner (1921-1940) and 
Tucker and Jordan (1979) for numerous citations; also see Weber (1981). 


EXSICCATI: USA: CALIFORNIA: San Mateo Co.: (Pigeon Pt.): Krypt. 
Exs. Vindob. 1870, Herre, s.n. (FH, G, M, US, WU seen); Alameda Co.: (Oakland 
Hills): Relig. Tuck. Exs. 116 (lectotype of L. thamnitis: FH; isolectotypes: COLO: 2 
packets; FH; GZU and UC seen). 


OTHER SPECIMENS EXAMINED: Unless stated otherwise, all specimens 
listed below were originally labelled as L. bolanderi and are + representative of that 
morphotype. 


USA: CALIFORNIA: Alameda Co.: (Oakland Hills): Bolander 125 (L. 
thamnitis)(FH); Bolander, s.n., undated (L. thamnitis) (FH; M). Marin Co.: Bolander 
29 eds of L. bolanderi: FH--specimen on card); Bolander, s.n., undated (COLO 
L-10312; FH; UC 666809); Bolander, s.n., 1867 (FH); Murray, s.n., 4.28.1982 
(SFSU); (Vicinity of Stinson Beach): Parks L-104 (COLO S-6136; UC); Parks 2697 
(UC); Peters 9168 (SFSU); Weber & Kunkel L-60850 (COLO); (Between Pt. Reyes 
& Inverness): Weber 75-93 (DAV); (1 mi E of Jct. Nicasio-Pt. Reyes Rd.): Weber 
& Kunkel L-60804 (COLO; M; US: 2 packets); (Hill E of Muir Woods): Wennekens 
64 (SFSU); (Mt. Tamalpais State Park): Riefner 84-75 (IRVC); (Ft. Cronkite): Setzer 
697 (ASU, SFSU); (Road to Stinson Beach): Ryan 21967, 21968; (Rocky Pt.): 
Volk 842 (SFSU); (Olema): Bolander, s.n., Tamales Bay, "Original Specimens!" (FH); 
Bolander, s.n. (H-Nyl. 28322), Bolander, s.n., 1864 (FH; UC 43264), Howe 15, (UC 
528927; US; specimen at FH contains no Cladidium), Bolander, s.n., 1868, "Boulders 
near Olyma [sic]" (G: 2 packets); (Pt. Bonita): Riefner 85-707 (COLO). Monterey 
Co.: (Monterey): Farlow, s.n., "Journey to California, 1885, Monterey" (as L. 
phryganitis) (FH: 4 packets; G; UPS); (Pt. Lobos): Ryan 21955 (ASU). San 
Francisco Co.: (Farallone Islands): Foreman, s.n. (COLO L-44,400); Lindahl, s.n., 
1886 (UPS); Tenaza 15526 (SFSU); Wheeler, s.n., June 1895 (UC 43266 & 43265); 
("San Francisco"): Bolander, s.n., 1867, "Californie" [sic], as L. thamnitis (G), 
Bolander, s.n., 1868, as L. thamnitis (G: 2 packets), Bolander, s.n., undated, (G), 
Bolander, s.n.? (51 on box inside), 1865 (US). San Luis Obispo Co.: (Morro 
Rock): Riefner 87-460 (IRVC; COLO). San Mateo Co.: (San Bruno Mt.): Bolander 
155 (as L. thamnitis) (FH); Herre 834 (FH; US: 2 packets); (Pigeon Point): Herre 
1308, (US); Herre, s.n. (7157), 1933 (US); (1 mi S of Pacifica City): Jordan 9 
(SFSU); (near Pt. San Pedro): Herre 506 (FH: 2 packets); Setzer 1953 (SFSU). 
Sonoma Co.: Jordan 3447 (GZU); Jordan 3461 (GZU); (near Jenner): Koch 479 
(UC: 2 packets); Halling 727 (SFSU); (Ft. Ross Historical Park): Riefner 85-608 
(IRVC); (Goat Rock State Park): Riefner 86-142 (IRVC; COLO); (Duncan’s 
Landing.): Halling 1296 (SFSU); Riefner 85-621 (IRVC); Riefner 86-159 (IRVC; 
COLO); (Carlevaro Rd.): Riefner 86-209 (IRVC); Riefner 85-625 (IRVC; COLO). No 
Locality Given: Bolander, s.n., undated, "ex. Willey herb." (as L. thamnitis) (FH); 
Bolander, s.n., 1865, "Coast of California" ("L. thamnitis", p.p.) (US); Hall, s.n., 1884 
(US); Herre, s.n., State Survey X-18 ("L. thamnitis’, p.p.) lc 208852); Bolander 
s.n., (H-Nyl. 28323); Bolander, s.n., "ex Willey Herb." (FH). ("San Francisco 
Watershed"): Thiers 15245 (SFSU). OREGON: Curry Co.: (Harris Beach State 
Park): Cooke 26803 (LAM); Cooke 26808 (LAM). 


VARIABILITY: Within Cladidium bolanderi sensu lato, variations occur in color 
and surface texture, morphology of branches and apothecia, size and shape of 
spoles and chemical content. These variations, which often occur side by side, 
show no obvious correlations with each other or with ecological factors, but more field 
study would be desirable. The two main morphotypes ("L._bolanderi" and "L. 
thamnitis"), their relationships to L. phryganitis, and the typification of these taxa, are 
discussed below. 


705 


x” 


ELEEELSAAEELL AALS ASS LEDERER SS LAKE LEER RTT OR ER ER ae AE RR RU Re eee ER oe ee 


FRARLALELELRADALE AGRE EARL IATAE LILI ARTE IET LITLE LALA AAR ARES PETA AEE ETAG AHL ELA ELAR ERT RIARGA URE I AERA PMH Oo EOE LEH. 


Figs. 9-12. Cladidium_bolanderi (Tuck.) Ryan: "Lecanora_thamnitis" morphotype. 
Scale = mm. -9. Lectotype of L. thamnitis Tuck. (Relig. Tuck. exs. 116, FH). -10. 
Representative thalli attached to rock (Bolander, s.n., Oakland Hills, FH). -11 . 
Peltate thalli (Bolander 125, FH). -12. Detail of branches from San Bruno Mountain 


collection (Bolander 155, FH). 


706 


DISCUSSION 
Relationships Among the Coastal Fruticose Lecanoras 


Tuckerman’s protologues distinguish L. phryganitis from the other two taxa by 
spore shape, position of the apothecia, and form and color of the thallus (Table 2). 
Later (1872) he stated: 


"...in L. phryganitis..we have neither the peltate fronds of L. Bolanderi, nor the 
effuse crust of L. thamnitis, but dense patches, made up, at the centre, of 
crowded erectish trunks, which are elongated, at the circumference, into finally 
decumbent branches." & 


My own study shows that the characteristics of L. phryganitis mentioned by 
Tuckerman may be correlated with other characters (Table 2). which make it very 
closely related to the lobate crustose species Lecanora pinquis Tuck., in spite of the 
conspicuous difference in growth form. In contrast, the characteristics originally 
observed by Tuckerman for L. bolanderi and L. thamnitis positively correlate with 
others, some of which (at least the ascus type) are sufficient for recognition of a 
separate genus for them, as discussed above. 


Although L. bolanderi and L. phryganitis have often been confused with each 
other, the protologues of the three fruticose taxa, as well as Tuckerman’s later 
comments (1872, 1882) and his annotations of specimens (see below) strongly 
suggest that Tuckerman applied the name L. bolanderi to material more similar to L. 
thamnitis than to L. phryganitis. 


Table 2 also compares L. thamnitis and L. bolanderi. Tuckerman’s protologue 


of L. thamnitis stated that his new species was similar to Cladonia papitata (now in 
eS and Lecanora aipospla (now in Lecania), both of which form mostly 
effuse, papillate crusts. Tuckerman’s protologue of L. bolanderi noted similarities 
between this new species and both Lecanora rubina (now Rhizoplaca_ chrysoleuca) 
and the Lecanora fruticulosa complex (now in Aspicilia). Later (Tuckerman, 1882) he 
discussed the analogous variation in growth form in these last two species (from 
effuse and stipitate-squamulose, to penate) The significance of such variation is still 


uncertain for these taxa (and in fact these inland species have sometimes been 
misidentified as one of his coastal taxal). 


Tuckerman (1872) characterized L. thamnitis as effuse and L. bolanderi as 
peltate, but stated: "There is yet no doubt, from specimens received since the 
publication of L. Bolanderi, that Its fronds are developed from scattered, papillaeform 
granules." In 1882, he stated: "L. Bolanderi offers etfuse conditions; as L. thamnitis, 
first observed only in such a state, passes finally into peltate ones, like the other.” 
(emphasis added). Although Tuckerman (1882) claimed: "the two lichens, in large 
sets of specimens, are, so far distinguishable," he considered all three new taxa (L. 
bolanderi, L. thamnitis and L. phryganitis) as closely related to each other, and 
"..probably only forms of one species", with apothecia similar to those of L. pinquis. 


Herre (1942), in discussing L. thamnitis under his description of L. bolanderi, 
stated: "An examination of specimens at the type locality revealed that a single 
colony showed every gradation from extreme bolanderi to extreme thamnitis." e 
went on to Say: Had Tuckerman, an able field worker, ever seen the plants 
growing, he would never have created L. thamnitis." This "type locality" (not 
specified), probably refers to San Bruno Mountain, where Herre had collected material 
somewhat transitional between the two forms (Herre 834, FH and US). Another 
Herre specimen from an unknown locality (UC 1208852) clearly demonstrates an 
almost complete range of morphotypes in a single patch. 


My own observations of highly variable morphology of such material in both the 
field and the herbarium indicate that L. thamnitis is probably at most only a form or 
variety of L. bolanderi (as lectotypified above). The "L. thamnitis" morphotype is 
presently known mainly from the Oakland Hills, where it was apparently abundant and 
fairly uniform. It would be useful to have further knowledge of both morphotypes in 
localities where both occur together, to better document the intergradations. 


Table 2. Comparison of Tuckerman’s coastal fruticose Lecanora species. 
from the protologues; those in parentheses ( ) 


bold, followed by asterisk (*) = 


followed by 172, T82 = from Tuckerman (1872, 1882); those in brackets [ ] = 
the author's own observations, where they ie 
numbers after colors refer to Kelly (1965). 

this study 
phryganitis from both L. bolanderi and L. thamnitis, and for distinguishing L 


707 


Items in 


from 


plement statements of Tuckerman; 
Underlined characters are considered in 


to be the most consistent and “REARS, useful for Seams ales | Bs 


from L. bolanderi. 


. thamnitis 


THALLUS 


Branches 


Color 


Surtace - 


Consistency 


APOTHECIA 


Discs 


Margin 


ANATOMY 


Spores 


Asa 
Spermata 
CHEMISTRY 


Lecanora_phryganitis 


fruticulose*; caespitose- 
Stipitate*; forming rounded 
patches® [usually with + 
definite jae (dense, 
crowded --T82) [to loose]; 
[peltate or compacted into 
crust]; [patches + uneven]. 


terete" [at least below]; 
erect, decumbent at margin 
[irregularly branched); 

to 1 inch tall* [to 510 mm 
in center, to 25 mm at 
margin); (tips obtuse --T82) 
[not papilliform; to 1-3.mm 
wide, to 25-50 per cm’]. 


ochre-white* [most le 


yellow-green (121) to 

greenish yellow (104), 

turning pale yellow (89) in 
erb.]; [not distinctly 

pale spotted]; 


rimulose-rugulose*; [often 
strongly verrucose to 
granular near tps}. 


{rather soft, crumbling}. 


lateral*; sessile* (sub- 
sessile --T82); large* (to 
medium --T82) [to 2-4 mm 
diam]. 


pruinose at first*; 
Ath flesh color* (pale 
rick colored --T82) [mostly 
Orange (53-54) to yellowish 
brown OST 
scarcely prominent’; 
subentire® (flexuously lobed 
-- T82); [< ca. 0.2 mm wide]. 


loose medulla}; [algae 
mostly in definite layer]. 


L:W_= 3-4:1*, oblong* ( 
oblong-ellipso ellipsoid -- mG ie 
-16 x 57 pm -T82) [mostly 
4-5 um wide). 

{axial mass narrow]. 
[(10-)12-15(-17) ym long]. 


[xanthones, zeorin and 
unknown LPN-1; usnic acid]. 


L. bolanderi 


fruticulose*; caespitose*; 
forming rounded patches* 
[usually with + definite 
margin]; (rather loose -T82); 
peltate, but effuse at first 

-- T82); [patches uneven]. 


terete* [compressed at margin]; 
erect*, [decumbent at margin]; 
dichotomously branched* [+]; 
[to 5-10{-15) mm in center, not 
muct longer at margin); tips 
papillitorm, obtuse® [0 1mm 
wide, to 25-50 per cm’. 


greenish straw color* [mostly 
grayish olive (109-111), 
ghee yellowish brown (74-75) 
in he Paine etd pale 
spotted] 


mostly continuous and even], 
not ee verrucose, not 
granular]. 


cartilaginous’. 


terminal*® (subterminal --T82): 
sessile* (to subpedicellate 
--T82); medium* (to large 

[to 2-4 mm diam_]. 


naked* [mostly pruinose!]: 
ellowish flesh color* (passing 
into tawny --T82) [most 
ellow (86-87) to yellowish 
rown (75-77)}. 


[somewhat prominent]: 
entire* [to crenate or lobed]; 


tumid®* {about 0.2-0.5 mm wide]. 


dense medulla’; [algal clumps 
scattered through thallus]. 


L:W = 1.5-2.5:1°, ovoid- 
ellipsoid [to ellipsoid]: 

10-14 x 6-8 ym —T82) [mostly 
5-6 um wide]. 


faxial mass broadened above]. 
[(12-)15-25 pm tong}. 
[no xanthones or zeorin; with 


unknown CBL-1; usnic acid or 
tsousnic acid]. 


L. thamnitis 


fruticulose*; caespitose- 
Stipitate*; forming verrucose 
crust without definite margin’; 
[very dense and crowded) 
ae also be peltate --T82); 


atches even). 


terete*, [also at margin}; 


erect® [also at margin]; 
[+ dichotomously branched]; 


somewhat over 1/4 inch talli* 
[+ uniformly 5 mm]; [tips 


papilliform], imostly < 0.5 mm 


wide, to 100-200 per cm’] 


greenish straw color* [mostly 
grayish olive (109-111), 

turning yellowish brown (74- 
75) in herb.; distinctly pale 
spotted]. 


mostly continuous and even], 
not strongly verrucose, not 
granular}. 


[cartilaginous]. 


terminal® (subterminal 
--T82); subpedicillate* [to 
sessile]; large* (medium 
--T82) [to 2-4 mm diam]. 


[mostly pruinose]; 

pale yellowish* (passing into 

tawny red --T82) [mostly 

ages (86-87) to yellowish 
own (75-77)). 


Sobeidatien prominent]; 
coming crenate® [to lobed]; 
[about 0.2-0.5 mm wide]. 


[dense medulla]; [algal clumps 
Scattered through thallus]. 


L:W = 2-2.5:1°, ovoid- 
ellipsold® {to ellipsoid]; 

10-14 x 5-7 yum --T82) [mostly 
5-6 um wide]. 

{axial mass broadened above]. 
[(12-)15-25 jum long]. 


[no xanthones or zeorin; with 
unknown CBL-1; usnic acid]. 


708 


Considerations in Typification of Tuckerman’s Taxa 


Tuckerman did not designate type specimens or even mention specific localities 
in many of his descriptions, nor date any of his annotations, resulting in much 
confusion. The following information regarding the Tuckerman herbarium was kindly 
supplied by Mr. Gennaro J. Cacavio, current curatorial assistant at Farlow Herbarium 
(pers. comm., 1988). Between 1937 and 1939, curators at FH (Dr. David H. Linder 
and others) reorganized and indexed the Tuckerman collection and affixed red printed 
"type" labels to many of the specimens. Although they did not explain their choices, 
Dr. Linder wrote: "All the type specimens were marked by the additional time- 
consuming labor of search through the literature to determine accurately the types." 
One of these curators, Mrs. L. W. Riddle, also prepared the "Reliquiae 
Tuckermanianae”" exsiccati (issued in 1942) from extra material which Tuckerman had 
set aside to distribute. In both cases the material was probably prepared without 
critical examination of specimens and distributed based on the data and determination 
provided by Tuckerman. 


According to Dr. Donald Pfister, current curator at FH (pers. comm., 1988), it is 
unlikely that these early curators found any additional information regarding these 
taxa in Tuckerman’s notes or correspondence. Because these "type" labels give no 
dates, names, or other information (other than citing the publication in some cases), 
this labelling by the early curators does not constitute valid lectotypification and 
without documentation does not offer proof of Tuckerman’s intentions. 


There is no reason to conclude that all of the original material of any of 
Tuckerman’s taxa is missing (necessitating selection of a neotype). Without proof 
that any of the known collections were "definitely studied by the author up to the time 
the name of the taxon was published", lectotypes must be chosen from among 
collections that at least were "clearly designated by the original author", giving 
preference to "any indication of intent by the author...unless contrary to the 
protologue.” (Voss, et al., 1983, Appendix 1.4). Except where noted otherwise, the 
annotations mentioned below are undated, and in several cases it is difficult to be 
sure who authored them, but in any case this is not essential, since none of the 
specimens were Clearly indicated as types by Tuckerman himself. Specimens that 
appear to be part of the same collection as the lectotype but do not have identical 
label data cannot be designated as isolectotypes. 


Typification of L. phryganitis 


Tuckerman (1866) cited the original material of L. phryganitis only as: "On 
sandstone rocks, coast; Mr. Bolander". The only locality mentioned by Tuckerman 
(1882) for this species was Mission Dolores (downtown San Francisco), which Herre 
(1910) referred to as the "type locality", stating that “material such as Bolander 
collected can no longer be obtained." (no material from San Francisco collected by 
anyone other than Bolander has been seen in this study). 


Lectotype: Bolander 63 (FH) (Figs. 3-4). 


The only collection labelled by the curators at FH as a "type" of L. phryganitis 
is Bolander 63, which contains four specimens (all similar to each other in anatomy 
and chemistry) on cards. One card has notes by Tuckerman comparing the species 
with L. thamnitis. The packet also has a note from Bolander, dated "1/12/64", saying: 
"No. 63. Quite abundant on metamorphic sandstone near the Mission Dolores and 
hence to the Ocean. Usually in depressions, forming there thick round patches. As 
much as possible | preserved that shape." (this note was cited by Tuckerman in 
1882, but not in the protologue). This collection was chosen as lectotype because it 
fits within the protologue of L. phryganitis, was collected prior to the publication, and 
: ean, ag ee (on all four cards) by Tuckerman, as: "No. 63. Bolander, California. 

. phryganitis T." 


Other Lecanora phryganitis specimens. 
Other Bolander specimens of L. phryganitis seen in this study are quite similar 


to the ones in the Bolander 63 collection (FH), but do not have collecting numbers 
and are not annotated by Tuckerman. A specimen at US, Boiander, s.n., "1864-70, 


709 


California" (labelled "authentic" by the curators at FH) is correctly identified (by 
Bolander?) as L. phryganitis. The ely clearly in Bolander’s handwriting, appears 
to be "Mission Hills" (possibly meaning Mission Dolores, in which case it could be 
part of the original material). However, "duplicates" of this specimen at FH and US 
are actually a Psora species!. At least one of several collections correctly labelled as 
L. phryganitis, from “San Francisco", Bolander, s.n., °/2 65 (sic; = unspecified date, 
February 1865) (FH) could have been seen by Tuckerman prior to publication. 


Typification of L. bolanderi 


The protologue of L. bolanderi cited only the _ following: "On rocks 
(‘metamorphic sandstone’) in Marin County, California, Mr. H. N. Bolander". 


Lectotype: Bolander 29--thalli on the card (FH) (Fig. 5). 


The iectotype of L. bolanderi consists of detached branches glued on a card, 
annotated by Tuckerman as “Lecanora (Cladodium) Bolanderi, Tuck., California, H. N. 
Bolander", and by Bolander as "No. 29. On metamorphic sandstone, Marin County". 
This specimen was chosen as lectotype of L. bolanderi for several reasons: 1) the 
thalli fit within the protologue, are from Marin County, and closely resemble other 
Bolander specimens labelled as L. bolanderi; 2) the card is clearly annotated by 
Tuckerman (although he referred to Cladodium instead of Squamaria as in the 
protologue); and 3) assuming the numbers are in chronological order, Bolander 29 
could have been collected (and seen by Tuckerman) before Bolander’s numbered 
collections of L. phryganitis (Bolander 63) and L. thamnitis (Bolander 125 and 155) 
and before publication of L. bolanderi. The material contains usnic acid and unknown 
CBI-1. Unfortunately, the Bolander 29 material is not of good quality: the thalli are 
moldy, do not show the growth form well, and bear only immature apothecia. 


Other Lecanora bolanderi specimens. 


Other Bolander collections of L. bolanderi from Marin Co. are similar to the 
lectotype in morphology and chemistry (usnic acid or isousnic acid; usually with 
unknown CBL-1) but are better specimens, and may be useful in understanding 
Tuckerman’s concepts. Some of these specimens may well be parts of the same 
original collection, but with neither collecting numbers nor annotations cannot be 
called isolectotypes. These critical specimens include : Bolander, s.n., Marin Co., no 
other data (FH; UC 666809: both labelled as "type" by the curators at FH) and 
Bolander, s.n., no other data (FH: “ex Willey herb."). Another specimen, Bolander, 
s.n., Tomales Bay, near oyna [sic]", undated (FH) (Fig. 6), is labelled “original 
specimens!", an expression that might have been used by H. Willey (at US) but not 
by Tuckerman (W. Culberson, pers. comm., 1988). This specimen, as is true for 
another one from this locality, Bolander, s.n., °/4 64 (sic; = unspecified date, March, 
1864) (FH), may have been collected too late to have been used in the protologue. 


Specimens excluded as types of L. bolanderi. 


The packet labelled Bolander 29 at FH contains, in addition to the material 
chosen as the lectotype of L. bolanderi, an unannotated packet bearing thalli closely 
matching my lectotype of L. phryganitis. The presence of unlabelled packets such as 
this one can be explained by the following note by Tuckerman (dated 19 March 
1876), from the archives at FH (Cacavio, pers. comm., 1988), which refers to his 
herbarium as a whole: 


"It does not follow, because specimens, evidently misdetermined, are placed in 
a certain wrapper, that | consider them to belong to the species for which that 
wrapper is designed, but only that a certain degree of resemblance leads me to 
place such specimens with such species till | examine it. It is just to me that 
this should be recognized; and throughout my collection." 


This unlabelled packet associated with Bolander 29 may be the source of Mrs. 
Riddle’s incorrect labelling of Relig. Tuck. Exs. 108 (Bolander, s.n., undated, Marin 
Co.) as "authentic" material of L. bolanderi. All specimens of this exsiccat seen are 
L. phryganitis. The one at FH contains an undated note (by Bolander?) saying: "100, 
California, Bolander, Placodium novum" (the "100" may be Bolander’s collection 


710 


number; Tuckerman would not have referred to "Placodium", which he applied only to 
species now in Caloplaca). 


Typification of L. thamnitis 


The original collection of L. thamnitis was cited in the protologue as follows: 
"Sandstone rocks, with the last [L. phryganitis], Mr. Bolander.” In the present study, 
no evidence was found that L. thamnitis and L. phryganitis were collected by 
Bolander from the same locality. Tuckerman (1882) mentioned two localities for L. 
thamnitis: 1) Oakland Hills (where apparently no specimens of fruticose Lecanoras 
have been collected since the time of Bolander, and suitable habitats may now be 
unavailable or inaccessible due to the build-up of the city), and 2) San Bruno 
Mountain (inferred by Herre [1945] to be the type locality; my search for L. thamnitis 


there turned up L. phryganitis!). 


In the absence of other evidence, one of Bolander’s Oakland Hills collections 
(discussed below) was chosen as the lectotype of L. thamnitis for the following 
reasons: 1) all of these collections are very similar to each other in morphology and 
chemistry (usnic acid and unknown CBL-1), are of good quality, and (except for for 
peltate thalli in some an show the distinctive characteristics of the taxon as 
expressed in the protologue; 2) several are clearly annotated by Tuckerman as L. 
thamnitis; and 3) the locality was cited by Tuckerman (1882). 


Lectotype: Relig. Tuck. Exs. 116 (FH) (Fig. 9). 


Of the Oakland Hills collections, the Relig. Tuck. Exs. 116, labelled by Mrs. 
Riddle as "determined by Tuckerman” and "type", was chosen as the lectotype of L. 
thamnitis for several reasons: 1) the specimen of this exsiccat at FH, labelled 
"isotype" by the curators, is annotated (by Tuckerman?): "L. thamnitis, Stock to 
publish, Bolander, California"; 2) although not showing the habit in situ, the thalli are 
of good quality and are only indistinctly peltate; and 3) choice of this specimen at FH 
as lectotype avoids further confusion, especially since Relig. Tuck. 116 (GZU) was 
the basis for the genus Cladidium Hafellner (see above). 


Other critical L. thamnitis specimens. 


One collection from Oakland Hills (Bolander, s.n.) shows especially well the 
effuse growth form of the attached thalli (Fig. 10). 


Another Oakland Hills collection, Bolander 125, undated (FH) consists of three 
parts, all annotated by Tuckerman as L. thamnitis: 1) a card with effuse thalli 
attached to rocks, with Bolander’s annotation: "125. This [is] the one | had in my 
former letters referred to. | send it now in all the various stages of growth."; 2) a 
similar set of effuse thalli, on a rock annotated by Tuckerman as: "“Lecanora 
thamnitis T. in litt. ad Bolander" and by Bolander as: "This | found now quite plenty 
on rocks, Oakland Hills"; and 3) a group of clearly peltate ull (Fig. 11) annotated 
by Bolander as: "On rocks (metamorphic sandstone), Oakland Hills. Is this Lecanora 
B.2?". The annotations, and the peltate thalli (neither mentioned nor explicitly 
excluded in the protologue) are interesting in view of Tuckerman’s changing concepts 
of the taxa (See above). 


The only known Bolander collection of L. thamnitis from San Bruno Mountain, 
Bolander 155 (FH--labelled as "co-type", i.e., syntype), is annotated by Tuckerman as: 
"California, Bolander 1865, L. thamnitis T." and by Bolander as: "155 L. thamnitis T. 
San Bruno Mts (sic). | send this also from this locality.". These rather fragmentary 
thalli (Fig. 12), which were collected prior to publication of L. thamnitis (but apparently 
sent to Tuckerman at the same time as Bolander 125), are somewhat similar to the 
lectotype of L. bolanderi, and unlike the other material of L. thamnitis, they partly 
contain isousnic acid instead of usnic acid. 


ACKNOWLEDGEMENTS 
This study was supported in part by a grant from the National Science 


Foundation (#BSR-851156) and in part by awards from the Smithsonian Institution, 
the Arizona Federation of Garden Clubs and the Graduate Student Association of 


711 


Arizona State University. | thank the curators of the various herbaria (ASU, CANL, 
COLO, DAV, FH, G, GZU, H, IRVC, LAM, M, SFSU, UC, UPS, US, WU) for loans of 
specimens, and the A.S.U. Media Production staff for photography. Special thanks 
also go to to C. Bratt, |. M. Brodo, G. Cacavio, J. Elix, J. Hafellner, M. E. Hale, Jr., 
S. Hammer, C. Leuckert, T. H. Nash Ill, D. Pfister, D. Pinkava, J. Poelt, and G. F. 
Ryan. 


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Culberson, C. F., W. C. Culberson and A. Johnson. 1981. A standardized TLC 
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7l2 


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Zahlbruckner, A. 1921-1940. Catalogus Lichenum Universalis, I--IX. Borntraeger, 
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Pflanzenfamilien 8, B. Spezieller Teil. 


ERRATA 


ors 


AUTHOR INDEX, VOLUME THIRTY-FOUR 


Ammirati, Joseph F., Dermocybe, subgenus Dermocybe, section Sanguineae in 
northern California 21-36 

Anon., Harry D. Thiers: Reminiscences about a teacher and friend 3-16 

Anon., Master’s degree students of Harry D. Thiers and thesis titles 267-268 

Anon., Remembering the morel grower: Ron Ower, 1939-1986 17-20 

Arambarri, Angélica M., and Marta N. Cabello, A numerical taxonomic study of 
some phialidic genera of hyphomycetes: cluster analysis 679-696 

Balazuc, Jean, see Tavares & Balazuc 

Baltzo, Doris E., Lichens of Mount Diablo State Park, Contra Costa County, Cali- 
fornia 37-46 

Baroni, Timothy J., and David L. Largent, The genus Rhodocybe: new 
combinations and a revised key to section Rhodophana in North America 47-53 

Barr, Margaret E., The genus Chaetomastia (Dacampiaceae) in North America 507- 
SLD 

Barr, Margaret E., The genus Dothidotthia (Botryosphaeriaceae) in North America 
517-526 

Borse, B. D., and K. D. Hyde, Marine fungi from India. III. Acrocordiopsis patilii 
gen. et sp. nov. from mangrove wood 535-340 

Brenneman, Timothy B., see Hanlin & al. 

Brusse, Franklin A., Two new species of Parmelia (Parmeliaceae, Lichenes), further 
new combinations and notes, and additional new lichen records from southernAfrica 
399-406 

Burge, Harriet A., Marion E. Hoyer, William R. Solomon, Emory G. 
Simmons, and Janet Gallup, Quality control factors for Alternaria allergens 
55-63 

Cabello, Marta N., see Arambarri & Cabello 

Cabral, D., see Romero & al. 

Calhoun, Cornelia J., Thaxterogaster thiersii: a new secotioid species from Cali- 
fornia 65-70 

Cavelo, Susana, and Laura Lorenzo, Noteworthy corticolous lichens in 
Nothofagus forests, north-western Patagonia 655-665 

Cole, R. J., see Wicklow & al. 

Constantinescu, O., and S. Ryman, A new Ophiostoma on polypores 637-642 

Desjardin, Dennis E., Species named for Harry D. Thiers 276 

Desjardin, Dennis E., and Ronald H. Petersen, Studies on Marasmius from 
eastern North America. II. New species 71-92 

Flock, JoAnn W., Lithographa, a lichen genus new to continental North America 
643-645 

Foos, K. Michael, and Judith A. Royer, A survey of Pilobolus from 
Yellowstone National Park 395-397 

Gallup, Janet, see Burge & al. 

Glawe, Dean A., and John P. Jones, The anamorphs of Diatrypella prominens 
and Eutypella sabalina 277-281 

Hale, Mason E., New species in the lichen genus Xanthoparmelia (Ascomycotina: 
Parmeliaceae) 541-564 

Halling, Roy E., A synopsis of Colombian boletes 93-113 

Halling, Roy E., see Thiers and Halling 

Halling, Roy E., and Barbara M. Thiers, Preface [to the Harry D. Thiers Fest- 
schrift] 2 

Hammer, Samuel, Cladonia thiersii: a new lichen from California 115-118 


See ERRATA 
714 


Hanlin, Richard T., Mei-Lee Wu, and Timothy B. Brenneman, The 
occurrence of Tuber texense in Georgia 387-394 

Heykoop, Michel, see Moreno & al. 

Honrubia, M., see Roldan & Honrubia 

Hoyer, Marion E., see Burge & al. 

Humber, Richard A., Synopsis of a revised classification for the Entomophthorales 
(Zygomycotina) 441-460 

Hyde, K. D., see Borse & Hyde 

Hyde, K. D., and E. B. G. Jones, Marine fungi from Seychelles. VIII. 
Rhizophila marina, a new ascomycete from mangrove prop roots 527-533 

Illana, Carlos, see Moreno & al. 

Jones, E. B. G., see Hyde & Jones 

Jones, John P., see Glawe & Jones 

Kerrigan, Richard W., Studies in Agaricus IV: new species from Colorado 119- 
128 

Korf, Richard P., see Zhuang & Korf 

Kuthubutheen, A. J., see Nawawi & Kuthubutheen 

Laferriére, Joseph E., Mnemonic three-letter acronyms for the names of fungal 
families 461-473 

Largent, David L, A new, lignicolous species of Entoloma (Entolomataceae, Agari- — 
cales) from California 129-131 

Largent, David L, see Baroni and Largent 

Lopez, S. E., see Romero & al. 

Lorenzo, Laura, see Cavelo & Lorenzo 

Mahu, Manuel, Pollution atmosphérique et lichens dans la ville de Santiago du Chile 
407-428 

Malloch, David, Notes on the genus Protubera 133-151 

McAlpin, Cesaria E., see Wicklow & al. 

Methven, Andrew S., Notes on Clavariadelphus. III. New and noteworthy species 
from North America 153-179 

Moreno, Gabriel, Carlos Ilana, and Michel Heykoop, Contribution to the study 
of the myxomycetes in Spain. I 623-635 

Nawawi, A., and A. J. Kuthubutheen, Canalisporium, a new genus of lignicolous 
hyphomycetes from Malaysia 475-487 

Nawawi, A., and A. J. Kuthubutheen, A new taxon in Colispora (Hyphomycetes) 
from Malaysia 497-501 

Nawawi, A., and A. J. Kuthubutheen, Quadricladium aquaticum gen. et sp. nov., 
an aquatic hyphomycete with tetraradiate conidia 489-495 

Nishida, Florence H., A key to the species of Inocybe in California 181-196 

Ovrebo, C. L., see Welden & Ovrebo 

Pande, Alaka, and V. G. Rao, Ascomycetes of western India XIII 503-505 

Petersen. Ronald H., see Desjardin and Petersen 

Reynolds, Don R., Foliicolous fungi 8: Capnodium in California 197-216 

Rodrigues, Katia F., Index to J. B. Ellis’ types of pyrenomycetes with amyloid ascal 
rings 577-599 

Rogers, Jack D., see San Martin Gonzalez & Rogers 

Roldan, A., A new addition to the genus Gorgomyces 381-385 

Roldan, A., and M. Honrubia, Varicosporium scoparium, a new staurosporous 
hyphomycete 375-379 

Romero, A. I., D. Cabral, and S. E. Lopez, Studies on xylophilous fungi from 
Argentina. IV. Anamorphs of Basidiomyceteson Eucalyptus viminalis (Myrtaceae) 
429-440 

Roquebert, Marie-France, see Wicklow & al. 


fat aes 


Royer, Judith A., see Foos & Royer 

Ryan, Bruce D., The genus Cladidium (lichenized Ascomycotina) 697-712 

Ryman, S., see Constantinescu & Ryman 

San Martin Gonzalez, Felipe, and Jack D. Rogers, A preliminary account of 
Xylaria of Mexico 283-373 

Schenck, Norman C., see Spain & al. 

Seidl, Michelle T., A new species of Gymnopilus from northern California 217-220 

Sieverding, Ewald, see Spain & al. 

Sigal, Lorene L., The lichens of serpentine rocks and soils in California 221-238 

Simmons, Emory G., see Burge & al. 

Soloman, William R., see Burge & al. 

Spain, Joyce Lance, Ewald Sieverding, and Norman C. Schenck, Gigaspora 
ramisporophora: anew species with novel sporophores from Brazil 667-677 

Sundberg, Walter J., Lepiota sensu lato in California. III. Species with a hymeniform 
pileipellis 239-248 

Tavares, Isabelle I., and Jean Balazuc, Sugiyamaemyces, a new genus of 
Laboulbeniales (Ascomycetes) on Clidicus (Scydmaenidae) 565-576 

Thiers, Barbara M., see Halling and Thiers 

Thiers, Barbara M., and Roy E. Halling, Type specimens of agarics, boletes and 
gasteromycetes in the San Francisco State University herbarium (SFSU) 269-276 

Tulloss, Rodham E., and Greg Wright, Amanita protecta — a new species from 
coastal southern California 615-622 

Vesonder, R. F., see Wicklow & al. 

Welden, A. L., and C. L. Ovrebo, Notes on tropical and warm temperate basidio- 
mycetes 601-614 

Wicklow, D. T., R. F. Vesonder, Cesaria E. McAlpin, R. J. Cole, and 
Marie-France Roquebert, Examination of Stilbothamnium togoense for Asper- 
gillus flavus group mycotoxins 249-252 

Wicklow-Howard, Marcia, The occurrence of vesicular-arbuscular mycorrhizae in 
burned areas of the Snake River Birds of Prey Area, Idaho 253-257 

Wong, George J., Compatibility and fruiting studies of an albino form of Auricularia 
cornea 259-266 

Wright, Greg, see Tulloss & Wright 

Wu, Mei-Ling, see Hanlin & al. 

Zhuang, Wen-ying, and Richard P. Korf, Notes on one lichenicolous and one 
fungicolous discomycete 647-653 


716 


INDEX TO FUNGOUS AND LICHEN TAXA, VOLUME THIRTY-FOUR 


This index contains the names of genera, infrageneric taxa, species, and infraspecific taxa. 
New names are in boldface, as are the page numbers on which such new taxa are 
proposed. New suprageneric names are in boldface CAPITALS. 


Acarospora 
chlorophana 38 
fuscata 38, 228, 234 
schleicheri 38, 228 
Acompsomyces 576 
stenichni 575 
Acremonium 
olidum 641 
Acrocordia 535, 538 
Acrocordiella 538 
Acrocordiopsis 535 , 538, 539 
patilii 535, 536, 539 
Acrogynomyces 575 
Acroscyphus 
sphaerophoroides 405 
Actinogyra 38 
polyrrhiza 45 
Aegerita 434 
Agaricus 11, 119, 120, 123, 127 
sect. Arvenses 119, 126 
sect. Spissicaules 119, 123 
amicosus 119, 120, 121, 123, 124, 269 
arorae 269 
augustus 126 
chlamydopus 119 
cinnamomeus 
var. semisanguineus 30 
cothumatus 119 
cuniculicola 119, 124-127, 269 
fuscovelatus 269 
haemorrhoidarius 123 
lanipes 123 
perobscurus 126, 269 
perrarus 126 
pilosporus 119 
praerimosus 119 
rubronanus 269 
rutilescens 119, 120, 123, 124 
sanguinea 28 
semotus 120 
sequoiae 269 
smithii 126, 127, 269 
solidipes 119 
sphaerosporus 119 
sterlingii 119 
summensis 269 
tabularis 119 
vinaceovirens 269 
Aigialus 
grandis 539 
parvus 539 
Aithaloderma 
rhododendri 214 
Aithalomyces 


rhododendri 214 
Alectoria 38 
oregana 38 
Aleuria 9 
Allescheriella 431,435  — 
cyanea 429, 434, 436 
Allescherina 
deusta 585 
eutypaeformis 584 
Alternaria 55-58, 60, 61 
alternata 55, 56, 58-61 
Amanita 9, 620 
sect. Amanita 620 
sect. Amidella 620 
sect. Vaginatae 616, 620, 621 
breckonii 269 
constricta 269, 621 
magniverrucata 270 
mortenii 620 
pachycolea 270 
pantherina 620 
peltigera 620 
phalloides 187, 620 
protecta 615 -617, 619-621 
pseudovaginata 621 
thiersii 276 
velosa 621 
Amaurochaete 
atra 623 
Amorphomyces 565 
Amphisphaeria 518 
aspera 519, 520, 578 
atrograna 578, 595 
bisphaerica 595 
confertissima 578 
deformis 578 
granulosa 578 
hypoxylon 578 
incrustans 579 
langloisii 579 
melantera 579 
nuda 579 
oronoensis 579, 595 
pilosella 579 
platani 579 
separans 579 
subiculosa 579 
Ancylistes 443, 455 
Antennariella 204, 205 
Antennatula 204, 205 
Antennularia 517 
Anthostoma 
acerinum 579 
ellisii 583 
flavoviride 579 


formosum 579 
microecium 580 
microsporum 583 
minor 581 
mortuosum 580, 595 
ontariense 580 
ostiolatum 581 
phaeospermum 584 
pholidigena 596 
picaceum 580, 596 
saprophilum 580 
stictoides 580 
Anthostomella 
albocincta 580 
brachystoma 580 
comicola 580 
eructans 580 
flavoviridis 579 
hypsophila 580 
inconspicua 596 
leucobasis 580, 595 
ludoviciana 580 
magnoliae 581 
mammoides 581 
melanosticta 581 
minor 581 
ostiolata 581 
pholidigena 581, 596 
picacea 596 
sabalensioides 581, 596 
suberumpens 581 
thyridioides 581 
tomicoides 595 
unguiculata 581 
xylostei 595 
Aposphaeria 509, 513,514 
Arcangeliella 
desjardinii 270 
parva 270 
saylorii 270 
variegata 270 
Arcyria 
cinerea 623 
incarnata 623 
obvelata 624 
pomiformis 624 
Armillaria 
olida 270 
Arthonia 704 
phaeobaea 702 
Arthrobotryum 
spongiosum 213, 214 
Asbolisia 205, 206 
ampullula 206 
Aspergillus 250, 251 
subg. Stilbothamnium 250 
flavus 249-251 
var. flavus 250 
var. parasiticus 250 
nomius 250 
ochraceus 249, 251 
parasiticus 251 


yAaws 


tamarii 249, 250 

togoensis 249, 250 
Aspicilia 42, 226, 235, 706 

caesiocinerea 228, 234-236 

calcarea 38, 236 

cinerea 38, 228 

crusii 235 

gibbosa 38 

laevata 38 

polychroma 235 

var. ochracea 235, 236 

serpentinicola 235, 236 
Asterostroma 608 
Astrocystis 

mirabilis 593 
Astrosphaeriella 539 
Auricularia 259-261, 265 

albicans 259, 260 

auricula-judae 260 

comea 259-262, 264, 265 

delicata 260 

eburnea 260 

fuscosuccinea 260 

leucochroma 259, 260 

mesenterica 260, 634 

polytricha 260, 265 

f. leucochroma 260 

tenuis 260, 265 
Austroboletus 93 

subvirens 94, 95 


Badhamia 
dubia 623, 624 
foliicola 625 
nitens 626 
panicea 625 
utricularis 625 
versicolor 625, 626 
Bahusutrabeeja 679, 684, 691, 695 
angularis 680, 691 
dwaya 680, 691 
Ballocephala 443, 456, 457 
Basidiobolus 443, 458 
Batkoa 441, 443, 445, 446, 447 
apiculata 446 , 447 
dysderci 446 
gigantea 446 
major 446 , 447 
papillata 446 , 447 
Berkleasmium 475, 477 
caribense 475, 477, 479 
concinnum 477 
pulchrum 475, 481, 484 
Bipolaris 251 
Biscogniauxia 
pezizoides 591 
Boletellus 93 
ananas 93, 95 
russellii 95 
Boletinellus 105 
Boletus 93 


718 


abieticola 270 var. paraguayense 655, 658, 660, 663 
amygdalinus 271 Caloplaca 39, 228, 233, 698, 710 
amyloideus 270 sect. Gasparrinia 39 
ananas 95 sect. Thamnoma 698 
atkinsonianus 95-98 bolacina 38 
barrowsii 270 bolanderi 227, 228, 233 
calopus 12 cerina 419, 422, 426, 427 
chrysenteron 102 chrysophthalma 38 
citriniporus 270 coralloides 699, 702 
coccyginus 270 decipiens 38 
dryophilus 270 elegans 39 
edulis 12 ferruginea 38 
fibrillosus 270 festiva 233 
fuligineotomentosus 93, 98 laeta 38, 228, 233 
haematinus 270 murorum 39 
mendocinensis 271 rosei 702 
morrisii 108 squamosa 228, 234 
mottiae 271 stanfordensis 38 
mottii 271 Calospora 
obsonium 97 clavispora 509 
orovillus 271 rhoina 584 
orquidianus 93, 98-100 Camarops 
piperatus 93 microspora 583 
pseudorubinellus 100, 101 ohiensis 588, 590 
pulcherrimus 271 polyspermum 589 
pulverulentus 100 tubulina 588 
puniceus 271 Canalisporium 475, 477, 486 
roseibrunneus 271 caribense 476, 479 
rubripes 12, 271 elegans 475, 483, 484, 485 
russellii 95 pulchrum 479-481, 482 
silvaticus 271 Candelaria 
smithii 271 concolor 39, 228 
spadiceus Candelariella 234 
var. rufobrunneus 271 vitellina 39, 228, 234, 235, 420, 422, 
subsolitarius 98 426, 427 
subtomentosus 101 Capnodium 197, 202-204, 207, 208 
truncatus 102, 103 araucariae 209, 212 
viridiflavus 98 australe 212 
Bolinia baccharidis 198, 208 
tubulina 588, 590 Caespitosum 209 
Botryobasidium citri 207, 210 
vagum 436 coffeae 210 
Botryosphaeria 517 dematium 197, 198, 200-203, 208, 21 
obtusa 517, 525 ellisii 202, 212 
Brachiosphaera 489 elongatum 209 
tropicalis 494 footii 210, 212 
Bryoria 38 ; heteromeles 206, 207, 212-214 
oregana 38 pelliculosum 202, 212 
Buellia 702 rhamni 205, 206, 213 
badia 228 salicinum 202, 203, 208, 213 
fuscula 419, 422, 426, 427 spongiosum 204, 205, 213, 214 
punctata 38, 228 tuba 211, 213 
spuria 228 walteri 202, 203, 212 
stellulata 228 Capnophaeum 
vilis 228 spongiosum 204, 213 
Burgoa 434 Capnophialophora 205 
Byssosphaeria Capronia 507 
macouniana 593 Caryosporella 
rhodomphala 579 rhizophorae 539 
Catapyrenium 
Calicium cinereum 228, 233, 234 


leucochlorum lachneum 39, 228 


Catillaria 
lenticularis 228 
Ceratocystis 640 
perparvispora 641, 642 
Ceriomyces 
atkinsonianus 95 
Ceriospora 
alabamiensis 581 
Cetraria 39 
chlorophylla 45 
merrillii 45 
orbata 45 
Chaetasbolisia 213 
californiana 207 
Chaetomastia 507, 508, 510 
aculeata 507 : 
clavispora 507, 509,510 
cucurbitarioides 507 
equiseti 507, 509, 510 
hirtula 507-510, 512 
hispidula 507 
juniperina 507 
phaeospora 507, 509, 510, 512 
pilosella 507 
sambuci 507, 509, 510, 513 
sambucina 507, 509, 510, 513 
typhicola 507, 509, 510, 514 
Chaetomium 251 
Chaetopsina 679, 684, 691, 694, 696 
auburnensis 680, 694 
catenulata 680, 694 
fulva 680, 694 
penicilliata 680, 694, 695 
polyblastia 680, 694 
splendida 680, 694 
unilateralis 680, 694 
Chaetopsis 679, 680, 684, 691, 696 
cubensis 683 
grisea 680, 694 
romantica 680, 694 
Chaetosphaeria 
callimorpha 681 
dingleyae 679, 683, 695 
pulchriseta 682 
pulviscula 683, 695 
talbotii 682 
Chitonomyces 574 
Chloridium 684, 691, 692, 694, 695 
atanagildae 680, 691 
botryoideum 680, 694 
caudigerum 680, 694 
chlamydosporis 680 
clavaeforme 680 
codinaecoides 681 
cubense 681 
cylindrosporum 681 
lignicola 681 
matsushimae 681 
pachytrachelum 681 
paucisporum 681, 691, 692 
phaeosporum 681 
preussii 681 


719 


reniforme 681, 694 
smithiae 681 
transvaalense 681, 694 
virescens 681 
Cladidium 697-699, 704, 710 
bolanderi 697, 699, 701-705 
thamnitis 697, 699 
Cladoconidium 
articulatum 379 
Cladodium 698, 699, 709 
hosseanum 698 
Cladonia 39 
sect. Perviae 115, 118 
carassensis 116 
cervicomis 
ssp. verticillata 39 
chlorophaea 39 
coniocraea 228 
crispata 118 
fimbriata 39 
macilenta 39 
papillaria 706 
pityrea 39 
pyxidata 228 
ramulosa 39 
santensis 115, 118 
squamosa 
var. subsquamosa 115, 116, 118 
subcericomis 39 
subsubulata 115, 116 
thiersii 115 -118, 276 
verticillata 39 
Clavaria 
pistillaris 159, 160, 173 
var. americana 154 
var. unicolor 160 
unicolor 160 
Clavariadelphus 153, 157, 161, 162, 169 
subg. Clavariadelphus 168 
sect. Cantharellopsis 178 
sect. Clavariadelphus 157, 166, 168, 173, 
174 
americanus 153, 154, 155, 161, 162, 
165, 168 
caespitosus 153, 155, 162, 165, 166, 
174 
cokeri 161, 162 
fasciculatus 166, 174 
flavidus 153, 162, 165, 166, 168, 169 
flavo-immaturus 169 
ligula 153, 165, 169, 174 
occidentalis 153, 161, 165, 168, 169, 
173-175 
pallido-incarnatus 153, 171, 174, 
175, 178 
pistillaris 153, 158-162, 164, 166, 168, 
172-174 
var. americanus 153, 154, 159-161 
f. americanus 159 
sachalinensis 169, 174 
subfastigiatus 166, 174 


720 


truncatus 153, 161, 178 

unicolor 161, 162 

xanthocephalus 169 
Clitocybe 

foveolata 272 

gibba 

var. occidentalis 272 

stercoraria 272 

thiersii 272, 276 
Clitopilus 48 
Clypeosphaeria 

imperfecta 581 

minor 581 

sanguinea 581 

ulmicola 582 


Codinaea 679, 683, 684, 691, 692, 694, 695 


apicalis 679, 683, 695 
apiculata 681 
aristata 681 
australensis 684 
botulispora 681 
brevisetula 681 
coffeae 681 
cylindrospora 681, 694 
dimorpha 682 
elegantissima 679, 683, 695 
eucalypti 682 
filamentosa 679, 683, 695 
glauco-nigra 679, 683, 695 
heteroderae 682 
hughesii 682 
illinoensis 682 
intermedia 679, 683, 695 
longispora 682 
lunata 682 
lunulospora 682 
maharashtrensis 679, 683, 695 
matsushimae 682 
obesispora 682 
pakhalensis 684 
parva 682 
septata 682 
setosa 682 
tortuosa 684 
unisetula 682 
vulgaris 683 
Colispora 497 
curvata 497 , 499, 500 
elongata 497, 500, 501 
Collema 704 
furfuraceum 39 
nigrescens 39 
Colloderma 626 
Completoria 443, 454 
complens 441, 454 
COMPLETORIACEAE 453 
Conidiobolus 441, 443, 444, 447, 455, 456 
subg. Capillidium 443, 456 
subg. Conidiobolus 443 
subg. Delacroixia 443 
apiculatus 447 
major 447 


utriculosus 456 
Coniochaeta 
kellermannii 593 
xylarispora 596 
Coniothyrium 509 
sambuci 513 
Coreomyces 565 
Corethromyces 
cryptobii 575 
Corticomyces 429, 431, 432 
xenasmatoides 429, 43 1, 432 
Cortinarius 8, 9, 21 
califomica 24 
phoeniceus 
var. occidentalis 26 
purpurascens 190 
sanguineus 28 
var. sierraensis 33 
semisanguineus 30 
thiersii 271, 276 
velatus 271 
verrucisporus 271 
Craterium 
leucocephalum 626 
Cribraria 
vulgaris 626 
Cryptandromyces 575 
Cryptosphaeria 
fissicola 582, 595 
juglandina 582 
subcutanea 580 
Cryptovalsa 
citricola 585 
deusta 585 
eutypaeformis 584 
pustulata 586 
sassafras 586 
Cucullospora 
mangrovei 539 
Cylindrotrichum 680, 684, 691, 694, 695 
clavatum 681 
curvatum 681, 691, 692 
ellisii 681 
excentricum 681 
fasciculatum 681, 694 
gorii 683-684 
helisciforme 684 
hennebertii 681 
oblongisporum 681 
oligospermum 681 
probosciophorum 681 
proliferum 681 
triseptatum 681, 694 
zygnoellae 681 
Cypheliopsis 
bolanderi 45 
Cytospora 
maclurae 597 


Dactylelia 497 
aquatica 497 
submersa 497 


fp al 


Deflexula 601 olivacea 584, 586 
nivea 602 phaeosperma 584 
subsimplex 601, 602, 614 prominens 583, 584 

Dendrospora 379 pustulans 584 
erecta 379 quercina 
polymorpha 379 var. lignicola 584 

Dermatocarpon 226, 234 radiata 584 
miniatum 39, 234 rhuina 584 

var. complicatum 228 sphaerospora 584 
reticulatum 39 texensis 584 
Dermocybe 9, 21 tiliacea 584 
subg. Dermocybe 21 tremellophora 585 
sect. Sanguineae 21, 22, 25-27, 30 trifida 585 
californica 21, 23, 24, 25, 30 tumida 585 
cinnabarina 26 vitis 585, 586 
phoenicea 32 Diatrypella 
var. occidentalis 21, 23, 26-28, 32 citricola 585 
var. phoenicea 28 comptoniae 582 
sanguinea 21-23, 28, 30, 35 decipiens 585 
var. vitiosa 35 demetrionis 585 
semisanguinea 21, 22, 28, 30, 32 deusta 585 
sierraensis 21, 23, 30, 33, 35, 271 fraxini 585 

Destuntzia herbacea 585 
saylorii 271 hysterioides 585 

Diacheopsis 626 irregularis 583, 585 
depressa 626 missouriensis 585 
nannengae 623, 626, 627 obscurata 586 
picninica 626 olivacea 584, 586 

Diaporthe 650, 652 paupera 586 
acervata 582 populi 586 
rhoina 584 prominens 277, 278 
tiliacea 585 prunicola 586 

Diatrype 641 pulcherrima 586 
acervata 582 pustulata 586 
aethiops 582 ramularis 586 
albopruinosa sassafras 586 

var. salicina 597 subfulva 586 
americana 582 tocciaeana 
bullata 583 var. subeffusa 586 
celastrina 582 verrucaeformis 586 
collariata 582 vetusta 586 
comptoniae 582 Vitis 586 
comuta 582 xanthostroma 586 
disciformis Dibotryon 517 
var. americana 582 symphoricarpi 517, 519 
var. magnoliae 585 Dichocantharellus 608 
fibritecta 582 Dichopleuropus 608 
flavovirens 583 Dichostereum 608 
hochelagae 583 Diclonomyces 575 
hullensis 583 Dictyochaeta 679, 681, 682, 684, 691-696 
infuscans 583 abnormis 681 
irregularis 583, 585 apiculata 681 
lateritia 583 aristata 681 
linearis 583 assamica 681 


botulispora 681 


maclurae 583 brevisetula 681 
macounii 583 clavulata 681 

megastoma 583 coffeae 681 

microspora 583 cylindrospora 681 , 694 
microstega 584 dimorpha 681 , 693 
minima 584 eucalypti 682 , 693 


nigerrima 584 


fertilis 682, 692 


722 


fuegiana 682 

gamundii 682 

heteroderae 682 

hughesii 682 

illinoensis 682 , 693 

longispora 682 

lunata 682 

lunulospora 682 

matsushimae 682 

menisporoides 679, 682, 695 

novae-guineensis 682 

obesispora 682 

parva 682 

querma 682 

septata 682 

setosa 682 

simplex 682 

tilikfrei 682 

triseptata 682 

unisetula 682 

vulgaris 682 
Diderma 

chondrioderma 623, 628, 629 

spumarioides 629 
Didymella 503 

capparidis 503,505 

stromatica 503 
Didymium 

bahiense 629, 632 

difforme 630 

karstensii 623, 629, 630 

marineri 623, 630-632 

melanospermum 632 

minus 632 

squamulosum 632 

vaccinum 633 
Didymosphaeria 518 

accedens 524, 525 

celtidis 520 

vagans 524, 525 
Dimelaena 

oreina 39 

radiata 228 
Dimerium 

hypoxylon 579 
Diplodia 517, 525 
Diploicia 

canescens 648 
Diplopodomyces 574 
Diploschistes 

actinostomus 39 

muscorum 39 

scruposus 39, 229 
Diplotomma 

alboatrum 229 
Distolomyces 574 
Dothidea 

clavispora 509 

insculpta 523 


Dothidotthia 517, 518, 520, 523, 524 


aspera 517-519, 520, 523 
celtidis 517,519, 520 


diapensiae 517, 518, 520, 522 
fruticola 517, 519, 520, 522, 523 
lasioderma 517, 518, 520, 523 
quercicola 517, 519, 520, 524 
ramulicola 517-520, 522, 524, 525 
symphoricarpi 518-520 

Dreschlera 251 


Echinochaete 601 
brachyporus 601, 602 
Elaphomyces 387 
Elasmomyces - 
stipitatus 272 
Elletevera 587 
Ellisia 578 
Ellisiella 578 
Ellisiellina 578 
Ellisiodothis 578 
Ellisiopsis 578 
Emericella 251 
Empusa 449 
americana 450 
apiculata 446, 447 
var. major 446 
dipterigena 452 
dysderci 446 
echinospora 452 
major 446, 447 
montana 451 
papillata 446, 447 
sciarae 451 
virescens 450 
Endogone 
heterogama 675 
Engizostoma 
aesculinum 587 
alpinum 587 
amorphae 587 
canodisca 597 
capillatum 597 
carpinicola 587 
coryli 587 
densisimmum 587 
deustum 597 
echinatum 587 
ellisii 597 
fissicola 595 
herbicola 588 
juglandinum 582 
lutescens 597 
maclurae 597 
rugiellum 598 
venusta 598 
Enteridium 
lycoperdon 633 
Entosordaria 
sabalensioides 596 
Entoloma 48, 129 
sect. Turfosa 131 
eulividum 130 
lignicola 129 , 130 
niphoides 130 


prunuloides 130 
rhodopolium 130 
sinuatum 130 
speculum 130 
trachyosporum 47-49 

var. griseoviolaceum 48, 49 

var. purpureoviolaceum 48, 49 

var. trachyosporum 48 
Entomophaga 441, 443, 445, 447, 448 
subg. Lichia 446 

aulicae 447 

batkoi 447 

calopteni 447 
conglomerata 447 

domestica 446, 447 

grylli 447, 448 

kansana 448 

limoniae 446, 447 

saccharina 448 

tabanivora 448 

tenthredinis 448 

tipulae 448 


Entomophthora 443, 445, 448, 449, 451 


anglica 453 
aphidis 451 
blunckii 
brahminae 452 
brevinucleata 448 
calliphorae 452 
calopteni 447 
crustosa 451 
culicis 448 
delphacis 452 
erupta 448 
gigantea 446 
gloeospora 453 
helvetica 448 
israelensis 448 
muscae 448 
phalangicida 453 
planchoniana 448 
scatophagae 448 
schizophorae 448 
thaxteri 448 
thripidum 448 
tipulae 448 
trinucleata 448 
weberi 448 
Entosordaria 
albocincta 580 
comicola 580 
magnoliae 581 
sabalensioides 596 
Erynia 441-445, 448-451, 453 
subg. Furia 450 
subg. Neopandora 451 
anhuiensis 453 
aquatica 449 
bullata 452 
conica 449 
creatonoti 450 


curvispora 449 
dacnusae 452 
delpiniana 449 
ellisiana 451 
erinacea 449 
formicae 452 
gracilis 449 
henrici 449 
ithacensis 451 
kondoiensis 453 
neoaphidis 452 
neopyralidarum 451 
nouryi 453 
ovispora 449, 450 
pieris 451 
plecopteri 449 
rhizospora 449 
sepulchralis 449 
suturalis 453 
variabilis 449 
zabri 451 
Eryniopsis 443 
Euantennaria 205 
rhododendri 204 
Euopsis 
pulvinata 229 
Eutypa 
echinata 587 . 
flavovirens 583 
heteracantha 587 
ontariensis 580 
Eutypella 
aequilinearis 584 
aesculina 587 
alpina 587 
amorphae 587 
canodisca 587, 597 
capillata 587, 597 
carpinicola 587 
cerviculata 583, 587 
collariata 582 
coryli 587 
densissima 587 
deusta 587, 597 
exigua 587 
fici 587 
goniostoma 588 
herbicola 588 
juglandicola 
var. juglandina 597 
juglandina 588, 597 
leprosa 582-584 
lutescens 588, 597 
maclurae 588, 597 
microcarpa 588, 597 
populi 588 
quadrifida 583 
rugiella 588, 598 
sabalina 277, 279, 280 
sarcobati 588 
scoparia 587, 588, 597 
tetraploa 587 


723 


724 


tiliae 588 
tumida 585 
venusta 588, 598 
Evernia 547 
prunastri 39 
var. sorediifera 39 


Fenestella 
princeps 525 
Fibulochlamys 429, 430, 431 
ferruginosa 429, 430, 432 
Flabellospora 489, 494 
crassa 494 
irregularis 494 
Flammula 
purpurata 219 
Flavoparmelia 43 
caperata 39, 547 
Flavopunctelia 43 
flaventior 39, 44, 229 
Fomitopsis 
pinicola 637, 639-641 
Fuckelia 
phaeospermum 584 
Fuligo 
septica 633 
Fumago 
fagi 212 
Fumagospora 202 
capnodioides 209 
cistophila 209 
gaultheriae 208, 209, 212 
Furia 441, 443, 450,451 
americana 450 
creatonoti 450 
crustosa 451 
ellisiana 451 
ithacensis 451 
montana 451 
neopyralidarum 451 
pieris 451 
sciarae 451 
virescens 450 
vomitoriae 451 
zabri 451 


Gasparrinia 
microphylla 420, 422, 426, 427 
Gastroboletus 
amyloideus 272 
citrinobrunneus 272 
suilloides 272 
turbinatus 12 
xerocomoides 272 
Gibbera 517 
andersonii 519, 520 
atrograna 595 
confertissima 578 
pilosella 579 
symphoricarpi 519 
Gibberidea 
symphoricarpi 523, 524 


Gigaspora 255, 668, 675, 676 
decipiens 674, 676 
margarita 675 


ramisporophora 667, 668, 669, 671- 


675 
Glomerella 531, 532 


Glomerulomyces 429, 432, 434 


fibulosus 429, 432 
Glomus 676 

microcarpus 255 
Gomphillus 

calicioides 405 
Gomphus 601 

cavipes 601, 604-606, 614 

subclavaeformis 605 

thiersii 272, 276 


Gonytrichum 679, 684, 691, 695, 696 


caesium 683 
chlamydosporium 683 
indicum 684 
macrocladum 683 
mirabile 683 
ypsilosporum 683 
Gorgomyces 385 
honrubiae 381 -385 
hungaricus 385 
Griphosphaerioma 
kansensis 520 
Gymnopilus 217, 219 
calobasis 219 
janthinosarx 219 
luteocameus 272 
luteofolius 219 
obscurus 272 
parvisquamulosus 272 
subtropicus 272 
thiersti 217 -219, 272, 276 
viridans 219 
Gymnopus 
aurantiacus 80 
Gyrodon 93 
exiguus 102, 104, 105 
metulioides 105 
monticola 93, 104-105 
Gyrodontium 601, 606 


versicolor 601, 605, 606, 610, 614 


Haematomma 

hilare 655, 659, 660, 663, 664 
Halosarpheia 

ratnaginiensis 539 
Haplotrichum 

curtisii 429, 436, 439 

gracile 429, 436, 439 
Hebeloma 187 
Heliscus 684 
Hendersoniella 199, 202 
Heptameria 

clavicarpa 509 
Hercospora 

tiliacea 585 
Herpomyces 565 


Herpotrichia 524 
rhodosticta 579 
separans 579 
symphoricarpi 523 

Herpotrichiella 
pilosella 507 

Hormonema 
prunorum 206 

Hydnellum 
pineticola 273 

Hygrophorus 
albinellus 273 
perfumus 273 
pyrophilus 273 
subellenae 273 

Hypocenomyces 42 
scalaris 39 

Hypocrea 
pulvinata 641 

Hypogymnia 
enteromorpha 39 
imshaugii 39 
pulverata 655, 659, 660, 664 
subphysodes 405 

var. austerodioides 405 
tubulosa 39 

Hypoxylon 
albocinctum 588 
albolanatum 592 
albostictum 591 
atrorufum 588 
atroviride 588 
bicolor 588 
bicoloratum 589 
bipapillatum 591 
californicum 589 
cinereum 589 
cohaerens 588 
commutatum 

subsp. holwayanum 589 
cylindrophorum 589 
discoideum 589 
fibuliforme 589 
geasteroides 593 
giganteum 593 
gigasporum 593 
glandiforme 593 
haematostroma 589 
holwayii 589 
hypophlaeum 589 
investiens 589 

f. bakeri 592 
langloisii 593 
limoniisporum 593 
lucidum 589 
mammatum 589 
melanaspis 592 
morgani 589 
multiforme 

var. effusum 589 
nicaraguense 590 


725 


notatum 590 
nuttallii 590 
occidentale 590 
ohiense 590 
pallidum 590 
papillatum 590 
piceum 590 
platystomum 590 
poliosum 594 
pulchrum 594 
rubiginosum 588-591 


_ rubrostromaticum 589 


sclerophaeum 590 
serpens 591 
stygium 590, 591 
subchlorinum 590 
subluteum 591 
thouarsianum 590 
tinctor 591 
truncatum 591 
vernicosum 591 


Hysterium 


clavisporum 509 


Immothia 


hypoxylon 579 


Inocybe 181-183, 192 


subg. Inocybe 183, 185, 190 
subg. Inosperma 183, 184 
subg. Mallocybe 183, 184 
sect. Cervicolores 183, 184 
sect. Inocybe 183, 190 
sect. Inocybium 183, 185 
sect. Rimosae 183, 184 
subsect. Acroconiatae 183, 187 
subsect. Cortinatae 184, 191 
subsect. Holoconiatae 183, 185 
subsect. Marginatae 184, 190 
acystidiosa 194 
adaequata 184, 192, 193 
agardhii 184, 192 
agglutinata 192 
albodisca 190, 194 
amblyspora 186, 192 
anomala 192 
auricoma 188, 192 
bakeri 186, 188, 193 
bresadolae 190, 194 
brunnescens 185, 193 
bulbosa 193 
calamistrata 184, 193 
californica 194 
chelanensis 192, 194 
chrysocephala 186 
cincinnatula 193 
cinnamomea 189, 193 
corydalina 193 

var. corydalina 188 
decipiens 191, 194 
decipientoides 194 
dulcamara 184, 193 
fallax 194 


726 


fastigiata 193 
fastigiella 185, 193 
flocculosa 190, 193 
fraudans 182, 188, 193 
fuscodisca 187, 193 
geophyila 182, 193 

var. geophylla 187 

var. lilacina 187, 193 

f. perplexa 193 
godeyi 185, 193 
hemileuca 189 
hirsuta 

var. maxima 193 
hirtella 193 

var. hirtella 185 
insinuata 187, 193 
jurana 193 
kauffmanii 187, 193 
lacera 183, 189, 193 
laetior 186, 193 
lanuginosa 191, 194 
leiocephala 186, 193 


leptocystis 182, 189, 192, 193 


leptophyila 191, 194 
lilacina 193 
longicystis 192, 194 
menthi-gustans 187 
mixtilis 182, 191, 194 
muricellata 185, 186, 193 
nigrescens 191, 194 
oblectabilis 190, 194 
Olida 190, 194 
olympiana 193 
petiginosa 192, 194 
phaeocomis 

var. major 188, 193 
phaeodisca 189, 193 
phaeoleuca 187, 193 
pudica 193 | 
pusio 188, 193 
pyriodora 193 
quietiodor 185, 193 
retipes 193 
rimosa 182, 185, 193 
serotina 193 
sororia 193 
splendens 187, 193 
subbrunnea 193 
subdestricta 189, 193 
subochracea 188, 193 
umbratica 190, 194 
vaccina 186, 193 
variabillima 182, 192, 194 
vinosistipitata 185, 193 
whitei 192, 193 

f. whitei 187 
xanthomelas 191, 194 


Julella 505 
multiloculata 503, 505 


Karstenula 508 


Keissleriella 
blepharospora 532 
Kionochaeta 679, 684, 691, 694, 696 
aristata 683, 694 
ivoriensis 683, 694 
keniensis 683, 694 
malaysiana 683, 694 
nanophora 683, 694 
pughii 683, 694 
ramifera 683, 694 
spissa 683, 694 
virtuosa 683, 694 
Kirschteiniella 
applanata 579 
Kirschteiniothelia 
aethiops 579 
Kobayasia 134 
nipponica 134, 143 
Koerberia 
biformis 42 
Kommamyce 
lateritia 591 
Kretzschmaria 
clavus 591 
pusilla 591 
spinifera 591 
turbinata 592 
Kruphaiomyces 574 


Laboulbenia 575 
Laccaria 
laccata 7 
Lachnocladium 601, 608 
schweinfurthiana 601, 606 
Lactarius 
argillaceifolius 
var. megacarpus 273 
californiensis 273 
cocosiolens 273 
subdulcis 13 
thiersii 276 
Lamia 449 
Lasiosphaeria 507 
Lasiosphaeris 507 
Lecania 706 
chilena 420, 422, 426, 427 
Lecanora 42, 229, 647, 697-699, 706, 707 
subg. Placodium 699 
sect. Cladodium 698, 699 
sect. Endochloris 699 
sect. Squamaria 709 
aipospila 706 
argopholis 227, 229 
atra 233 
bolanderi 697-699, 702-704, 706, 707, 
709, 710 
bolcana 42 
caesiorubella 
ssp. merrillii 42 
carpinea 647 
chlorophana 38 


cinerea 38 

dispersa 420, 422, 426, 427 

effusa 647, 648, 650 

fruticulosa 706 

garovaglii 42 

gibbosula 38 

hagenii 42 

laevata 38 

mellea 42 

muralis 42, 234, 702 
var. diffracta 42 
var. versicolor 42 

pacifica 42 

pallida 42 


phryganitis 697-699, 701, 702, 704, 706- 


710 
pinguis 699, 702, 706 
polytropa 229, 233 
pulicaris 229 
rubina 706 
rupicola 42, 229 
scotopholis 229 
thamnitis 698-700, 702, 704-710 
Leccinum 93, 108 
sect. Luteoscabra 108 
aeneum 273 
andinum 93, 106-108 
armeniacum 273 
brunneum 273 
califomicum 273 
constans 273 
insigne 
var. brunneum 273 
largentii 273 
manzanitae 2, 274 
var. angustisporae 274 
var, angustisporum 274 
montanum 274 
subalpinum 274 
subglabripes 108 
Lecidea 42, 226, 229, 233, 234 
atrobrunnea 38, 42, 44, 229 
carpathica 234, 236 
demissa 43 
fuscoatra 38, 42, 229 
globifera 44 
glomerulosa 42 
lurida 39 
mannii 42, 45 
novomexicana 44 
scalaris 39 
stigmatea 234 
tesselata 38, 229 
Lecidella 42 
carpathica 229, 234, 236 
euphora 42 
stigmatea 229, 234 
subincongrua 702 
Leioderma 
amphibolum 655, 660, 664 
Lemonniera 489, 494 
terrestris 494 


vel 


Lempholemma 421, 424, 426 
Leocarpus 
fragilis 633 
Lepiota 239 
sect. Cristatae 239, 242 
sect. Lilaceae 245, 248 
sect. Stenosporae 242 
castaneidisca 239, 244 
cristata 239-242, 244, 246 
cristatoides 245 
luteophylla 239, 274 
neophana 239, 245-247 
thiersii 239, 242-246, 274 
Lepraria 
neglecta 42 
Leptocapnodium 
krameri 198, 202 
Leptochidium 44 
albociliatum 42, 229 
Leptogium 229 
califomicum 42, 229 
comiculatum 42, 229 
furfuraceum 42 
laceroides 655, 660, 662, 663 
lichenoides 42 
minutissimum 42 
palmatum 42 
Leptonia 
thiersii 276 
Leptosphaeria 
subg. Massariosphaeria 508 
baldingerae 514 
clavicarpa 509 
lasioderma 523 
occidentalis 514 
phaeospora 512 
ramulicola 524 
sambucina 513 
typhicola 514 
Leptoxyphium 210 
Letharia 
columbiana 42 
Licea 
minima 633 
Lichinella 
stipatula 42 
Limacinia 
alaskensis 204 
multiseptata 213, 214 
Limacinula 207 
anomala 207 
Lithographa 643 
tesserata 643-645 
var. nivalis 644 
var. petraea 644 
Lojkania 
nuda 579 
separans 579 
Lopadostoma 
caespitosum 592 
conorum 596 
formosum 580 


728 


microecium 580 
Lulworthia 539 

grandispora 532, 539 _ 
Lycogala 

epidendrum 633 

flavofuscum 633 


Macrobiotophthora 443, 456 
vermicola 456 
Marasmiellus 
ramealis 
var. californicus 274 
Marasmius 71 
sect. Globulares 85, 100 
sect. Marasmius 71, 74 
sect. Neosessiles 90, 91 
sect. Sicci 71, 80, 84, 85, 90, 91 
subsect. Neosessilini 90 
subsect. Pararotulae 74 
subsect. Siccini 80, 90 
subsect. Spaniophyllini 90 
ser. Actinopodes 80 
ser. Haematocephali 90 
ser. Leonini 90 
applanatipes 274 
armeniacus 90 
capillaris 75 
ciliatomarginatus 71, 76-78, 80 
cormugatus 
var. aurantiacus 80 
decipiens 85 
falcatipes 71, 85, 87-91 
glabellus 81, 83, 84 
ilicicola 71, 72-75 
ilicis 75 
paludigenus 71, 81, 82, 84, 85 
polycystis 90, 91 
pusio 90 
var. guatopoensis 90 
var. pusio 90 
rotalis 72 
scototephrodes 74, 75 
siccus 76, 81, 85 
similis 84 
sullivantii 80 
tetrachrous 74, 75 
thiersii 274, 276 
Massalongella 
carpinicola 587 
Massarina 497 
Massariosphaeria 507, 508 
phaeospora 508, 512 
rubicunda 508 
typhicola 514 
Massospora 443 
Megalospora 
tuberculosa 405 
Melanconis 
tiliacea 585 
Melanelia 43 
fuliginosa 42 
glabra 43 


glabroides 43 
incolorata 43 
multispora 43 
subargentifera 43 
subaurifera 43 
subelegantula 43 
subolivacea 43 
Melanoleuca 
reai 
var. texana 274 
Melanomma 507 
subg. Chaetomastia 507, 50 
atrogranum 595 a 
canescens 507 
cucurbitarioides 507 
glaciale 512 
hirtulum 507, 512 
hispidulum 507 
hypoxylon 578 
pilosellum 507 
plejosporum 507 
sambuci 513 
suldensis 512 
Meliolopsis 
heteromeles 207, 212 
Melogramma 
aethiops 582 
campylosporum 583 
lateritia 583 
vagans 583 
Menispora 679, 683, 684, 691, 695, 696 
britannica 679, 683, 691, 695, 696 
ciliata 683, 695 
glauca 683, 695 
manitobaensis 683, 695 
tortuosa 683, 695 
uncinata 679, 683, 691, 695, 696 
Menisporopsis 679, 684, 691, 694, 696 
ludoviciana 683 
novae-zelandiae 683 
pirozynskii 684 
pleiosetosa 683 
profusa 683 
theobromae 683 
MERISTACRACEAE 456 
Meristacrum 443, 456, 457 
Micromphale 
arbuticola 274 
sequoiae 274 
Microthelia 
accedens 524 
celtidis 520 
incrustans 579 
vagans 524 
Mollisia 647 
lesdainii 647, 648 
Montagnula 507, 508 
hirtula 512 
infernalis 508 
Morchella 17, 18 
esculenta 17 


729 


Morfea 211 Otthia 517, 518 
alaskensis 204 clematidis 522, 523 
hendrickxii 208 distegiae 519, 520 

Mucilago fendleraecola 519, 520 
crustacea 633 fruticola 522 

Mutinus hypoxyloides 578 
caninus 148 lisae 523 

Mycena 3 quercicola 524 

Mycosphaerella spiraeae 517 
acervata 582 symphoricarpi 519, 520 

Mycothyridium 
americanum 596 Pagidospora 431 
antiquum 595 Pandora 441, 443, 446, 451 
pallidum 596 blunckii 452 
stilbostomum 596 brahminae 452 
syringae 597 bullata 452 
vitis 597 calliphorae 452 

dacnusae 452 

Naetrocymbe delphacis 452 
scoriadea 202 dipterigena 452 
stevensonii 207 echinospora 452 

Nanomyces 565 formicae 452 

Neodeightonia gammae 453 
ramulicola 524 gloeospora 453 

Neofuscelia 43 kondoiensis 453 
loxodes 43 neoaphidis 452 

Neozygites 442, 443, 457 nouryi 453 

Nephroma phalangicida 453 
laevigatum 38 suturalis 453 

Nesolechia Pannaria 42, 43 
lesdainii 647, 648 amphibela 660 

Niebla 702 amphibola 660 

Normandina leucophaea 43, 229 
pulchella 43 leucostictoides 43 

Nummularia praetermissa 43 
albosticta 591 thysanota 663 
lateritia 591 Pannularia 
pezizoides 591 amphibela 660 
repanda 591 Papulospora 434 

var. zonata 591 Paraparmelia 400 
rufa 591 subtropica 400 
subapiculata 591 Parmelia 43, 399, 402, 421, 424, 426 
vemicosa 591 sect. Paraparmelia 400 

Numulariola adhaerens 404 
albosticta 591 adplanata 404 
cylindrophora 589 agamalis 399 -401 

applicata 403 

Ochrolechia amoldii 43 
subpallescens 43 caliginosa 400 
upsaliensis 43 caperata 39 

Opegrapha conspersa 402 
tesserata 644 cumberlandia 45 

Ophiocapnocoma elegantula 43 
multiseptata 214 eruptens 402 

Ophiostoma 637, 641 flaventior 39 
angusticollis 641 geckonalis 399, 400-402 
epigloeum 641 glabra 43 
grande 641 glabratula 42 
microsporum 641, 642 inconspicua 403 
nigrocarpum 641 ioannis-simae 45 
polyporicola 637 -641 ischnoides 400 


roraimense 641 isidiotyla 43 


730 


kurokawae 405 
lesothoensis 403 
lineola 45 
mexicana 45 
mixta 399, 402 
mougeotii 402 
mudata 
var. pulverata 659 
multispora 43 
natalensis 402 
neoquintaria 400 
novomexicana 45 
numinbahensis 400 
paradoxa 402 
perfunctata 399, 402, 404 
perlata 43 
phyllodactylaris 399, 402 
protodysprosa 399, 403 
pseudoaspera 43 
pseudoglabra 43 
quercina 43 
rugulosa 403 
salax 399, 403 
saniensis 399, 403, 404 
saxatilis 43 
spissa 404 
stuppea 43 
subargentifera 43 
subaurifera 43 
subolivacea 43 
subramigera 404 
subrudecta 44 
sulcata 43 
tantillum 399, 403, 404 
taractica 45 
umtamvuna 399, 403, 404 
Parmeliella 43 
amphibola 660 
cyanolepra 229 
praetermissa 43 
thysanota 655, 660, 663, 664 
Parmelina 43 
quercina 43 
Parmotrema 43 
amoldii 43 
chinense 43 
stuppeum 43 
Parodiella 518 
fruticola 522 
Peltigera 
collina 44 
Peltosphaeria 
canadensis 596 
Peltula 
bolanderi 229 
euploca 44 
omphaliza 227, 230 
zahlbruckneri 38 
Penicillium 250 
Penzigia 
turbinata 592 
Peridiothelia 538 


Peroneutypa 
heteracantha 587 
Peroneutypella 
capillata 597 
microcarpa 597 
Pertusaria 
albescens 44 
amara 44 
chiodectonoides 44 
lecanina 44 
Phaeochaetia 
arbutifoliae 207, 209 
Phaeophyscia 702 
orbicularis 44 
Phaeosaccardinula 
anomala 214 
dematia 198, 202, 208, 213, 214 
Phaeosphaeria 
baldingerae 514 
berlesei 510 
clavispora 509 
typhicola 514 
Phaeoxyphiella 202-205 
fisheri 197, 199, 201-203, 205, 206 
morototoni 199, 208 
walteri 199 
Phallus 
impudicus 148 
Phlyctis 
argena 44 
Phomatospora 
wistariae 592 
Phragmodiaporthe 
tiliacea 585 
Phylloporus 93, 99, 100, 102 
boletinoides 274 
Physarum 
cinereum 633 
compressum 633 
leucophaeum 634 
nutans 634 
pezizoideum 634 
pusillum 634 
straminipes 634 
vernum 634 
Physcia 226, 704 


adscendens 44, 230, 423, 424, 426, 427 


aipolia 44 
alba 44 

var. obsessa 44 
albinea 44 
caesia 234 
callosa 44 
cascadensis 44 
grisea 

f. detersa 44 

f. enteroxanthella 44 
leptalea 44 
mexicana 44 
millegrana 44 
orbicularis 44 


phaea 44 
pulverulenta 44 
semipinnata 44 
stellaris 44, 230 
tenella 44 
Physconia 
detersa 44 
distorta 44, 230 
enteroxantha 44 
grisea 44 
f. isidiigera 230 
muscigena 44 
Pilobolus 395-397 
crystallinus 396 
kleinii 396 
roridus 396 
Piptoporus 
betulinus 633 
Placodium 709, 710 
sect. Thamnonoma 698 
saxicola 234 
Placographa 
tesserata 644 
Placynthium 
nigrum 230 
Pleosphaeria 
salicina 208 
Pleospora 
antiqua 595 
Plowrightia 
fruticola 522, 523 
symphoricarpi 518-520 
Podosordaria 
leporina 592 
mexicana 592 
Podoxyphium 210 
yuccae 210 
Polycauliona 698 
bolanderi 699 
thamnitis 699 
Polychaetella 
araucariae 209 
Polychaeton 210 
Polychidium 44 
albociliatum 42 
Polycladium 379 
equiseti 379 
Polyporus 634 
Poronia 
leporina 592 
minuta 592 
turbinata 592 
Porphyrellus 
subvirens 94 
Porpidia 
urbanskyana 405 
Protogautieria 
substriata 274 
Protophallus 135 
brunneus 137 
jamaicensis 138 
Protoventuria 517 


731 


Protubera 133-138, 146, 148 
africana 133-135, 140, 149 
borealis 133, 134, 136, 137, 140, 143, 
144, 149 
brunnea 133, 135, 137, 138, 149 
clathroidea 133-135, 138, 140, 146, 148, 
149 
jamaicensis 133, 135, 138-140, 143, 149 
maracuja 133, 134, 138, 140, 142, 144, 
148, 149 
nipponica 133, 134, 139, 140, 143, 144, 
149 
sabulonensis 133, 144, 146, 148, 149 
Protuberella 134 
borealis 134, 136 
Psalliota 120 
Psathyrella 
atrospora 274 
ellenae 
var. yubaensis 275 
griseopallida 275 
lithocarpi 275 
sequoiae 275 
texensis 275 
thiersii 276 
Psathyrophlyctis 
serpentaria 405 
Pseudevernia 39 
Pseudocyphellaria 
anomala 44 
anthraspis 44 
Pseudotthia 517 
symphoricarpi 517, 519-520 
Pseudovalsa 
comptoniae 582 
texensis 584 
Psora 42, 709 
califomica 44 
globifera 44, 230 
nipponica 44 
Psorula 
rufonigra 230 
Pulveroboletus 100 
Punctelia 43 
stictica 704 
subrudecta 39, 44 
Pycnoporellus 
metamorphosus 436 
Pycnothelia 706 
Pyrenopsis 
phaeococca 230 
Pyrenula 
imperfecta 581 
Pyricularia 
aquatica 497 
submersa 497, 500, 501 
Pyrrhoglossum 219 
lilacinum 219 
lilacipes 219 


Quadricladium 489 
aquaticum 489, 490, 492, 494 


foe 


Ramalina 
ecklonii 423, 424, 426 
farinacea 44 
leptocarpha 44 
menziesii 44 
Ramaria 601 
subg. Echinoramaria 609 
ser. Grandisporae 609, 610 
amyloidea 610 
apiahyana 610 
aurea 610 
celevirescens 610 
claviamulata 610 
cyanocephala 601, 608, 610 
flavobrunnescens 610 
formosa 610 
grandis 609 
guayensis 610 
mollerana 610 
nigrescens 610 
stricta 610 
thiersii 276 
velocimutans 610 
verna 610 
zippelii 601, 609, 610 
Ramonia 
gyalectiformis 226, 230 
Requinella 538 
Rhizocarpon 232, 235 
bolanderi 42, 44, 230, 234, 235 
effiguratum 235 
ferax 44 
geographicum 230, 232, 236 
grande 230 
sphaericum 235 
viridiatrum 230, 232, 235, 236 
Rhizophila 527 , 531, 532 
marina 527, 528, 529, 531, 532 
Rhizoplaca 
chrysoleuca 230, 706 
glaucophana 38 
marginalis 38 
melanophthalma 230, 698 
Rhizopodomyces 565 
Rhizopogon 134, 136 
Rhodocybe 47, 48 
sect. Rhodophana 47, 50 
carlottae 48 
var. carlottae 47, 48 
var. vinacea 47-49 
eccentrica 50 
mycenoides 50 
nitellina 50 
priscua 50 
speciosa 50 
trachyospora 47, 48, 50, 52 
var. griseoviolacea 49 , 52 
var. purpureoviolacea 49 , 50, 52 
var. trachyospora 47, 49, 51, 52 
var. vinacea 49, 52 


Rhopographus 
clavisporus 509 
Rinodina 
hallii 45 
tephraspis 230, 232 
Rosellinia 539 
abietina 


var. trichota 592, 595 


albolanata 592 
arctispora 592, 595 
bakeri 592 
bicolor 592 
bigeloviae 592 
caespitosa 592 
compressa 592 
confertissima 593 
geasteroides 593 
gigantea 593 
gigaspora 593 
glandiformis 593 
hystrix 593 
kellermannii 593 
langloisii 593 
limoniispora 593 
macouniana 593 
macra 593 
megaloecia 593 
melaleuca 594 
muriculata 594 
obliquata 

var. americana 594 
ostiolata 594 
ovalis 594, 595 
parasitica 594 
pinicola 594 
poliosa 594 
pulcherrima 594 
pulveracea 595 
subcompressa 594 
thelena 

var. terrestris 594 
trichota 594, 595 
xylarispora 594, 596 


Scirthia 
insculpta 523 
Scolecoxyphium 202 
americanum 210 
Scutellospora 675 
heterogama 675 
pellucida 675 
Scytinostroma 608 
Skyttea 647, 648 
cruciata 648 
lesdainii 648 , 649 
Sphaeria 
abietina 


var. trichota 592, 594 


antiqua 595, 596 
arctespora 592, 595 
atrograna 578, 595 
bisphaerica 595 


fissicula 582, 595 

hirtula 512 

leucobasis 580, 595 

mortuosa 580, 595 

multiformis 

f. effusa 589 

oronoensis 579, 595 

ovalis 594, 595 

pholidigena 581, 596 

picacea 580, 596 

ramulicola 524, 525 

sabalensioides 581, 596 

xylariaespora 594, 596 
Sphaerophorus 

globosus 38 
Sphaeropsis 525 

celtidis 522 
Sphaleromyces 

lathrobii 575 
Splanchnonema 

melanterum 579 
Sporothrix 637, 639, 641 

curviconia 641 
Sporotrichum 435 

aurantiacum 429, 436 

azureum 429, 435, 436 
Stemmatomyces 575, 576 
Stemonitis 

virginiensis 634 
Sticta 

fuliginosa 45 
Stigmatomyces 566, 570, 572, 575, 576 
Stilbothamnium 250 

togoense 249-251 
Strigopodia 

batistae 205 
Strobilomyces 93 

confusus 109 
Strongwellsea 443 
Sugiyamaemyces 565, 566, 572, 574- 

576 

orousettii 565, 566-568, 570, 572 
Suillus 

acerbus 275 

fuscotomentosus 275 

glandulosipes 275 

kaibabensis 275 

luteus 93 

monticola 275 

occidentalis 275 

pungens 275 

reticulatus 275 

riparius 275 

volcanalis 275 

wasatchicus 276 
Symphytocarpus 

flaccidus 634 


Tarichium 443 
gammae 453 
Teloschistes 
chrysophthalmus 423, 424, 426, 427 


733 


Tephromela 

atra 230 
Tetraposporium 489, 494 
Thaxterogaster 

sect. Aporpogaster 66 

conicum 65-67, 70 

porphyreum 65-68 

thiersii 65 -70, 276 
Thaxterosporium 442, 443 
Thelephora 601, 612 

aurantiotincta 601, 610, 612 
Thelidium 227, 230 
Thelomma 

mammosum 45 
Thyridaria 

comptoniae 582 

lateritia 583 

texensis 584 
Thyridella 

canadensis 596 
Thyridium 

americanum 596 

antiquum 595, 596 

canadense 596 

pallidum 596 

stilbostomum 596 

syringae 596 

tuberculatum 525 

Vitis 597 
Toninia 

aromatica 230 

squalida 230 
Trametes 

trogii 623 
Trematosphaeria 

phaeospora 512 
Tremella 

fuciformis 641 
Trichia 

varia 634 
Tricholoma 130 
Tuber 

candidum 392 

harknessii 392 

melanosporum 392 

texense 387, 388, 391-393 
Tubifera 

ferruginosa 634 
Tuckermannopsis 39 

chlorophylia 45 

mermillii 45 

orbata 45 
Tulostoma 

thiersii 276 
Tylopilus 93 

ammiratii 276 

humilis 276 

niger 111 

nigerrimus 111, 112 

nigropurpureus 111 

obscurus 93, 109-112 
Typhula 


734 


idahoensis 260, 264 Varicosporium 379 
ishikariensis 260, 264 delicatum 379 
Tyromyces elodeae 379 
stipticus 637, 638, 640 scoparium 375 -379 
Verrucaria 226, 233 
Umbilicaria 38 aethiobola 231, 233 
krascheninnikovii 231 margacea 231, 233 
phaea 45, 231 maura 704 
polyphylla 45 muralis 231 
polyrrhiza 45 nigrescens 231 
Unguiculariopsis 647, 648, 650 viridula 231 
jamaicensis Vertixore 
thallophila 647, 648 atronitidum 206, 207 
Unguiculella 650, 652 
aggregata 650 Xanthoparmelia 43, 541, 542, 544, 546- 
jamaicensis 650, 652, 653 548, 550, 553, 555-557, 560, 562, 563 
meliolicola 650 afrolavicola 541 ,542 
oregonensis 650, 652 amphixanthoides 558, 561 
Urosporella amplexula 543 
alabamiensis 581 amplexuloides 541 , 542, 559 
Usnea 38, 45 angustiphylla 556 
arizonica 45 applicata 403 
cavemosa 45 ausiana 541, 543,544 
igniaria 423, 424, 426, 427 brevilobata 403, 404 
trichodea 45 catarinae 541, 543, 544 
chalybaeizans 561 
Valsa chlorochroa 551, 561 
acervata 582 cirrhomedullosa 541, 544, 545 
canodisca 587, 597 colorata 543, 552, 554, 556 
capillata 587, 597 coneruptens 541, 545, 546 
caryigena 582 conjuncta 541, 545, 546, 562 
ceanothi 525 crassilobata 552 
celastrina 582 cumberlandia 45, 226, 231, 236 
collariata 582 diffractaica 403 
comula 582 eruptens 402, 545, 558 
comuta 582 filarszkyana 554 
deusta 587, 597 granulata 541, 545, 546 
fibritecta 583 harrisii 541,546, 547 
hochelagae 583 hypopsila 543 
juglandina 588, 597 idahoensis 541, 547, 548 
lutescens 588, 597 imbricata 541, 547,548 
maclurae 588, 597 inconspicua 403, 404 
macounii 583 inflata 541,548, 549 
megastoma 583 isidiascens 543 
microcarpa 588, 597 keralensis 564 
microstega 584 khomasiana 541, 548, 549 
minima 584 kiboensis 559 
radiata 584 kotisephola 541, 549,550 
rugiella 588, 597 lavicola 541 
tumida 585 lesothoensis 403, 545 
venusta 588, 598 lineola 45 
Vitis 586 lipochlorochroa 541, 550, 551 
Valsaria luderitziana 541,550, 551,554 
aethiops 582 luminosa 561 
insitiva 582 ned etae a 


mapholanengensis 541, 550, 551 


magnoliae 524, 525 maxima 541, 552,553 
pustulans 584 mexicana 45 
symphoricarpi 519 microspora 562 

Valsella mougeotii 562 
pulcherrima 586 mougeotina 562 


Vararia 608 multipartita 556 


natalensis 402, 404 
naudesnekia 403, 404, 562 
neosynestia 559 
neowyomingensis 541, 552,553 
norcolorata 541,553, 554 
norlobaronica 541,553, 554 
norwalteri 541,551, 554,555 
novomexicana 45 
paradoxa 402 
prodomokosii 549 
protodysprosa 403 
protolusitana 541, 554,555 
protoquintaria 541,555, 556 
pseudocongensis 562 
psornorstictica 541,555, 556 
pustulifera 402 
pustulosorediata 541, 556,557 
rugulosa 403 
salamphixantha 541,557, 558 
saleruptens 541,557, 558 
salkiboensis 541,557, 558 
saniensis 403, 404 
scabrosa 543 
shebaiensis 562 
springbokensis 541, 559, 560 
stenosporonica 562 
subamplexuloides 541, 543, 559, 560 
subbullata 541, 559, 560 
subcolorata 545 
subconvoluta 541, 560, 561 
subdecipiens 549 
subluminosa 541, 561, 563 
submougeotii 541, 561,563 
substenophylloides 541, 562, 563 
taractica 45 
tsekensis 541, 562, 563 
umtamvuna 403, 404 
walteri 551, 554 
weberi 549 
wyomingica 552, 561 

Xanthoria 231 
candelaria 45, 704 
fallax 45 
lobulata 45 
parietina 45, 423, 425, 426 
polycarpa 45, 425, 426 

Xenasma 432 

Xerocomus 
globuliger 100 
subtomentosus 101 
truncatus 102 

Xylaria 283, 284, 289, 292, 295, 296, 300, 

306, 316, 320, 324, 326, 330, 334, 343, 
353, 358, 362-369 

adscendens 290, 301, 302, 364 
aemulans 363 
alata 283, 293, 301, 302, 334 
allantoidea 286, 302, 304 
amphithele 283, 300, 304, 334 
anisopleura 285, 289, 304, 308, 310, 367 
apiculata 318 
arbuscula 293, 295, 299, 304, 306, 310, 


i fale. 


312, 14,516; 301 

aristata 298, 314, 343, 366 

bambooensis 294, 306, 316, 336 

berkeleyi 367 

biformis 367 

brachiata 291, 318, 320 

brevipes 296, 306, 320 

chardoniana 592 

claviceps 283, 290, 308, 320, 336 

coccophora 297, 308, 324, 352 

comosa 285, 308, 324, 326 

comosoides 326 

compressa 324 

comiculata 287, 308, 310, 328 

comiformis 334 

cubensis 286, 328, 329, 346 

curta 291, 310, 329, 330 

cylindrica 598 

dealbata 345 

dichotoma 294, 310, 330 

enterogena 286, 310, 312, 332, 333, 338 

feejeensis 290, 296, 312, 316, 320, 333, 
334, 336, 365, 371 

fockei 284 

globosa 308, 310, 367 

gracillima 293, 312, 336, 338, 344 

grammica 287, 312, 314, 338, 340, 345 

guazumae 283, 298, 316, 336, 340, 
343 

guyanensis 287, 314, 341, 342 

heloidea 341 

hyperythra 322 

hypoxylon 347 

ianthina-velutina 299, 316, 330, 342, 343 

inaequalis 299, 338, 343, 344 

juruensis 297, 314, 336, 344 

kegeliana 287, 314, 338, 344, 345 

laevis 286, 316, 336, 345, 346 

longiana 295, 326, 346, 347 

longipes 283-285, 288, 316, 347, 348 
var. tropica 283, 288, 316, 336, 348 

luteostromata 365 

magniannulata 283, 294, 318, 334, 
348, 349 

magnoliae 285, 298, 316, 318, 330, 343, 
349, 350 

maitlandii 292, 318, 350, 351 

mellisii 318, 351 
var. nuda 297, 318, 351 

microceras 297, 320, 351 

multiplex 293, 320, 352, 353, 364, 598 

muscula 292, 320, 353 

myosurus 368 

nigrescens 285, 320, 338, 346, 354 

obovata 361 

ophiopoda 288, 359 

oxyacanthae 285, 300, 320, 354, 355 

pallida 292, 322, 355 

pallide-ostiolata 288, 322, 338, 356 

persicaria 285, 299, 322, 356, 357 

phyllocharis 301, 322, 357, 358 

poitei 287, 342, 358 


736 


polymorpha 285, 288, 324, 358-362, 365 hirschiopori 683, 694 
rhytidophloea 371 Zanclospora 684, 691, 695, 696 
scabriclavula 283, 294, 324, 336, 360 austroamericana 683, 694 
schweinitzii 289, 324, 359, 361 brevispora 683, 694 


scruposa 289, 324, 328, 356, 360, 362 mystica 684 
sicula novae-zelandiae 683, 694 
f. major 366 Zeugandromyces 575 


squamulosa 283, 291, 330, 334, 368 Zoophthora 441-444, 450, 451, 453, 459 


telfairii 287, 332, 369 

teres 598 

uniapiculata 283, 290, 330, 336, 370 

venustula 287, 340 
Xylosphaera 338, 351, 356 

mellisii 318 

var. nuda 351 

Xylosphaeria 

americana 596 

antigua 595 

brachystoma 580 

mortuosa 595 

pallida 596 

stilbostoma 596 

syringae 596 

Vitis 597 


Zakatoshia 684, 691, 695 


subg. Erynia 448 
subg. Furia 450 
subg. Pandora 451 
anglica 453 
anhuiensis 453 
aphidis 453 
canadensis 453 
crassitunicata 453 
geometralis 453 
lanceolata 453 
occidentalis 453 
orientalis 453 
petchii 453 
phalloides 453 
phytonomi 453 
radicans 450, 453 
vomitoriae 451 


Zygnemomyces 443, 456, 457 


Yor 


REVIEWERS, VOLUME THIRTY-FOUR 


The Co-Editors express their appreciation to the following individuals who have, prior to 
acceptance for publication, reviewed one or more of the papers appearing in this volume: 


R. K. BENJAMIN T. LUMBSCH 
M. E. BARR BIGELOW L. MARANOVA 
C. BAS T. NASH, III 
G. L. BENNY N. E. NANNENGA-BREMEKAMP 
J. R. BOISE M. E. PALM 
I. M. BRODO R. H. PETERSEN 
B. E. CALLAN D. PINKAVA 
M. CAMPOS J. POELT 
L. M. CARRIS J. E. PURKYNE 
D. S. DE HOOG D. REYNOLDS 
E. DESCALS R. G. ROBERTS 
O. E. ERIKSSON J. D. ROGERS 
T. L. ESSLINGER C. T. ROGERSON 
AG ELIX G. J. SAMUELS 
W. FERREN J. A. STALPERS 
W. GAMS I., I. TAVARES 
I. J. GAMUNDI H. D. THIERS 
D. W. GRUND J. W. THOMSON 
M. E. HALE G. VOBIS 
R. E. HALLING R. WATLING 
D. L. HAWKSWORTH W. A. WEBER 
D. T. JENKINS J. WEBSTER 
W. JULICH T. E. WEIER 
J. W. KIMBROUGH D. T. WICKLOW 
J. KOHLMEYER N. WILDING 
K. E. KOSKE J. WRIGHT 


M. F. LOPEZ ARMENGOL 


MYCOTAXON PUBLICATION DATES 


Volume 33 (October-December 1988) November 28, 1988 
Harry Thiers Festschrift January 20, 1989 


Volume 34(1) 


738 


ERRATA, VOLUME THIRTY-ONE 


Page 485 fig. 1 scale bar should read 1.5 mm, not 1.5 Um 
489 line 44 for (1986): read (1987): 
46 for 23:501-507. read 28:501-507. 


ERRATA, VOLUME THIRTY-FOUR 


Page 38 line 41 for tesselata read tessellata 
49 line 17 _ for trachyosporum var. trachyosporum 
read trachyospora var. trachyospora 
27 for trachyosporum vars. griseoviolaceum and purpureoviolaceum 
read trachyospora vars. griseoviolacea and purpureoviolacea 
lines 29,32,33,37,38 for trachyosporum _ read trachyospora 
line 34 for trachyosporum var. griseoviolaceum 
read tracyospora vat. griseoviolacea 
36 for trachyosporum var. purpureoviolaceum 
read trachyospora vat. purpureoviolacea 
38 for griseoviolaceum read griseoviolacea 
50 line 1 _ for trachyosporum var. purpureoviolaceum 
read trachyospora var. purpureoviolacea 


TOs eines | for ligulus read ligula 
197 line 11 for dematum read dematium 
198 lines 18,21 for dematum read dematium 
200 line 3 for dematum read dematium 
201 line 20 for dematum read dematium 
202 lines 20, 22, 40 for dematum read dematium 
203 line 47 for dematum read dematium 
208 lines 8, 14 for dematum read dematium 
line 28 for dematum read dematia 
213 lines 9,12,15 for dematum read dematium 
line 37 for Arthroboytrum read Arthrobotryum 
214 slines 9.10 for multiseptatum read multiseptata 
234 line 12 for saxicolum read saxicola 
23 for Lecidella carpathica read Lecidella carpathica 
37 for Aspicilictum read Aspicilietum 
255 line? 12 for ochraea read ochracea 
02 for spahaericum read sphaericum 
lines 37, 39 for Aspicilictum read Aspicilietum 
236 lines 2, 3, 6, 12 for Aspicilictum read Aspicilietum 
line 9 for carpatica read carpathica 


250 line 13 for togoense read togoensis 


MY COTAXON 


AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION 
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS 


ACTA TANG FSS MD, GD, Gl FAV IE oe, 


COMPLETE IN TWO ISSUES, THE “THIERS FESTSCHRIFT’ 
AND ONE QUARTERLY ISSUE, 


CONSISTING OF v + 738 PAGES INCLUDING FIGURES 


CO-EDITORS 


G. L. HENNEBERT 
French Language Editor & Book Review Editor 


Laboratoire de Mycologie systématique et appliquée 
Université de Louvain, B-1348 Louvain-la-Neuve, Belgium 


RICHARD P. KORF 
English Language Editor & Managing Editor 


Mycotaxon, Ltd., P.O. Box 264 
Ithaca, NY 14851, USA 


SUSAN C. GRUFF 
Associate Editor & Index Editor 


Plant Pathology Herbarium, Cornell University 
Ithaca, NY 14853, USA 


Published by 


MYCOTAXON, LTD., P.O. BOX 264 
ITHACA, NY 14851, USA 


Printed in the United States of America 


TABLE OF CONTENTS, VOLUME THIRTY-FOUR 


No. 1. January 20, 1988 


A FESTSCHRIFT IN HONOR OF HARRY D. THIERS 
Guest Co-Editors: ROY E. HALLING and BARBARA M. THIERS 


RE ke ocd ca eats Bite ted ee ks is eh Petcesn ccm S viele cags ks 
Harry D. Thiers: Reminiscences about a teacher and friend ............... 
Remembering the morel grower: Ron Ower, 1939-1986 ................. 
Dermocybe, subgenus Dermocybe, section Sanguineae in northern 
Oe TTT PS FOSS aN Re POM ead Mp) 7 ae Joseph F. Ammirati 
Lichens of Mount Diablo State Park, Contra Costa County, California. 
Doris E. Baltzo 
The genus Rhodocybe: new combinations and a revised key to section Rhodo- 
phana in North America.. Timothy J. Baroni and David L. Largent 
Quality control factors for A/ternaria allergens. 
Harriet A. Burge, Marion E. Hoyer, William R. Solomon, 
Emory G. Simmons, and Janet Gallup 
Thaxterogaster thiersii: anew secotioid species from California. 
Cornelia J. Calhoun 
Studies on Marasmius from eastern North America. II. New species. 
Dennis E. Desjardin and Ronald H. Petersen 
Pures Or OlOMDIAN DOLCIES = 7.) os uc es chs ee eae ore Roy E. Halling 
Cladonia thiersii: a new lichen from California......... Samuel Hammer 
Studies in Agaricus IV: new species from Colorado.. Richard W. Kerrigan 
A new, lignicolous species of Entoloma (Entolomataceae, Agaricales) from 


ET ee oe as stistnek Sach ei'g age cetoleul « reth.o David L. Largent 
Ee UOTOCUUS IPOLUDETQ ..c «cisterns oe cs os owe wie to oge David Malloch 
Notes on Clavariadelphus. Ill. New and noteworthy species from North 

ORCS 5 2 teil aia pe aati iiral ara irs Mara an Andrew S. Methven 
A key to the species of Inocybe in California....... Florence H. Nishida 
Foliicolous fungi 8: Capnodium in California......... Don R. Reynolds 
A new species of Gymnopilus from northern California.. Michelle T. Seidl 
The lichens of serpentine rocks and soils in California..... Lorene L. Sigal 


Lepiota sensu lato in California. III. Species with a hymeniform pileipellis. 
Walter J. Sundberg 
Examination of Stilbothamnium togoense for Aspergillus flavus group 
PIV COLOXING 04. sk. sie s s D. T. Wicklow, R. F. Vesonder, Cesaria E. 
McAlIpin, R. J. Cole, and Marie-France Roquebert 
The occurrence of vesicular-arbuscular mycorrhizae in burned areas of the 
Snake River Birds of Prey Area, Idaho..... Marcia Wicklow-Howard 
Compatibility and fruiting studies of an albino form of Auricularia cornea. 
George J. Wong 
Master’s degree students of Harry D. Thiers and thesis titles.............. 
Type specimens of agarics, boletes and gasteromycetes in the San Francisco 
State University herbarium (SFSU). 
Barbara M. Thiers and Roy E. Halling 
Spevies named tor Harry:D. Thiers.) 7°. 62... Dennis E. Desjardin 


No. 2. January-March 1989 


The anamorphs of Diatrypella prominens and Eutypella sabalina. 
Dean A. Glawe and John P. Jones 
A preliminary account of Xylaria of Mexico. 
Felipe San Martin Gonzalez and Jack D. Rogers 


iil 


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129 
135 


133 
181 
197 
PAG 
pp 


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249 
253 
Zo 
267 


269 
276 


214 
283 


iv BEE ERRATA 


Varicosporium scoparium, a new staurosporous hyphomycete. 
A. Roldan and M. Honrubia 
A new: addition tothe genus Gorgomiycesy. 04). so. ee A. Roldan 
The occurrence of Tuber texense in Georgia. 
Richard T. Hanlin, Mei-Lee Wu, and Timothy B. Brenneman 
A survey of Pilobolus from Yellowstone National Park. 
K. Michael Foos and Judith A. Royer 
Two new species of Parmelia (Parmeliaceae, Lichenes), further new 
combinations and notes, and additional new lichen records from southern 
AEP CAN Oy hee weet MeL) atten ache neat Uaihee 3 Franklin A. Brusse 
Pollution atmosphérique et lichens dans la ville de Santiago du Chile. 


Manuel Mahu % 


Studies on xylophilous fungi from Argentina. [V. Anamorphs of Basidiomycetes 
on Eucalyptus viminalis (Myrtaceae). 
A. I. Romero, D. Cabral, and S. E. Lopez 
Synopsis of a revised classification for the Entomophthorales (Zygomycotina). 
Richard A. Humber 
Mnemonic three-letter acronyms for the names of fungal families. 
Joseph E. Laferriére 
Canalisporium, a new genus of lignicolous hyphomycetes from Malaysia. 
A. Nawawi and A. J. Kuthubutheen 
Quadricladium aquaticum gen. et sp. nov., an aquatic hyphomycete with 
tetraradiate Conidid 2. awa. o. A. Nawawi and A. J. Kuthubutheen 
A new taxon in Colispora (Hyphomycetes) from Malaysia. 
A. Nawawi and A. J. Kuthubutheen 
Ascomycetes of western India XIII........ Alaka Pande and V. G. Rao 
The genus Chaetomastia (Dacampiaceae) in North America. 
Margaret E. Barr 
The genus Dothidotthia (Botryosphaeriaceae) in North America. 
Margaret E. Barr 
Marine fungi from Seychelles. VIII. Rhizophila marina, a new ascomycete 


from mangrove prop roots........ K. D. Hyde and E. B. G. Jones 
Marine fungi from India. II. Acrocordiopsis patilii gen. et sp. nov. from 
MAaNPTOVE. WOOd Fy ole cet een. Lean ee B. D. Borse and K. D. Hyde 


New species in the lichen genus Xanthoparmelia (Ascomycotina: Parmeliaceae). 
Mason E. Hale 
Sugiyamaemyces, a new genus of Laboulbeniales (Ascomycetes) on 
Clidicus (Scydmaenidae) .... Isabelle I. Tavares and Jean Balazuc 
Index to J. B. Ellis’ types of pyrenomycetes with amyloid ascal rings. 
Katia F. Rodrigues 
Notes on tropical and warm temperate basidiomycetes. 
A. L. Welden and C. L. Ovrebo 
Amanita protecta — a new species from coastal southern California. 
Rodham E. Tulloss and Greg Wright 
Contribution to the study of the myxomycetes in Spain. I. 
Gabriel Moreno, Carlos Illana, and Michel Heykoop 
A new Ophiostoma on polypores .... O. Constantinescu and S. Ryman 
Lithographa, a lichen genus new to continental North America. 
JoAnn W. Flock 
Notes on one lichenicolous and one fungicolous discomycete. 
Wen-ying Zhuang and Richard P. Korf 
Noteworthy corticolous lichens in Nothofagus forests, north-western Patagonia. 
Susana Cavelo and Laura Lorenzo 
Gigaspora ramisporophora: anew species with novel sporophores from Brazil. 
Joyce Lance Spain, Ewald Sieverding, and Norman C. Schenck 
A numerical taxonomic study of some phialidic genera of hyphomycetes: 
cluster analysis ..... Angélica M. Arambarri and Marta N. Cabello 


The genus Cladidium (lichenized Ascomycotina) ........ Bruce D. Ryan 


BY 
381 


387 
395 


399 
407 


429 
441 
461 
475 
489 


497 
503 


507 
517 
527 
535 
541 
565 

ey) 
601 
615 


623 
637 


643 
647 
655 
667 
679 
697 


IRB ESE te at eh a See OA See en) Soa et Pht et EE ENE A LY gatelat fe al The: 
Se PUN POUS ANGIACHKCN LANA je cil) isle) vie sie, c bate & arated s ier) elaine casio re 716 
RN ree By FO Ce FS ites Pu et RAISE © tee nes phe dene ter ee. ae TolK weukhe 5 ToL 
Publication dates, MYCOTAXON Volumes 33 and 34(1) .............. Ta 
Errata 


an 


CO-EDITORS oF MYCOTAXON 


RICHARD P. KORF SUSAN C. GRUFF G. L. HENNEBERT 
English Language Editor Associate Editor French Language Editor 
& Managing Editor & Index Editor & Book Review Editor 
P.O. Box 264 Plant Pathology, C.U. UCL, Place Croix du Sud 3 
Ithaca, NY 14851 USA Ithaca, NY 14853 USA B-1348 Louvain-la-Neuve, Belgium 


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