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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.
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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.
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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-
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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.
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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,
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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.
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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
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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,
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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.
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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.
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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
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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.
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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-
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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).
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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}.
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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.
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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
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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.
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Pileus 4-13 cm broad, convex then