CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 FESTSCHRIFT IN HONOR OF HOWARD A. CRUM IN CELEBRATION OF HIS 70TH BIRTHDAY University of Michigan Herbarium Ann Arbor, Michigan 22 August 1992 322 22 z 392 342227 73 if, $555 £93 Howarb A. CRUM AND Hattry CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 FESTSCHRIFT IN HONOR OF HOWARD A. CRUM IN CELEBRATION OF HIS 70TH BIRTHDAY University of Michigan Herbarium Ann Arbor, Michigan 22 August 1992 CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM ISSN 0091-1860 Copyright 1992 University of Michigan Herbarium All rights reserved Printed in the United States of America Volume 18 Editor: Wilham R. Buck New York Botanical Garden For information about the availability and prices of previous volumes of the Contributions, address the Director, University of Michigan Herbarium, University Building, Ann Arbor, Michigan 48109-1057, U.S.A North 22 August 1992 CONTENTS Foreword WILLIAM R. Buck A tribute to Howard Crum Lewis E. ANDERSON Crumuscus vitalis gen. et sp. nov. (Ditrichaceae) WILLIAM R. Buck & JERRY A. SNIDER Key to the moss genera of North America north of Mexico Dace H. Virt & WILLIAM R. Buck A contribution toward a history of the Arctic moss flora Norton G. MILLER Hypnum plicatulum in eastern North America W. B. SCHOFIELD Observations on Sphagnum fitzgeraldit Lewis E. ANDERSON What is Lejeunea trigona? S. RoB GRADSTEIN Mosses as useful plants A. J. SHARP Studies on Geocalycaceae VII. Subspecific differentiation of Chiloscyphus semiteres together with further refinements in Chiloscyphus (s. lat.) JOHN J. ENGEL A revision of Ochrobryum (Leucobryaceae) Bruce ALLEN Reproductive biology of the rare “copper moss” Mielichhoferia mielichhoferiana A. JONATHAN SHAW, N. J. NiGuipuLA & T. M. WILSON Beeveria (Hookeriaceae), a new genus from New Zealand ALLAN J. FIFE = 4] a FOREWORD I can’t remember when I first met Howard Crum, but after my initial year of graduate study at the University of Michigan I went to the Biological Station at Douglas Lake and there took two of Howard’s courses, Bryophytes and Lichens, as well as Ed Voss’s Boreal Flora. Although I had had some preliminary contact with bryophytes, I had never sat down and identified them myself. Suddenly [ was in class six days a week, and spending evenings and the one “free” day secluded in a dank laboratory keying out moss after moss. Howard had said that his classes always found lichens easier than mosses, and it was true, at least in this pair of classes. It may have been that Howard knew more about mosses and thus taught more, but at least in part lichen species indeed have a more distinctive aspect than moss species. Why, then, would anyone spend their life studying a group of plants with little apparent distinction? The late William Campbell Steere, one of How- ard’s earliest mentors, was once asked “What good are mosses, anyway?” His response, “They’ve supported me and my family all these years,” has become a classic. However, it is more than the lure of easy money that seduces the unwary into bryology. Whether it be in the field, on hands and knees and peering through a hand-lens, or in the laboratory, gazing through a microscope, mosses have an intrinsic beauty which can be a true Siren to those unafraid to use magnification as an intermediary. I don’t know anyone who, from childhood, decided to study mosses. They are too subtle for children, or for most adults for that matter. They are a cultivated taste, but an addictive one. It is to Howard Crum that I owe (or blame) for developing my own fascination with this group of plants. As a beginning student of mosses, I would work through great piles of unidenti- fied mosses, doing my best to put names on them. Many were from exotic lands (and in those days I had never ventured into the tropics), and identification was further encumbered by guides in German or French. I remember as clear as if it were yesterday, because it happened so many times, that I would struggle for hours with a single specimen until I had exhausted what seemed to be all possible ave- nues. I would then trundle myself and the vexatious specimen from my basement cell up to Crum’s office for assistance. After briefly glancing at the specimen and my prepared slide he would look over, roll his eyes, and say “Everyone knows what that is!” The obvious implication being, everyone but me! Nevertheless, my atten- tion was held so that next time I would know. I dreamed of the day when I could reciprocate. It only happened once as a graduate student, but it was sweet. In June of 1975 I was going to visit another Michigan graduate student, Joe Beitel, on Long Island, New York. Howard asked me to look for Sphagnum macrophyllum and send it to him as living material. The name meant nothing to me at the time, so | went scrambling to the herbarium to see if I couldn’t learn to recognize the aspect of the plant. In the pine barrens of Long Island I was delighted to find what I felt surely was S. macrophyllum. | put it in a plastic bag and mailed it back to Ann Arbor. Upon returning to Michigan several weeks later I went to Howard's office and there he had the Sphagnum, floating in a bowl of water. The first thing he said was “Thanks for the Sphagnum henryense.” A little disappointed, but not willingly to totally retreat quite yet, I asked if he had examined it with a microscope. He admitted that he hadn’t but didn’t think he needed to. Nevertheless, I asked that he 2 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 just look at it for my sake, and he did. It was Sphagnum macrophyllum! Howard mumbled some excuse about not actually knowing the species in the field and never having seen it alive. It was OK. I would never have to endure the “Everybody knows... .” again. From the very beginning Howard Crum and | got along well. We both have a similar sense of humor, and although my expression of it is a bit more tactless and crude, Howard never fails to laugh. (Although usually voting Republican, he ts a closet liberal.) We both have what might be called an addiction to gossip. At the Biological Station, Howard had a saying, “If you havent heard a good rumor by noon, start one!” Whenever I call him, he always wants to be sure he has exracted all the juicy tidbits floating around the bryological community. Early on, Howard became more than just a colleague or mentor, he became a friend. It may seem odd, then, that up until about five years after leaving Ann Arbor, and taking up resi- dence at the New York Botanical Garden, I continued to call him Dr. Crum. There was no implication of formality, it was just the name I used. Finally, in August, 1984, I wrote to ask if I could use his Christian name, since it had become awkward in talking to colleagues about a man I knew better than they did, even if they used his first name. Howard wrote back: “No, since you are no longer so low, I do not mind your calling me by my first name. I understand the awkwardness. I went through it, for example, with in-laws!” Although we continue, as colleagues, to send specimens back and forth for opinions, our friendship has grown. I have shamelessly telephoned him at home to see how his recovery is progressing, from one affliction after another. I got to know his children when they were both young (and Roger a real hellion), so it’s still amazing to me that they are grown and married. I have talked with his wife Irene any number of times when Howard was ill, and now I have been conspiring with her about arrangements for the party we have planned for Howard’s 70th birthday. hus, in late 1990-early 1991, at the mecting of the American Bryological and Lichenological Society at Wakulla Springs, Florida, Dale Vitt and I talked about what we should do to celebrate Howard’s 70th birthday. We decided on a combina- tion Festschrift and party at the Biological Station. Unfortunately we were not able. due to space restrictions, to open up these pages to all those who would have liked to contribute. For that I apologize. We decided only to invite manuscripts from former students and close colleagues. Although in the midst of editing the massive Moss Flora of Mexico that Howard has been shepherding through publication, | was pleased to take on the editorial tasks engendered by this Festschrift. It provides tangible evidence of my affection, admiration and respect for Howard Crum. On this occasion of Howard’s 70th birthday and approaching retirement, I, and the entire bryological community, wish him congratulations and many more happy years of bryology. William R. Buck Contr. Univ. Mich. Herb. 18:3-38. 1992. A TRIBUTE TO HOWARD CRUM Lewis E. Anderson Department of Botany Duke University Durham, NC 27706, U.S.A. In his tribute to me in a Festschrift honoring my 70th birthday, Howard Crum (1985) wrote that “it is difficult to write about a friend or to evaluate his career when he is still capable of making rebuttal.” This was a prescient statement be- cause, now, ten years later, I have been asked to write a comparable piece for a Festschrift celebrating his 70th birthday. There is not really much to rebut, how- ever; for the most part he was kind and generous to me in his tribute. To my surprise, he used discretion and tact in selecting stories about me and his exaggera- tions were generally within the established boundaries that we tolerate from each other. I will try to be equally discriminating and cautious in the things I write about him; nevertheless, I will probably touch a few tender spots. I think he will expect that and I wouldn’t want to disappoint him. Because we have known each other for so long and have worked together so closely over the years we each have thick folders of incriminating letters from the other that reveal items and thoughts about colleagues and others that neither of us cares to have disclosed. Most of our letters contain, in addition to the business of the moment, ridicule of each other, continuing arguments, attempts at humor, and a large amount of foolishness. As Howard pointed out in his article about me, they contain material that should not be taken out of context. So, I, too, will refrain from mining those letters for that sort of material. A short while after the aforementioned tribute to me appeared, I received a letter from a friend of his who stated that he learned more about Howard Crum from the article than he did about Lewis Anderson. He probably got this impres- sion because our lives for the past 44 years have been closely intertwined both socially and professionally. A great deal about me will, of necessity, surface in the present article about him. After all, during these years we have collaborated on 23 papers, 10 fascicles of exsiccati and the two volumes of Mosses of Eastern North America. The latter work began in 1959 and continued more or less continuously until it was published in 1981. We have been guests in each others homes countless times; we have watched our respective children mature and drift away; we have spent endless hours together in the field and even more hours working side by side in the herbarium, passing slides back and forth; we have had countless extended telephone conversations; and, as noted above, we have carried on a prolific corre- spondence over the years. We have exchanged philosophies and concepts and we have learned a great deal from each other. We share the same kind of humorous outlook on life and we have long been able to overlook each other’s shortcomings and imperfections. We probably understand each other as well as two friends and colleagues can. Furthermore, we both genuinely enjoy our work and we have had fun in our careers and probably wouldn’t do anything much different if we had a chance to relive our lives. Our first meeting was in 1948 at the University of Michigan, where Howard was 4 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 FIG. 1. Howard Crum at age 7 (left) and about ten (right). a graduate student working with Bill Steere. Howard was among a group of stu- dents and myself who were invited by Bill and Dorothy for one of their famous evenings at home. As always at the Steere home, it was a fun evening, with Dorothy plying her guests with food and drink, and Bill cleverly maneuvering the conversa- tion in and out of a delightful mix of light and humorous anecdotes and serious bryology. Students learned without knowing it. Bill had already told me that How- ard Crum was by far the best student that had come his way. He cautioned me that Howard was shy and quict but that underneath there was a deep and penetrating intellect. Steere was right. IT hardly knew that Howard was there that night. He scarcely said anything the entire evening; he listened and learned. Later, he told me that he was in awe of Steere and me. How our close friendship and working relationship has survived these many years 1s a puzzle to most of our friends and acquaintances. I am not sure that even we understand it. We criticize each other’s opinions, ideas, concepts, and even personalities, openly, loudly, and with brutal frankness. We often disagree and our arguments range from playful to vehement. Rarely do either of us win. Those within earshot of our snipes at each other are often astounded, fearful that we are about to end a friendship. We both enjoy telling embarrassing stories about each other. Our colleagues are amazed that our collaboration has endured for so long and that it has been so productive. I think our friendship and collaboration is sustained mainly because we share a mischievous, derisive sort of humor. We work together well because we are comfortable with each other. We say outrageous things about and to each other without malice. I can’t recall that we have ever been really mad at each other, although Howard hides his feelings much more than I do. Howard is easily bored. When we work together, he will often say, “Anderson, you bore me,” then tidily put away his tools and depart or demand that we work on something more interesting. Another of his favorite expressions is, “Anderson, | understand you a lot better after I’ve had a couple of beers.” After a time, in a conversation group, he has been known to say, “You people bore me,” and promptly leave. Such frankness can be puzzling to some, but however true the 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM ) expressions of boredom may be, they are always uttered in jest and reflect partly his puckishness. Mostly, however, his real need is to get away from people. It was a side of Howard that Bill Steere could never understand. When they were working together at Stanford, for example, Howard’s need for privacy would on occasion drive him to absent himself without telling Steere or anyone else where he was going or when he was coming back. He would just disappear. Howard might take the bus to Monterey, to the Big Sur, to Muir Woods, to Sonoma Valley or elsewhere and explore the wineries, or to any place that he could be alone. Wilf Schofield spent the summer of 1957 collecting with Howard in the Cana- dian Rockies and the Yukon and encountered the boredom trait in Howard. In Wilf’s own words: “Peggy [Wilf’s wife] joined us in the Yukon for a month. We tented at Mile 1022 on the Alaskan Highway and did most of our collecting there. Peggy would go fishing near a culvert and bring back our meals while Howard and I explored another collecting area. I recall an incident when Howard told us “You folks bore me; I’m going to go away for a few days.’ Since Peggy and I were newly married, I’m sure Howard wanted to give us some time by ourselves, but I suspect that to a degree, we did bore him! He returned refreshed, bearing, among other things, fresh Splachnum luteum, which I had not seen before”. Students and colleagues who have never met Howard often ask me, “What is Crum like?” Even those who are acquainted with him will sometimes ask me, “What is Crum really like?” I only know one Crum: highly intelligent, charming, gracious, courteous, kind, polite, tasteful, cultured, sensitive, and possessed of a tremendously individualized sense of humor. As I have already noted, he is a listener, not a talker. I have never known him to interrupt or break into a conversa- tion and he scarcely ever raises his voice. He is articulate, but soft-spoken with a slight drawl, enough on occasions to be mistaken for a Southerner, which he does not consider flattering. He is not very effective in an oral argument; when pushed into a corner, he simply clams up and grins at you, which takes all the fun out of the dispute. Howard’s listening qualities make him a very effective student advisor. The University of Michigan soon discovered this gift and tapped him for the onerous task of freshman-sophomore counseling. For years he has spent an afternoon or more each week in these counseling sessions. I have learned not to telephone him when he is in an advising session because then he is strictly unavailable to anyone. Those students admire and appreciate him because he is genuinely interested in their lives and problems and he gives them a sympathetic ear. Besides, Howard has an ingrained curiosity. In 1978, he was honored with the prestigious Ruth M. Sinclair Memorial Award for Freshman-Sophomore Counseling, a tribute to his effectiveness. Along these lines, Jerry Snider reminded me of a story that Irene, Howard's wife, tells about one of Howard’s many hospital stays in which he shared a room with an interesting but uneducated man. Howard was very nice to the man, prying with his usual curiosity into every facet of his life. The man was appreciative of Howard’s interest and they got to know each other well. The man left the hospital before Howard and, some weeks later, Irene met him on the street. He asked Irene about Howard’s health. She replied that he was in the field (meaning with his class ona field trip). The man, looking very downcast, said, “Oh, too bad; oh, how sad.” having interpreted Irene’s comment to mean that Howard had died and was now in Potters iBicid.” Next to coffee, beer is Howard’s favorite drink, although, during his excursions 6 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 G. 2. Howard Crum in uniform during WWII. He served in the Middle East in an intelligence unit in the United States Army Air Force, 1942-1945, in the southern states, he developed a mild fondness for sour mash bourbon. In the South the tiny, almost microscopic larvae of the chigger or “red bug.” are very prevalent in summer. They attach themselves to animal hosts that brush against plants on which the larvae lie in wait. Plant collectors are especially vulnerable as they walk through graminoids and brush, generally at a slow and deliberate pace. The tiny larvae are transferred to shoes and clothing from which they crawl and attach themselves to skin surfaces, causing intense itching and pink or red spots. Humans vary, apparently, in their susceptibility to these pests, and they seem to 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM Z FIG. 3. Howard Crum during undergraduate days at Western Michigan Teachers College (now Western Michigan University). have a particular affection for Howard. Reasonably effective preventive chemical applications are available but he never seemed to learn how to apply them. On one of our collecting trips to Florida he accumulated a particularly heavy infestation, claiming that he had counted more than 100 of them on his body, some of which he claimed were inhabiting rather embarrassing places. When a motel clerk asked us what we intended to do that evening, Howard replied that he didn’t know what Andy was going to do but he intended to retire to his motel room, drink bourbon, and scratch his bites. I quickly explained to the clerk the kind of bites he was speaking about before we were tossed out of the place. Howard’s puckish humor inevitably surfaces at appropriate and often unex- pected times and generally cuts right at the heart of the situation. An example is a story that Jack Sharp tells with great relish. Jack, Howard, and the late Bill Fox were collecting in Mexico in 1951. One of their stops was at the Rancho del Cielo, Tamaulipas (Frank Harrison’s home), high on the Eastern Escarpment above Antiguo Morelos. They had rented horses and a guide from an Indian village across the river and started toward Rancho del Cielo late in the afternoon (too late for the distance they had to travel, according to Howard’s version). It turned out that there were not enough horses for all of them and their luggage and collecting gear. Howard, being the youngest, volunteered to walk. The weather was warm and humid, and the trail was very steep and rocky. As they ascended the high ridge, the trail become rougher and, as darkness approached, one of Howard's legs began to give him trouble. Finally, after several stops because of Howard’s leg, Sharp, be- cause he was accustomed to such trails in the Smoky Mountains, suggested that they trade places, which they did. It soon began to rain and as they gained altitude, fog settled in and visibility became zero. Sharp stayed on the trail by holding on to the horse’s tail. It continued raining, the fog became even denser, and the guide, 8 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 obviously having difficulty keeping on the trail, called a halt to the procession while he did some scouting. After what seemed to be a long period of silence, out of the black dismal night came Howard’s voice, loud and angry, “What am I doing here? | hate botany!” On our field excursions I always drove the vehicle, even if it was Crum’s. He will admit under duress, or perhaps several beers, that his driving is somewhat erratic. Those who have ridden with him would call that an understatement. After riding with him in a van at the University of Michigan Biological Station for the duration of his bryology course one summer, the class gave him a Saint Christopher Medal! One summer he wrecked a National Museum of Canada van by crashing into the rear of a car ona straight, level, prairie road. I was never able to wring the particulars out of him. All he would ever say was that he was not picking his nose at the time of the accident. I never would have even learned about the accident but for the fact that he was on his way to Winnipeg, where he planned to leave the van, take the train to Fort Churchill and join Wilf Schofield and me (later, Bill Steere joined us for a while). At the end of our stay in Fort Churchill, he and I rode the train back to Winnipeg to pick up the van and drive to Ottawa. He gave some lame excuse that mechanical trouble had forced him to leave the van east of Winnipeg and that we would have to take a bus to this small town. Of course, there, I learned that in the interim, a garage had made the necessary repairs. In his account of this trip (Crum 1985), these details are omitted. Howard has had at least two other narrow escapes in automobile accidents (there may have been others that I don’t know about), but I hasten to add that he was not driving in either of these. Remarkably, both accidents occurred in Tennes- see in different years and both happened when he was returning from collecting trips to Mexico. In 1949, he joined Bob Wilbur, a fellow graduate student at Michigan, ina joint collecting trip to Mexico. They were accompanied by Bob’s brother, who was an engineering student with little direct interest in botany. His and Howard's religious philosophies differed widely, and, according to Bob, they argued almost incessantly throughout the trip. On the return trip, during a particularly high-pitched moment in their argument, with the brother driving, the car ran off the road and crashed into a side ditch. Bob blamed them both equally for the accident. Returning from Mexico with Jack Sharp and the late Bill Fox in 1951 (the same trip referred to above), nearing Chattanooga, with Fox driving, Sharp sitting beside him and Howard in the rear seat, Fox attempted to pass to the right of a stopped milk truck. The right tires left the pavement and the van flipped over on its side. As soon as he saw that he and Fox were uninjured, Jack looked back and was alarmed that he didn’t see Howard. Presently, however, they were relieved to see him emerging from a complete cover of black beans in the corner of the van. Before leaving Mexico, they had bought ten kilos of dried black beans and had placed them on the shelf behind the back seat. The accident ruptured the bag and Howard, unharmed, was buried in the beans. In 1953, Virginia Bryan, then a graduate student with me at Duke, and I spent most of the spring semester at Stanford University working with Bill Steere on chromosome investigations of California mosses. Bill had obtained one of the very first grants from the newly established National Science Foundation to support the chromosome project. Howard had gone to Stanford after receiving his Ph.D., in 19ST, to work with Bill on, among other projects, a flora of Puerto Rico (Crum and Steere 1957), and to help with the identification of Alaskan and other speci- 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM 9 4. Howard Crum as a beginning bryologist, in the field (left, 1946) and at the microscope FIG. (right, 1947). mens that Bill had accumulated over the years. Howard was in his last year there when we arrived. At that time Stanford was a very active place, bryologically. Wilf Schofield, Ed Ketchledge, and Grace Iverson were graduate students working with Bill. Gilbert M. Smith, whom Bill had succeeded, although retired, was still very active. He was revising the bryological chapters of his Cryptogamic Botany text and he and Howard had developed a particularly nice and mutually profitable friendship. Douglas Houghton Campbell, who was Smith’s predecessor at Stan- ford, lived on the campus when Virginia and I arrived, but was not active; in fact, he died shortly afterwards. Thus, Campbell, Smith, and Steere, three giants of cryptogamic botany who had occupied successively the same professorial post at Stanford were together at Stanford for a period of time. Having been a member of the faculty there since Stanford opened in 1891, Campbell lived to be 95. In 1959, however, Steere resigned to become Director of the New York Botanical Garden and he was not replaced. The bryological tradition at Stanford, alas, abruptly came to an end. Howard’s three-year stay at Stanford was productive and enjoyable, but it was not without certain frustrations. He had gone to Stanford with high expectations of working closely with Steere and learning from the one-on-one relationship he envis- aged. When they were together at Michigan, Steere was chairman of the depart- ment and laden with administrative responsibilities. Their contacts there were brief, intermittent, and unpredictable. Among Steere’s reasons for leaving Michi- gan for the post at Stanford was a strong desire to escape the administrative burdens of chairmanship. He claimed that he had no time for research at Michigan, and Stanford promised that they would relieve him of all administrative responsibilities. But, little did they know the strong attraction that administration held for Bill Steere. In a letter to Oswald Tippo, though, Professor H. H. Bartlett, who knew Bill well, summed it up correctly: “[Steere] is a natural-born administrator who can no more keep away from it than he can from his research” (Crum 1977). At Stanford, Steere attracted administrative duties like iron filings to a magnet. It was not long after Howard arrived that Bill, through no fault of Stanford, became as busy, administratively, as he had been at Michigan. (Within a short time he became 10 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 FIG. 5. Howard Crum as a graduate student in botany at the University of Michigan, 1949. Dean of the Graduate School!) As Howard put it, “[Steere] was too busy to help me interpret [species] or to join me in close collaboration” (Crum 1977) At Stanford, Howard worked in what was called the “back room,” which was Steere’s laboratory, accessed more or less through Steere’s office. The cytological lab, where Virginia Bryan and I worked, was in an adjacent building, but the four of us (and often others) gathered in the back room at mid-morning and mid- afternoon for coffee. These were about the only times that Howard could capture Steere long enough to discuss problems of identification and interpretation encoun- tered in several investigations they were carrying out. One of these occasions pro- duced a Steere expression that Howard and I treasure to this day. Howard had laid aside several sterile specimens that he was unable to identify and hoped that Steere could suggest names. Particularly challenging was a ditrichaceous-looking moss from Panama that had baffled Howard for some time. He reviewed a long list of characters that he had noted, after which Steere turned to him and in all seriousness asked, “But Howard, have you checked the rhizoids?” Of course, Howard had not checked the rhizoids, and there the matter rested! [The moss turned out to be Garckea phascoides (Hook.) C. Mull.| Since then, when either of us sends the other a specimen for verification or hopeful identification, the reply is often, “I don't believe you checked the rhizoids!” Thereafter, when Bill would overhear us use this expression, he would always appear puzzled, then give us his characteristic quizzical look, followed by “Oh, really?” Although Bill Steere had a vigorous sense of humor and could and did spiel off one joke after another, they were never jokes on himself. His shortcomings, blunders, and human frailties were not amusing t ~ himself. The piantord ve was os and Sreniee for all of us. More- evenings ae the cuisine a splendid wines in the aa fine resturants in the 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM 11 area. In a small group such as this, Howard is at his best, and it was at these gatherings that I got to know him well. I discovered that, unlike Bill Steere, he could laugh at himself and that, indiscriminately, he could combine fun and serious work. Moreover, we liked each other’s company. Howard put it best: “Our friend- ship has been based on mutual interests, scientific and cultural, as well as mutual tolerance for personal foibles, his and mine. We have a similar way of viewing the world and each other, with puckish good humor” (Crum 1985). Thus, when the opportunity came to collaborate, scientifically, we felt that we could work together pleasantly and profitably by combining somewhat diverse talents and knowledge. Howard spent the summer of 1953 in Alaska with Bill Steere, based at the Arctic Research Laboratory at Point Barrow. They collected together at Barrow and Cape Lisburne and Howard collected extensively at Meade River and the Anaktuvik Pass in the Brooks Range (Crum 1977). Actually, Bill did little collecting that summer. Using the techniques we had worked out at Stanford, he counted chromosomes from material brought in from the field by Howard. This was How- ard’s first field experience in the High Arctic and the time he had spent identifying Steere’s Alaskan material at Stanford provided him with a background knowledge of the moss flora there that greatly enhanced his collecting effectiveness. He also learned to recognize capsules in the proper meiotic stages for chromosome study. Over the years he has sent me much living material for cytological study and, together, we published a long paper on the chromosomes of a large number of mosses of the Canadian Rocky Mountains, based entirely on collections of his that were shipped to me at Duke. Howard’s post-doctoral with Steere ended in 1953 and, after much job-hunting, he accepted a position in the Department of Biology of the University of Louisville. He stayed only one year, however. A position opened at the National Museum of Canada when I. Mackenzie Lamb accepted the directorship of the Farlow Library and Herbarium at Harvard University. Howard applied for the position vacated by Lamb and was appointed Curator of Cryptogams, beginning in the fall of 1954. In the meantime, I was able to obtain meager funds to support Howard at Duke for the summer of 1954. He helped with the determinations of mosses that I had collected on a previous summer’s excursion to the Ozark Mountains. He was grate- ful for the financial support, but when his first check arrived he was stunned to discover that he was being paid out of a petty cash fund. To this day he reminds me of this humbling experience. Rudy Schuster was in residence at Duke then and it was a memorable summer. Rudy occupied two small rooms adjoining the then small herbarium. Olga, Rudy’s wife, did much of his typing directly across the hall from his work room. I know of no couple more devoted and respectful of each other than Olga and Rudy. Their loud, reciprocal arguments, replete with mutual denuncia- tions, strong profanity and vilifications when working together, however, are notori- ous. The exchanges are totally harmless, but this was Howard’s first experience with the Schusters and, on occasion, he would get so nervous he would have to leave and take a walk Generally, we used Grout’s flora in our determinations and its deficiencies became even more apparent as we worked through the Ozarkian specimens. We soon came to the conclusion that it needed to be revised. The long discussions and experiences we had that summer led eventually to Mosses of Eastern North Amer- ica. First, we decided simply to revise and tidy up Grout’s work, a project that we estimated would take no more than three years. Very quickly, we abandoned the idea of a revision of the Grout treatment and decided on an original flora scaled 12 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 a ay on Mont Tremblant, Quebec, during the Pre-congress foray, LX International FIG. 6. A rainy ¢ Botanical Congress, August, 1959. Howard organized and led the foray. Harry Williams, Howard Crum, Seville Flowers, Fred ction (left to right). down to include only eastern North America. We realized that such a flora would take more time, but even so, we couldn't have realized that we were beginning a 27- year project. During that eventful summer of 1954, I discovered many of the talents and skills that contribute to Howard’s scientific and scholarly proficiency. I found that he has a broad liberal background in arts and sciences, especially in classics and languages, which is quite unusual for someone trained in the sciences. His tremen- dous command of the English language is reflected in all his writings. A natural editor, he cannot refrain from tinkering with careless and slovenly writing. He attacks manuscripts with a sort of vengeful glee. When he spots an ungrammatical lapse his lips curl into a determined but pleasing grin. Thus, when Bill Steere called me in July to report his appointment as Dean of the Graduate School at Stanford and to tell me that he could not possibly edit The Bryologist any longer, he and | agreed that Howard would make a good editor. When I broached this to Howard, he was dumbfounded. Out of graduate school only three years, he was stunned at the prospect of editing a professional journal at this early stage in his career. | reminded him that all the while he was at Stanford he had more than assisted Steere in editing The Bryologist; he had handled a lot of it on his own. At the time, I was acting as Business Manager of the American Bryological Society and the journal was being published in Durham. With some prodding from Steere and after taking Howard to the publishing firm, where he met the printing staff and publisher and saw the production operation, he was persuaded to take the job. It was not difficult LODZ ANDERSON: A TRIBUTE TO HOWARD CRUM Iss} FIG. 7. The hair gets thinner. Howard Crum in 1952 (left) and 1970 (right). to convince the executive committee of the Society to name him the new editor. He began immediately, editing Number 3 of Volume 57 (September, 1954), while he was in Durham that summer. He was able to work directly with individual members of the publishing firm and gained valuable experience that would help him in future editing. By chance, I was the author of the first paper in that number and I do not need to add that he tackled all its deficiencies with delight. As Bill Steere and I expected, Howard became an excellent editor. He edited with a firmer pencil than Steere and may have ruffled the feathers of a few authors with his grammatical and stylistic tinkering, but the quality of papers improved during his editorship and The Bryologist became a better journal. Among other changes, Howard shifted the journal to a quarterly in 1962. Founded by Abel Joel Grout in 1898, it had begun as a department in the Fern Bulletin, which was a quarterly. In 1900, however, The Bryologist became an independent journal and, in 1902, it went to six numbers per year, where it remained until Howard’s change. In the December, 1955, issue he introduced a valuable “News and Notes” section. Its chatty and informal style has continued through a number of editors to the present. He kept the editorship for two terms or a total of eight years and was succeeded by William Louis Culberson in 1962. During the summer of 1954, Howard became a fixture in the Anderson house- hold. He was 32 and unmarried. Although he lived in a dormitory room on the Duke campus he spent much time in our home. There were five children, three girls and two boys, seven to twelve years old, including a pair of twins. It was a busy, noisy place, and Howard was fascinated, especially with the girls, seven, eight and eleven. One evening when he was having dinner with us a girlfriend of our oldest boy, Philip, was also a guest. She and Philip were 12. There was a bouquet of snapdragons on the table. Our little guest was sitting next to Howard and at some point asked Howard a question about the snapdragon flower. Always the ready educator, he began explaining the structure of the snapdragon flower to her at a technical level that was beyond anything the poor girl could comprehend. Also, it soon became clear that she knew very little about the sexual life of the flower or for that matter, animals or humans. As her questions became more pointed and pre- cise, Howard found himself in the awkward position of having to explain sex to this innocent child. He made a number of clumsy and embarrassing starts, but he was totally unprepared for the challenge that soon unfolded. He looked to all of us in I4 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 despair, expecting help that never came. He never forgave us for the silence and for giving him no help whatsoever. Finally, my wife, Pat, came to the rescue by telling the girl that the flower produced seed, which, when planted, produced another plant. That satisfied the youngster and Howard was finally off the hook. After he was married and had children of his own such conversations with them would become more natural. Howard told me rather recently of an incident that occurred when his children, Roger and Mary, were in their pre-teens. Roger’s elementary school teacher (and presumably others) had given a sex education lecture to his class and some of the parents were apprehensive about its content. The parents were invited to come to the school one evening and listen to the lecture as it was given to the students. Roger’s parents attended. When they returned, Roger asked his father how he liked the lecture. Howard replied, “Well, it was very instructive and educational, but they didn’t tell you how much fun it is!” This episode illustrates the kind of open and frank relationship he always kept with his children, even when they were very young. He always treated them as adults. In early September, 1954, we went to the Gainesville, Florida, AIBS meeting with Heinie Oosting and Terry Johnson. [Howard, in his account of this trip (1985), incorrectly included Bill Culberson in the group.] Terry and Howard were graduate students together at the University of Michigan and knew each other quite well. Howard, in his account, refers to“. . . night disturbances resulting from sopho- moric humor,” without further elaboration except that “The situation would not have arisen except for Heinie’s cautious use of the dollar, which resulted in our sleeping five [actually four] to a room.” Terry and Howard shared one bed and Heinie and I the other. Howard is a light sleeper. After retiring, he would stay awake for a long time, get out of bed often, and walk around in the room until he felt sleepy. Terry went to bed early, slept soundly, and arose at an unearthly hour, such as four A.M., wide awake and ready to go! As Heinie complained, by the time Howard quit getting up and bumping around in the room, Terry was up, shaving and showering. On one particularly bad night, Howard, still wide awake at mid- night or one A.M., gave the sleeping Terry several stiff jabs with his elbows. Terry sat upright, inquiring, “What’s wrong, Howard?” Howard said, “Terry, I’ve just wet the bed.” Terry jumped out of the bed, knocking over a table and lamp and waking Oosting and me. Of course, Howard had not wet the bed, but there was little sleep for any of us thereafter. Sophomoric humor? Almost immediately after arriving at the National Museum Howard made plans for Canadian field work, which the Museum encouraged. He spent succeeding summers collecting in the Gaspé and Laurentian Mountains, the Bruce Peninsula and Niagara Escarpment, the Canadian Rocky Mountains, the Yukon, and north- ern Manitoba. In 1957, he and I made a firm commitment to work on the moss flora of eastern North America. We applied for and received a grant from the National Science Foundation and very shortly signed a contract with Columbia University Press to publish the work. I spent most of the winter and spring of 1959 in Ottawa. We worked together, side by side, each with microscopes, passing slides back and forth and trading jibes and information. This was the real beginning of the flora. Howard had just bought and moved into a new house when I arrived and I occupied one of the bedrooms during my stay, which extended into the spring. It was during this stay that Howard claims I gave him stomach ulcers, which ultimately necessitated rather drastic surgery. This was followed by a siege with his gall blad- der, which I maintained was brought on by his greasy cooking. Actually he was a a) 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM good cook and produced some real culinary treats. But, unfortunately, he was fond of fatty foods, including something the English call fried bread, which consists of soaking and frying slices of bread in left-over bacon grease. I found the result utterly loathsome, and rejected it on sight. e brightest parts of our working days at the National Museum came when Irene McCarthy stopped in and visited with us. She was Administrative Secretary in the Department of Botany and we looked forward to her visits. Attractive, viva- cious, cheerful, spirited, extraordinarily intelligent (much smarter in a non- academic way than Howard), incredibly organized, the neatest and tidiest person I have ever known, she managed the affairs of that department with stern, unyielding efficiency, and with a tremendous sense of humor. She was not intimidated in the slightest by a scientific staff of all-male prima donnas, including the chairman, A. E. Porsild, a noted authority on arctic plants. She radiated brightness and cheerfulness in the otherwise gloomy atmosphere of the then old and stuffy museum building with its high, multistoried lobby overcrowded with tall, dusty totem poles. I soon noticed that she and Howard were quite friendly. They enjoyed sharing gossip; they joked and kidded each other; they had formed an alliance against staff members they disliked; they obviously liked each other. He called her McCarthy then, never Irene. Of course, I joined the good-natured bantering, which provided intermittent relief from the daily tedium of looking at herbarium specimens, super- vising a recalcitrant but gifted artist, and constant decision-making. At first I thought something might be developing between them, but soon dismissed it because I knew they were not seeing each other except in the herbar- ium. That was certainly no place to carry on a romance. Howard did ask me once if I thought McCarthy would make a good wife. My answer was a stream of praise for her and an enumeration of all the splendid qualities that I had observed during our brief acquaintance. I reminded him, however, that she was too smart to marry him, a deep-rooted, fastidious bachelor, whose habits and mores were indelibly fixed. As it turned out I was quite wrong. They were married on December 26, 1960, after what has to be one of the strangest courtships on record. She recently filled me in on the particulars, and they are worth recording. At the time, the fall of 1960, the Canadian government was in the midst of a campaign to encourage its employees to become bilingual and were providing free language courses for them. On this particular day Irene and Howard were discussing the possibilities of enrolling. Irene had already decided that she didn’t want to register for a French course, which was the principal language being pushed. Rather, she told him, she wanted to be different and intended to register for a biology course, which might help her in her work. Howard replied, without any hesitation or change in inflection, “Well, if you want to be different, then why don’t you marry me?” At first she thought he was joking, but he was serious. Moreover, he wanted to get married that weekend! She would have none of that, so they were married the day after Christmas, with all of the appropriate rituals of a traditional wedding. Irene tells me that when Howard first joined the museum staff she thought he was pretty cute. Presently, she began to make a conscious effort to attract his attentions toward her. As time went on, however, she saw no results of this encour- agement and after a couple of years decided that it was a hopeless cause. Although they established a pleasant relationship at the Museum, it never went beyond that. He never dated her once; he didn’t take her anywhere, not even to a movie; they never had lunch or dinner together; they never even took a walk together. It is, 16 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 FIG. 8. Irene and Howard Crum, married and intent on starting a family. therefore, no wonder that she had given up. When the off-handed proposal finally came she was stunned and she could scarcely wait to tell her mother. After dashing into the house and loudly exclaiming that she was getting married, her mother, having forgotten about Howard, replied, “To whom, for goodness sake?” I was right about one thing, though. Irene turned out to be the kind of wife that T envisioned and much more. She is the only woman that I can think of that could have sustained a happy marriage with Howard. They have a fantastic marriage and a mutually satisfying home life. Pat and I have been guests on numerous occasions over the years and have watched their children mature. [rene is a wonderful cook. Her specialties are desserts, and: she and I share a love for them. We also share a struggle with our respective weight levels. Irene is an impeccable housekeeper. She is even more fastidious than Howard. Her house is as clean as an operating room in a hospital. She has looked after him, [am sure, as well as his mother did. He has survived an unbelievable number of life-threatening illnesses principally because of her agressive vigilance. The two Crum children, Roger and Mary, are highly talented. I don’t know of any parents who have enjoyed and relished their children more than Howard and Irene. The children have responded in kind. Roger, now married to Robin, re- ceived his Ph.D. in art history from the University of Pittsburgh in 1991, and is now Assistant Professor at the University of Dayton. Mary, a music major with a degree 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM 17 FIG. 9. Irene and Howard Crum learning to deal with toddlers Mary and Roger (above); and o Mackinac Island, summer, 1974, Mary, Irene, Linda (Mrs. Jerry) Snider, Roger, Howard (left Caane from the University of Michigan, teaches music in the public schools (she is an accomplished clarinetist, and plays beautifully) and is married to Brian Scholtens, who, in 1991, received his Ph.D. in zoology from the University of Michigan, where he now holds a teaching position. A quote from a recent letter from Roger provides a much better insight into family life than I could possibly write. “One thing that I’ve always enjoyed telling people is that, while my father is a botanist, there have never been any plants in our home. You can well imagine why this is so, given that my mother ts definitely in charge. Since she grew up on a farm, 18 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 FIG, 1 . A beautiful bride; Howard, Mary Crum Scholtens, Irene. dirt of any nature in the house represents something with which to wage constant battle. “There was never any bryological shop talk at our home, even though hours upon hours of bryological research and writing occurred on my father’s lap in the living room. My father could work while any sort of noise, playing or watching of television, was going on. He could work through it all without being disturbed in the least. It seems to me that my father has always worked best at home, in his living room chair (and at times with the two of us in the chair with him), with a manuscript set on a kitchen cutting board on his lap. In fact, as you well know, since Dad has been seriously ill so many times, a good deal of his work has by necessity been carried out at home. “Finally, bryology has been Dad’s thing and Dad’s thing alone. He has never spoken of his academic stature or success, and it was years before I understood the importance of his contributions to the field. He showed by his example the joys of an academic life style but he never urged Mary or me to become interested in science or botany in particular. His major interest was that we pursue that which interested us most. As things have turned out, this has been music for Mary and art history for me.” Howard Alvin Crum was born on July 14, 1922, at Mishawaka, Indiana. His mother, Eunice Eva Crum, died in 1949, at age 51; his father, Earl Earnest Crum, called Jack by his friends, died in 1964, at age 73. The father worked as a riveter for the Studebaker automobile factory in South Bend, Indiana, during the 1920's and into the early years of the Great Depression. They had four children, Chester, Roger, Howard and Roland. Howard grew up in Mishawaka, except for one year in which his father moved the family to Howard’s grandfather’s farm in Burr Oak, Indiana. That one year on the farm made a deep impression on Howard. For years, 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM 19 FIG. 11. Roger Crum’s wedding. Above, Howard, Irene, Roger, Mary. Below, Howard, Irene Roger, Robin, Mary and Brian Scholtens. 20 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 2. Irene and Howard Crum after Roger’s wedding (left) and a happy Howard (right). FIG. I thought he grew up on a farm, because he spoke so much about farm life. After that year the family moved back to Mishawaka. Howard had a special affection for his mother and speaks of her often. He can get misty-eyed while discussing her. | remember an incident when Pat and I were visiting the Crums in Ann Arbor. I went to the kitchen to mix my five o'clock drink only to find that the ice cubes had frozen into a solid bar of ice. Reaching into the nearest drawer, I pulled out an old, nondescript wooden spoon and proceeded to whack away at the ice to try to break it apart. The spoon shattered into splinters just as Howard walked in. Instead of playing the perfect host and reassuring me that it was nothing, just an old wooden spoon, he gave me a despairing look, and in the saddest voice, said, “It belonged to my mother; you clod, you've destroyed It.’ There was sadness, of course, over the loss of the sentimental spoon, but mostly he wanted to embarrass me. I reminded him of the time that he was admiring a particularly beautiful and expensive mahogany salad bowl at a dean’s house when it inexplicably fractured in his hands. After attending Kennedy School for grades one through eight, Howard went to Mishawaka High School, graduating in 1939. His two older brothers, Chester and Roger, as well as an aunt, had all attended Western Michigan Teachers College (now Western Michigan University) at Kalamazoo, so Howard followed them there, entering in the fall of 1939. He became a German major, but his university education was interrupted by World War I]. He is very proud of the fact that he did not wait to be drafted. He volunteered for the United States Army Air Force in 1942, and served in the Intelligence Division, where he became an expert cryptogra- pher, with duty in North Africa and the Middle East. As his son Roger has reminded me, Howard, in a very quiet, apolitical way, Is a very patriotic person. I have seen him become maudlin upon viewing the American 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM PA flag. The flag means more to him than it does to most Americans. He can get genuinely sentimental about the origins and preservation of the ideals of his coun- try. This may surprise people who do not know him well. Howard returned from military service in late 1945, and spent some time with his mother, who was in ill health in Mishawaka. Like many returning veterans, he began to have some uncertainties about his academic directions. In the spring of 1946, he visited the University of Chicago and interviewed for medical school there. Based on his superb record to that point, they accepted him. He never had any real interest in medicine, however, and eventually decided against it. This was a lucky break for bryology. Finally, he decided to resume his studies at Western Michigan University, but enrolled at the University of Michigan Biological Station, at Douglas Lake, for the summer session as a student disjunct from Western. He registered for Anatomy, taught by C. D. LaRue, and Systematic Botany, taught by none other than Bill Steere. He claims that “the excitement of being there, in a research environment with the best of students and the best of teachers, predestined me to a career in botany” (Crum 1977). When he returned to Western Michigan it is not surprising that he changed his major from German to botany. Somewhat defensively, he brags that he had always had a real love for plants. He speaks proudly of the fact that both his parents knew the common names for most of the plants in their neighborhood and that his mother was a good gardener. Contrary to what one might think, Bill Steere did not introduce Howard to bryophytes when he took his systematics course in the summer of 1946. He discov- ered them the following winter when he returned to Western Michigan. Bill Buck, who knows everything about everybody, told me that Howard’s interest in mosses developed as follows. Howard and another botany major, a girl, by the way, that fall began a routine of walking in the woods in order to strengthen their knowledge of plants. With the onset of winter, the tree leaves and flowers disappeared. The only green remaining were mosses and liverworts. They began collecting them and made attempts to identify them, probably with the aid of Mosses with a Handlens. They may have had some help from Professor Leslie A. Kenoyer, who was the principal botanist there, but it would have been minimal. There is no indication that he knew much about bryophytes. At some point Howard did accompany Kenoyer on a short trip to Mexico. The professor was a generalist with very broad interests, including systematics, floristics, ecology, morphology, anatomy, and pollination biology. He undoubtedly provided some encouragement for Howard’s leanings toward botany, but his interest in mosses was evidently self-induced. Howard be- came an instructor his senior year at Western Michigan and received the B.S. degree, magna cum laude, in 1947. Meanwhile, he was awarded a prestigious Rackam State College Scholarship at the University of Michigan, to begin in the fall of 1947. Curiously, in his application for the scholarship he indicated that he wished to major in plant pathology. After graduation, however, Howard decided to spend the summer at the Biological Sta- tion, where he enrolled in Margaret Fulford’s course in bryophytes. This was the first time she offered it at the Station. Concurrently with her bryology course, he took Professor Gerald Prescott’s course in aquatic flowering plants, for which he originally received a grade of A+. Later, however, Prescott changed it to A—. For Howard, this was an unforgiving act, and he complains to this day about it. When Howard arrived in Ann Arbor he discovered that the University of Michigan did not offer a program, not even a course, in plant pathology. He says 22 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM pare) VOLUME I8 3. Howard, the Ulota Hunter, University of Michigan Biological Station, FIG. hat was a birthday present from his class that year; the raw egg was received in the _ summer, 1985. The mail.” 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM O58 eventually he “blundered into Steere,” which was an “accident of fate.” Of course, he would have known Steere from the previous summer’s course in taxonomy at the Biological Station, so it probably didn’t involve too much “blundering.” In any event, he soon became interested almost exclusively in mosses and began his gradu- ate work as Bill Steere’s student. He entered Michigan toward the end of a great period in bryology there under Steere’s leadership. Steere attracted a host of students immediately before and after World War Il. Howard was certainly the star of the group. Almost immediately he became interested in the moss flora of Mexico. Altogether, he made four collecting trips to Mexico and collected a large number of mosses. The experiences he gained on these trips expanded greatly his knowledge of tropical mosses and their ecology. He also visited the major herbaria of this country and compiled a wealth of distribu- tional information. Jack Sharp tells with great delight that Howard came to the University of Tennessee to spend a week identifying Jack’s collection. He was there almost a month, staying at the Sharps, whose home is called the “Sharp Motel” by bryologists. Howard had not known that Jack’s unidentified material filled several large barrels. It was Howard who introduced the term “dung ball” to describe Jack’s habit of collecting several handfuls of mosses, squeezing them all together and stuffing them, often in wet condition, into a paper bag or wrap them, unpressed, in folded newspapers. Howard’s thesis, “The Appalachian-Ozarkian Element in the Moss Flora of Mexico with a Check-list of All Known Mexican Mosses,” was finished under Professor Harley H. Bartlett’s (“Uncle Harley” as he was fondly known) direction because of Steere’s departure for Stanford. Unfortunately, Howard’s thesis was never published. It is undoubtedly the most-cited thesis in the history of bryology. A great deal of its substance was eventually lifted and included in one of Howard's more significant contributions, “The Geographic Origins of the Mosses of North America’s Eastern Deciduous Forest,” published in 1972, as a part of a symposium in Tokyo, which was held the previous year. Howard’s eleven years in Ottawa were very pleasant ones. He and Irene both loved the city, they had a nice home, albeit a bit small for an expanding family. She was a Canadian and “at home.” While the National Museum was in antiquated quarters, there was generous financial support for technicians, equipment, and field work, and he lectured occasionally at Carleton University. He left an imprint on the National Museum that is well described in a letter from Wilf Schofield: “He built the bryological collection of the National Herbarium of Canada to world-class stature through active collecting and exchange. His organization of the collection and his great enrichment of the library are of profound significance to bryology.” Meanwhile, the bryology program at the University of Michigan became dor- mant after Steere’s departure. In 1964, however, Rudolph M. Schuster was hired to revive the program, but, after a tumultuous year in Ann Arbor, Rudy departed for the University of Massachusetts. Howard was then offered the job. He couldn't resist the lure of his alma mater and accepted the offer of Associate Professor of Botany, beginning in the fall of 1965. He and Irene purchased a beautiful home in a very nice section of Ann Arbor where they still reside. Their years in Ann Arbor have undoubtedly been their happiest and most cherished. There the children grew up, married, and fledged. He became Curator and Professor in 1969 and served as Chairman of the Department of Botany from 1981-83, interrupted by a serious illness. When he arrived in Ann Arbor, Howard quickly revived the graduate program 24 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 aco GY He si . Bhe Sackit } Se \ erench Began gto oites | creak ‘ oe nS First Sa FIG. 14. Howard makes the front page news in Mackinaw City, summer, 19835. in bryology and promptly set about revamping the quiescent bryophyte herbarium. Howard has supervised six Ph.D. students: Dale H. Vitt, 1970, now Professor of Botany at the University of Alberta, and currently President of the American Bryological and Lichenological Society and the International Association of Bryolo- gists; William R. Buck, 1979, now Curator of Bryophytes at the New York Botani- cal Garden; Jeffrey W. Holcombe, 1980, now teaching in a private school in New England; Allan J. Fife, 1982, now on the staff of the Botany Division, Department of Scientific and Industrial Research, Christchurch, New Zealand; Jonathan Shaw, 1983, now Associate Professor of Biology at Ithaca College; and Joseph R. Rohrer, 1985, now Associate Professor of Biology at the University of Wisconsin-Eau Claire. Marie Cole was his only Masters student. Howard’s relations with his students were what they all describe as “hands-off.” Following are some excerpts from a letter from Dale Vitt: “Howard always had time to talk, whether it was about bryology or just plain gossiping. From him I learned that there is only one way to be a good bryological taxonomist and that is to look at specimens. The more specimens one examines the better one will under- stand the species at hand. Surely no one else in the world has been able to name as many specimens from throughout the world as Howard. I rarely saw a specimen from really anywhere that he couldn’t determine to genus and often a good guess to species.” Dale goes on to say: “I think Howard has been a remarkably efficient editor over the years (often without getting the credit that he deserved, e.g., the Flowers book). His thesis is surely one of the better ones to have been written at its time: it brought Mexican bryology into the 20th century. Although I don’t agree with all his ideas, his ‘Origins of Eastern North American mosses’ paper is superbly written. in) nA 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM FIG. 15. Summer visitors at the Biological Station. Howard with Mason Hale in 1969 (above) and Dale Vitt in 1989 (below). The Great Lakes Flora is a perfect flora down to the wonderful stories that it contains (including my favorite about Mungo Park). Of course your joint work couldn’t be better!” All his students emphasize his professionalism, which, with amazing success, he was able to pass along to them. Their published dissertations reflect this. He went 26 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 G FIG. 16. Howard Crum with Bill Steere at the latter’s 80th birthday celebration at the New York Botanical Garden, 4 November 1987. through each thesis with a fine-toothed editorial comb. Individually, his relation- ship with his graduate students was mostly at the professional level, they tell me, but in a group he generally let his hair down and joined them in fun and clowning. Sometimes he surprised them with his “odd humor.” Bill Buck tells of introducing his parents to Howard, who, after a studied look at them paused and remarked, to Bill’s discomfort, “Well, you two look normal enough.” He was at his best at the Michigan Biological Station, where he still teaches every summer that health permits. Joe Rohrer describes his summer there as fol- lows: “My fondest memories of Howard come from the summer of 1980, which I spent at Douglas Lake. I was to begin my doctoral work with him (at your sugges- 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM a7. tion if you remember) that fall, and Howard strongly encouraged me to join him at the ‘bug camp’ for the summer so that I could get acquainted with him and the local bryophytes. He soon started calling me ‘Jo Jo the Dog-faced Boy,’ Howard has an odd sense of fun, but it also included poking fun at himself. For a class picture that summer, he allowed the students to adorn his shoulders with military-style maca- roni of Prtilium and Sphagnum. The field trips that summer were great. We were in the field five or six days a week. Howard was famous for never neglecting to stop at a doughnut shop in the morning and often for ice cream in the afternoon. One student commemorated the experience in a song he titled the ‘Bryophyte Blues.’ One verse went: Drive all day and we don’t stop Until we get to a donut shop, Buy a dozen, or maybe two And we hunt for mosses ‘til the day 1s through.” In the summer of 1967, Howard invited Wolfgang Maass and me to the Biologi- cal Station for a month. Howard was teaching both bryology and lichenology with assistance from Norton Miller and Dick Harris. Dale Vitt was also there working on his dissertation. It was a great summer. I spent most of the time with cytological studies on mosses (still unpublished) and Maass and Howard spent a great amount of time collecting and arguing about Sphagnum concepts. We all had fun. I had brought along a bottle of Howard’s favorite bourbon and suggested that we file it away under the letter “B” for some wet and cold day. “No way,” he said, and added that a year or so before a friend had brought him an expensive bottle of scotch and that they had filed it under “S” for that cold, rainy day. At an informal break in the bryology class Howard had told the students and others hanging around about some of Elizabeth G. Britton’s interesting traits. She made a habit of swiping small samples of enticing specimens, including types, that she had bor- rowed, but she would always write across the original specimen in her large, famil- iar handwriting, “Took Piece.” Soon, that cold rainy day arrived at the Station, and Howard went to the filing cabinet to retrieve the unopened bottle of scotch. To his amazement, it had been opened and was only three quarters full, but stuck to it was a slip of paper that, in a large handwritten scrawl, said “Took Piece.” The “honest” culprit was never discovered. The long tradition of bryology and lichenology at the Michigan Biological Station has been carried on most ably by Howard, and has to be listed as one of his major contributions to the field. He has made it a major training facility for bryology in this country. I encouraged all my students to go there and nearly all of them did so. Howard has invited colleagues and graduate students to use the Station as a base for their researches and in many instances he has obtained funds to support them. Howard, at 69, is still teaching a rigorous field course at the Biological Station. l.iary says “.... he is particularly proud of a nickname he acquired from his bryology students two years ago when they complained that they could not keep up with him in the field. These twenty year olds dubbed this 67 year old ‘Hurricane Howard’ and they called themselves the ‘brood bodies.’ We all have the class T- shirts to prove it!” Because the Crum children returned to the Station summer after summer, essentially through their entire childhood, it became a second home to them. They became a part of his professional and social life and were aquainted 28 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 FIG. 17. Bryophyte Reunion of the Summer Society honoring Howard Crum at the Biological Station, August, 1989. Above, Howard at his favorite thing, lecturing in a peat bog, Mackinac County, Michigan; and below, reminiscing with his first two Ph.D. students, Bill Buck (left) and Dale Vitt (right). with students and colleagues. Mary says that at lunch one day a student told her “I just love groveling at the base of trees with your Dad. We shouldn't be getting credit for having this much fun.” Howard was a member of Bill Anderson’s doctoral committee at Michigan. Bill 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM 29 FIG. 18. The Bryophyte Reunion continues, August, 1989. Bill Buck, Lewis Anderson, Howard Crum, Grace Blanchard Iverson and Norton Miller (left to right). wrote me the following. “One day I went to Howard and told him I wanted to change the subject of my dissertation. My first thesis was not working out well so I was switching to a problem that was sure to result in a publishable monograph in a reasonable time. Howard said, ‘That’s all very well, but what’s going to be your gimmick in this new thesis?” ‘My what?’ ‘Your gimmick. You know, the bit you'll use to convince the non-taxonomists on your committee that you did some science. Every taxonomist needs a gimmick, at least until he gets tenure.’ I have remem- bered Howard’s advice, made good use of it in my own career, and passed it along to my students. I have also borne in mind that while Howard cultivates the image of a vague and dithering sweetheart, he is actually a foxy strategist whose survival among the piranhas at the University of Michigan was not entirely a matter of good fortune.” Howard’s contribution to Sphagnology is enormous. Completely self-taught, he approached the genus from an ecological vantage point. He soon recognized that Andrews’ ultraconservative approach was an overreaction to Warnstort’s exces- sively narrow species concepts and did not mesh with what Crum saw in the field. On the other hand, he is troubled by more recent tendencies to edge back toward Warnstorfian attitudes, although some might call this edging forward. Howard has attempted to strike a middle ground in his concepts, although some can see his species concepts narrowing as his interests move into South America. Whatever, his North American treatise with its accurate descriptions and magnificent illustrations is the most useful work on Sphagnum that has been produced anywhere this cen- tury. He has set a standard that will be difficult to exceed. I agree with Wilf Schofield, Dale Vitt, and just about everyone who has a copy, that his Great Lakes moss flora is a gem. As Wilf wrote me, it is“. . . a delight to read and browse for intriguing observations derived from a rich background of reading and [experiences] with living plants.” All that is lacking is a comprehensive index to this huge cache of fascinating information. It is the only flora that I know of 30 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 that can be read with pleasure. It has and will continue to pique the interest of many a student, professional, and non-professional. A habitual note-taker, Howard carries a notebook with him wherever he goes. He and I have collected together over much of the southeastern United States. I drove, of course, and he took notes. I think he wrote down every thought I had on the trips. “I didn’t quite get that. Would you mind repeating it?” he would say. My copy of Grout’s moss flora is so marked up with Howard’s pencilings that I can scarcely use it. If he has a Bible, and Iam sure he has, | would not be surprised if it is filled with marginal notes. If I send him a specimen that is somewhat unusual and doesn’t quite fit a particular species, it comes back to me with a full penciled description which he jotted down while examining it. Once he sent me a description like this based on a Sphagnum that someone had sent him from Florida and that he thought was a new species. It proved to be ordinary S. molle. He had forgotten that there is a member of Section Acutifolia whose branch leaves have a resorption furrow. It is not often that he can be caught like this. With the help of Bill Buck, Howard’s published contributions to bryology are enumerated at the end of this article. Considering the seriousness and frequency of his illnesses they are nothing short of amazing. There are scarcely any breaks in the flow of publications through these years, indicating that he worked through and around his indispositions. He continues to work at the same pace. I can detect no let-up in quantity or quality of his output and there are still many more contribu- tions to come. | hesitate to attempt an assessment of Howard Crum’s contribution to Bryology because I have a strong prejudice in his favor. Wilf Schofield, who has been one of Howard's associates over the years, has this to say: “I consider Howard to be one of the major bryologists of the 20th Century. He is intensely engaged in a fascination for bryophytes. I can attest that, once contracted, this condition is usually incur- able. This commitment shows in his teaching, his lucid style of writing, and his sharing of his considerable knowledge through these potent means of communica- tion.” This says it well. Too, I think the scholarly qualities of his papers, reviews, and books, as well as those of his students, have set an example that has influenced bryology immensely, especially in North America. His professional experience, services and awards are summarized below. Instructor, Western Michigan College, 1946-47; Rackham State College Scholar, University of Michigan, 1947-48; Teaching Fellow, University of Michigan, 1948-49; Rackham Pre-Doctoral Fellow, 1949-S1; Research Biologist and Acting Assistant Professor, Stanford University, 1951-53; Assistant Professor, University of Louits- ville, 1953-54; Research Assistant, Duke University, Summer 1954; American Bryological and Lichenological Society (Editor, The Bryologist, 1954-62, Associate Editor, 1962—76; President, 1962-63); Curator of Cryptogams, National Museum of Canada, 1954—65; Visiting Lecturer, University of Michigan Biological Station, Sum- mer 1958; Lecturer, Carleton University, 1961-62; Curator, University Herbarium and Associate Professor of Botany, University of Michigan, 1965-69, Curator and Professor, 1969—present; Chairman, Department of Botany, University of Michigan, 1981-83; British Bryological Society (Associate Editor, Journal of Bryology, 1972— 77); Academic Counselor, College of Literature, Science and Arts, University of Michigan, 1973-76, 77—present; Nordic Bryological Society; Southern Appalachian Botanical Club; Sigma Xi; Michigan Botanical Club (Board of Directors, 1977-84; Associate Editor, The Michigan Botanist, 1975-76, 85—present, Editor, 1977-84; President, Huron Valley Chapter, 1974-75); Editorial Board, North American Flora, 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM 3] 1985—present; Ruth M. Sinclair Memorial Award for Freshman-Sophomore Coun- seling, 1978; H. A. Gleason Award for Excellence in Botanical Publication, 1981 (presented to H. Crum and L. E. Anderson); Abstracting for Excerpta Botanica (1959-83); Convener for Bryological Foray and Sessions, LX International Botanical Congress, Montreal, 1959. Crum! You bore me. I’m quitting. ACKNOWLEDGMENTS [am not sure that I can ber all the kind people that have helped me with this article. Because of the surprise element, I was not able to contact the best source of information, Howard Crum himself. | hope he will ne errors. Foremost, I am indebted to William R. Buck. He has not only been a constant nag, his help with the bibliography greatly improved its accuracy and completeness and he has been a durable source of information and has helped in countless other ways. Irene Crum has provided me with a wealth of information that only she could supply. She has added greatly to the spice of the account. Bil Anderson kindly provided me with a copy of Howard’s CV. The letters that Roger Crum, Mary Crum Scholtens, Bill Anderson, Joe Rohrer, Wilf Schofield, Jack Sharp, Jerry Snider, and Dale Vitt wrote me have been invaluable and their consent to be quoted is most appreciated. Pictures have been provided by Irene Crum, Mary Crum Scholtens, Bill Buck, Jerry Snider, and Dale Vitt. Finally, my wife, Pat, acted as chief censor, and Molly McMullen helped with the manuscript. I am grateful to both of them. — BIBLIOGRAPHY OF HOWARD A. CRUM Compiled by Lewis E. Anderson and William R. Buck 1949 The discovery of fruiting Gymnostomiella orcuttii in Mexico. Bryologist 52(4): 208-211. 1950 Publication date of the Prodromus Bryologiae Mexicanae. Bryologist 53(1): 60, 61. Additions to the moss flora of Panama. Bryologist 53(2): 139-152. (H. Crum and W. C. Steere) Mosses from Honduras. Bryologist 53(4): 292-295. Nota floristica de una associaci6n importante del suroeste de Tamaulipas, México. Bol. Soc. Bot. México 11: 1-4. (A. J. Sharp, E. Hernandez X., H. Crum and W. B. Fox) The Aponiecniane -Ozarkian element in the moss ee of Mexico with a nen list of all known Mexican mosses. Ph.D. Dissertation, University of Michigan, Ann Arbor. 504 pp. Mosses of Baja California. Rev. Bryol. Lichénol. 1934. 188-192. (L. : Koch and H. Crum) A checklist of the mosses of northeastern Mexico. Bol. Soc. Bot. México 12: 1-27. A be art ei area in Tamaulipas. Bull. Torrey Bot. Club 78(6): (ge 463. (E. Hernandez X., H. W. B. Fox and A. J. Sharp) 195] Notes on Mexican Jayloriae, with the description of a new species. Bryologist 54(4): 269-273. 1952 Bees notes on Palisot de Beauvois’ Prodrome. Bull. Torrey Bot. Club a ee 409. f ican mosses collected by Aaron J. ab oe 55(1): 5 ee aan: in Middle America. ante aor 13 A preliminary survey of the Nicaraguan moss flora. ie ee Pa 159-165. Mosses from Honduras. II. Bryologist 55(4): 279-285. _~ 1953 The propagula of Prerigynandrum filiforme. Bryologist 56(2): 98-100. Garckea phascoides in Panama. Bryologist 56(3): 204-207. Additional bryophytes from Panama. Rev. Bryol. Lichénol. 22(3/4): 148-159. (H. Crum and C. B. Arzeni Otros musgos de Tamaulipas. Bol. Soc. Bot. México 15: 13-16. (H. Crum and A. J. Sharp) CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 oS) Nw 1954 Additions to the moss flora of Kentucky. Trans. Kentucky Acad. Sci. 15(1/2): 24-26. Mosses of Mexico. I. Species new to the country. Rev. Bryol. Lichénol. 23(3/4): 256-264. 1955 Taxonomic studies in the Funariaceae. Bryologist 58(1): 1-15. CH. Crum and L. E. Anderson) Courses in bryology and lichenology to be offered during the summer of ee icc: S58(1): 78. [Review of] Franz Schommer: Kryptogamen-Praktikum. Bryologist 58(3): 26 Taylorta splachnoides and T. acuminata in America. Rev. Bryol. Lichénol. ae : 215- pt 1956 Syrrhopodon alatomarginatus. A new species from Trinidad. Trans. Brit. Bryol. Soc. 3(1): 7 Hedwig’s Species Muscorum and its effective date of publication. Taxon 5(6): 130-133. (L. FE Koch and “rum) Notes on Hypnodon, a genus of Orthotrichaceae. mae S59(1): 26-34. Two rare ae new to Canada. Bryologist 59(1): 3 [Review of] L. I. Savicz-Ljubitzkaja: Musci frondosi (1). "Shag. Bryologist 59(2): 148, 149. [Review of] at Bastin: Plants without flowers. Bryologist 59(2 Di: [Review of] G. M. Smith: car botany. Volume IT. Brienne and pteridophytes. Second edi- tion, Bryologist 59(2): | 150. e i Two new species of Barbula vee from the Southwest. cars Naturalist 1(1): 35-38. Lindbergia brachyptera in North America. Bryologist 59(3): 203-212. Puerto Rican Musci. New species and new ae Bry ae 59(4): 246-255. (H. Crum and W. Steere) eave of] E. Nyholm: /lustrated moss flora of Fennoscandia, Fascicle I. Bryologist 59(3): 226, 227. [Review of] Josef Podpéra: Conspectus Muscorum Europaeorum. Bryologist 59(3): 227, 228. Additional records of ffypnodon perpusillus. Bryologist 59(4): 255, 256. [Review of] A. Boros: Magyarorszag Mohat (Bryophyta Hungartae). Bryologist 59(4): 285. Taxonomic studies on ee th American mosses. I-V. Jour. Elisha Mitchell Sci. Soc. 72(2): 276-291. (H. d LE: rson) Mae to ai moss tee of Kentucky. Il. Trans. Kentucky Acad. Ser. 17(3/4): 131-134. Bryophytes from Guadalupe Island, Baja Califormia. Southw. Naturalist 1(3): 116-120. (H. Crum and H Miller) 957 A contribution to the moss flora of Ecuador. Svensk Bot. Tidskr. 51(1): 197-206. Some additions to the California moss flora. Madrono 14(2): 74-79. s Brazilian Expedition. Botany: Musci. Los Angeles Co. Mus. Contr. Sci. [8: 1- The mosses ay Porto Rico and ig Virgin Islands. N. Y. sae Sci. Scientific Surv. Porto ao and the Virgin Islands 7(4): 396-599. (H. Crum and W. C. Review of] Arthur Jewell: he »bserver’s book of mosses a liverworts. Bryologist 60(1): 4 Review of] Sigfrid Arnell: Hlustrated moss flora of ee ga I, Hepaticae. Bryologist sey 166. Review of} H. S. Conard: How to know the mosses and liverworts. Bryologist 60(2): 166, 167. Review of] Henry A. Imshaug: ons of West Indian lichens. Bryologist 60(4): 3 Review of] W. H. Welch: Mosses of Indiana: an illustrated manual. Bryologist 60(4): 367, 368. Review of] Z. Pavletic: Prodromus flore briofita Jugoslavije. Bryologist 60(4): 3 1958 Canadian bryological notes. 1. Bull. Natl Mus. Canada 147: 116-123. A contribution to the bryology of Haitth. Amer. Midl. Naturalist 60(1): 1-51. (H. Crum and W. € Steere re Asurvey of the moss flora of Jamaica. Bull. Inst. Jamaica, | Ser. 8: 1-90. (H. Crum and E. B. Bartram) Anacamptodon in North America. Bryologist 61(2 36-140. Taxonomic studies on North Armerican mosses. VI-XIILL pe Elisha Mitchell Sci. Soc. 74(1): 31-40. ( and L. E. Anderson) Some ane from Baja C Sitom Southw, Naturalist 3(1/6): 114-123. (CH. Crum and W. C. Steere) 1959 Cytotaxonomic studies on mosses of the Canadian Rocky Mountains. Bull. Natl Mus. Canada 160: 1— 89. (L. E. Anderson and H. Crum) es) SS) 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM The mosses of Gillam, Manitoba. Bull. Natl. Mus. Canada 160: 91-106. (H. Crum and W. B. Schofield) 66. yn) Bryoceuthospora nom. nov. Bryologist 62(1): . Crum and L. E. Ander [Review of] J. J. Barkman: Phytosociology and ecology ao ry ypige ure “pipes Bryologist 62(1): 74. [Review of] E. Nyholm: //lustrated moss flora of Fennoscandia. I. Musct. Fasc. 3. Bryologist 62(1): 75. A contribution to the Jamaican moss flora. eeu 62(3): 166—- 79, A small collection of West Javanese mosses. Bryologist 62(3): 188-190. [Review of] J. Szweykowski: Prodromus florae hepaticarum Poloniae. Bryologist 62(3): 195. [Review of] C. C. Cunningham: Forest flora of Canada. Bryologist 62(3): 196 1960 A bryological contribution from Florida. Bryologist 63(1): 32-46. (H. Crum and L. E. Anderson) Bryophytes of the Mont Tremblant region. Bry aa ne 59-63. (H. Crum and H. Williams) Three new species of Fissidens. eae 63(2): S [Review of] Z. Pilous & J. Duda: Klié k uréovani mechs CSR. Bryologist 63(4): 255. [Review of] B. Szafran: Mchy Me 1). Bryologist 63(4): 25 [Review of] H. Weymar: Buch der Moose. Bryologist Le PSS), [Review of] K. Bertsch: Moosflora von Stidwestdeutschland. Ed. 2 feo ag 63(4): 255, 256. [Review of] W. B. Turrill, editor: Vistas in botany. Bryologist 63(4): 2 1961 The vascular flora of Liard Hotsprings, B.C., with notes on some bryophytes. Bull. Natl. Mus. Canada 171: 131-197. (A. E. Porsild and H. Crum) Luisterella barbula in Georgia. Bryologist 64(4): 315-320. (H. Crum and L. E. Anderson) Mosses and liverworts of the Ottawa District—tentative list. December 1961. Department of Biology, Carleton University. 10 pp. mimeograph. [Review of] E. Nyholm: /llustrated moss flora of Fennoscandia. I. Musci. Fasc. 4. Bryologist 64(4): 384, 385. [Review of] S. Flowers: The Hepaticae of Utah. Bryologist 64(4): 385 [Review of] L. F. Koch: Bryology. Bryologist 64(4) [Review of] W. H. Welch: A monograph of the fC eee Bryologist 64(4): 385, 386. new Fissidens from Louisiana. neater st 64(4): 345-348. (H. Crum and L. E. Anderson) A bryological contribution from Florida. II. Bryologist Ae ): 368-370. (H. Crum and L. E. Anderson) [Review of] E. H. Walker: A bibliography of eastern Asiatic botany. Supplement I. Bryologist 64(4): 386. 1962 Mosses ae the Douglas Lake region of northern Michigan. A Preliminary Survey. vii + 87 pp. mimeo- h. University of Michigan. 1963 Encalypta brevicolla and E. longicolla in North America. Bull. Natl. Mus. Canada 186: 35-44. Mosses previously unrecorded for Panama. Bryologist 66(2): 66-68. (W. H. Welch and H. Crum) 1964 Mosses of the Douglas Lake region of Michigan. Michigan Bot. 3(1/2): 3-12, 48-63. G. Taylor: British ferns and mosses. eieeae 67(2): ia 242. Review of] E. F. Warburg: Census catalogue of British mosses. Ed. - Erolone 67(2): 2 Review of] B. R. Vashista: eve (for degree re Part LT. Bryophyta. Bavalocict 67(3): 382. Review of] [D. K. Zerov:] [Flora of Hepaticae and ties ee ae Ukraine. ! rae 67(3): 382. Review of] P. A. Florschiitz: The mosses of Suriname. Part I logist 67(3): 382, 383. Review of] Pang-Chieh Chen (chief compiler): Genera Mu iscorum ee Brot 67(3): 383. New names for some North American mosses. Bryologist 67(2): 162-164. (H. Crum, W. C. Steere and L.E.A on) Notes on ieee collenchymatum. Bryologist 67(3): 350-355. (H. Crum and L. E. Anderson) 1965 James Kucyniak (+ March 11, 1962). Rev. Bryol. Lichénol. ae 614-617, pl. XXVI. Additions to the Doves! Lake bryoflora. Michigan Bot. 4(1): 2 Hyophilopsis, a new genus of Pottiaceae. pce 68(1): 68- ai Mnium nudum in Japan. Bryologist 68(1): 1 Fissidens eee new to the United ce eee 68(1): 100-105. (H. Crum and L. E. Anderson) 34 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 The taxonomy and distribution of Acaulon schimperianum, Bryologist 68(2): 208-211. CH. Crum and L. A re- eralaaeen of Glossadelphus andersonu. Bryologist 68(2): 219, 220 [Review of] R. S. Breen: Mosses of Florida. An ate er lia J. eiiean Bot. Lab. 28: 146. Brotherella tenuirostris in Canada. ae eine 68(2): 220, Grimmia oe v. indianensis review es ig mie )f 2332235. ecords from Haiti. Bryologist 68(2) ; eratcnae on the Jamaican bryoflora. Trans. eis ae Soc. 4(5): 790-793. Barbula johansenti, an arctic disjunct in the Canadian Rocky Mountains. Bryologist 68(3): 344, 345. [Review od F. H. Perring, P. D. Sell and S. M. Walters: A flora of Cambridgeshire. Bryologist 68(2): 271, [Review of] W. W. Judd & J. M. Speirs (eds.): A naturalist’s guide to Ontario. Bryologist 68(2): 274, 275. Barbula eustegia, a moss new to Canada. Canad. Field-Naturalist 79(2): Dicranella crispa, a moss new to eastern Canada. Canad. Field-Naturalist 79(3): 2 A list of mosses of North America. Bryologist 68(4): 377-432. (H. Crum, W. : Steere and L. E. — WG ~—l Some additional new names for North American mosses. Bryologist 68(4): 432-434. (H. Crum, W. C. St gt a sie nderson) Hyophilopsis—a nomenclatural correction. Bryologist 68(4): 470. [Review of] A. E. Jensen: The how and why wonder book of mushrooms, ferns, and mosses. Bryologist 68(4): 485. | A tribute to Edwin B. Bartram, Bryologist 69(1): 124-134. Evolutionary and phytogeographic patterns in the ene moss flora, pp. 28-42 in R. L. Taylor and R. A. Ludwig (eds.), The evolution of Canada’s flora. Toronto. The relationships of Tetraplodon aul 'ARICUS. a bei 205-207. A note on the taxonomy and distribution of Polym >, Advancing Frontiers Pl. Sei. 17: 23-26. Bryophytes of Labrador and Ungava. Bull Natl. Mus. Canada 216: 87-101, (H. Crum and P. Kallio) Bryophytes of Owen Sound and the Bruce Peninsula, Ontario. Bull. Natl. Mus. Canada 216: 102-122. A taxonomic account of the genus Thelia. Bull. Natl, Mus. Canada 216: 123-127. Bryophytes new to the Douglas Lake area. Michigan Bot. 5(2): 57-61. (N. G. Miller and H. Crum) Edwin B. oe (1878-1964). Rev. Bryol. Lichénol. 34(3/4): 931-936. 1967 The status of Hylocomium splendens var. tenue. Bryologist 70(1): 78-101. (GH. Crum and L. E. Anderson) Studies in North American Bryaceae I-I. Bryologist 70(1): 106-110. Three propaguliferous Pohlias from Michigan. Bryologist 70(1): 118, 119. (GH. Crum and N. G. Miller) Barbula ee a new species from Peru. Bryologist 70(2): 235-237. A new spe of Prerigoneurum trom California. Madrono 19(3): 92-94. A variety - pet (Musci) new to North America. Canad. Field-Naturalist 81(2): 113-115. Bryophytes from the Upper Falls of the Tahquamenon. Michigan Bot. 6(2): 54-57. (H. Crum and N. G. Miller) Fifty-six mosses new to Nicaragua. Bryologist 70(3): 365-369. (M. R. Crosby and H. Crum) — 968 Funarta flavicans, a moss new to Canada. Michigan Bot. 7(1): 24. (H. Crum and W. Botham) ryvophytes new to Michigan. Michigan Bot. 7(3): 132-134. (H. Crum and N. G. Miller) x new species of Drummondia from Mexico. plea 71(2): 117-119. (D. H. Vitt and H. Crum) Two new mosses from Hawai. J. Hattori Bot. Lab. 31: 293-296. (H. Crum and D. cei Dombots) Mosses from Honduras. HII. Na acne Frontiers PL - 21: 189-193. ae Bryophytes new to Michigan. [H]. Michigan Bot. 8(1): 2 9. (H. Crum and N. G. er) A reconsideration of the relationship of Barbula Scene (Musci). Canad. Field- ae 83(2): 156, L37 s Nomenclatural notes on North American mosses. Bryologist 72(2): 240-246. Recent sie saute collections in Barbados. J. Barbados Mus. & Hist. Soc. 33: 86, 87. (W. H. Welch and H. Crum) 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM 3 1970 Re ae in Charlevoix County, Michigan. Michigan Bot. 9(2): au 116. (W. Zales and H. Crum) Mosses new to the flora of Michigan. See Bot. 9(2): 139-14 et, tomentosa new to the United States. Bavolosict 73(1): eer (D. H. Vitt and H. Crum) A new species of Cryphaea from Florida. ere 73(2): 380, 381. Recent cryptogamic collections in the West Indies. Rev. Bryol. Lichénol. 37(2): 223-235. (W. H. Welch and H. Crum) 1971 Brachymenium andersonii, a new species. Bryologist ae 47-49. Nomenclatural changes in the Musci. Bryologist 74(2): 165-174. “Leucodon sciuroides” in North America. Bryologist te ): 373, 374. (H. Crum and L. E. Anderson) Three new moss species from the Hawaiian Islands. Bryologist 74(4): 484-489. (W. J. Hoe and H. Crum) 1972 The geographic origins of the mosses of North America’s eastern deciduous forest. J. Hattori Bot. Lab. 35: 269-298. Validation of Leucodon brachypus var. andrewsianus. Bryologist 75(1): 101. (H. Crum and L. E. Anderson) Mosses new to the state of Michigan. Michigan Bot. 11(3): 123, 124. Mosses of unusual interest from Baja California. Madrono 21(6): 403, 404. A taxonomic account of the Erpodiaceae. Nova Hedwigia 23(2+3): 201-22 seed magellanicum new to the West Indies. Bieler 75(3): 359, 360. (H. Crum and F. J. mann) A new at ee from Virginia. Bryologist 75(3): 360-362. The dubius origin of Glyphomitrium canadense Mitt. J. Bryol. 7(2): 165-168. Splachnum ampullaceum in West Virginia. Castanea 37(4): 253-257. (H. Crum, E. G. Fisher and H. C. Burtt [Review of] E. Lawton: Moss Flora of the Pacific Northwest. eee 75(2): 205-207. Disjunctions in bryophytes. Ann. Missouri Bot. Gard. 59(2): 1 204. 1973 Leucobryum albidum new to Michigan. Michigan Bot. 12(2) A new list of mosses of North America north of Mexico. Bryologist 76(1): 85-130. (H. Crum, W. C. Steere and L. E ; Mosses of the Great Lakes Forest. Contr. Univ. Michigan Herb. 10: 1-404. [Review of] J. Taylor and A. P. Underhill: Liverworts, a photographic study. Bryologist 76(4): 590, SOL. 1974 eens muehlenbeckii, a moss newly discovered in Michigan. Michigan Bot. 13(2): 64. v Grimmia from Michigan's Upper Peninsula. Michigan Bot. 13(4): 171-174 nue ‘| O. Degener, I. Degener and H. Hérmann: Mosses of Hawaii. Bryologist 77(1): 103. Some Cee of Silumiut Island, Northwest Territories. Canad. Field-Naturalist 88(3): 361. 362. (N. McCarthy and H. Crum A new ee from high altitude Costa Rica. Ann. Missouri Bot. Gard. 61(3): 904-906. (H. Crum nd M. R. Crosby) — — 1975 The vertical Sane iNe of eee rae in pane hollow complexes in northern Michigan. Michi- gan Bot. 14(4): 190-200. (D. H. Vit . Crum and J. A. Snider Notes on the distribution of Sphagnum hee Contr. Univ. Michigan Herb. 11(2): 85-87. Comments on Sphagnum capillaceum. Contr. Univ. Michigan Herb. 11(2): 89-93. Seligeria tristichoides, a moss new to the West. Canad. Field-Naturalist 89(3 ): 317, 318. 1976 Mosses of the Great Lakes Forest. Rev. ed. University of Michigan ges Ann Arbor. 404 pp. Distribution of Grimmia cribrosa in North America. Bryologist 79(3): 355-357. (R. R. Ireland and H. Crum) Revision of the genus Rozea (Musci). Bryologist 79(4): 406-421. (W. R. Buck and H. Crum) 36 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 Sphagnum taxa and their distribution in Towa. Proc. lowa Acad. Sei. 83(3): 98-101. CH. Crum, N. R. Lersten and G. H. Crum) 1977 William Campbell Steere. pein 80(1): 1 Grimmia pulvinata in eastern North America. Bryologist 80(1): 152, 153. CH. Crum and Y. Mescall) [Review of] A. Boros and i Jarai-Kornlodi: An atlas of recent European moss spores. Rey. Palacobot. Palynol. 23(5): 404-406. Sphagnum la an in the American West. Bryologist 80(1): 156-158. (H. Crum and J. A. Snider) Meiothecium, a new record for North America. Bryologist 80(1): 188-193. New eine sat new taxa of mosses proposed by Nils Conrad Kindberg. Mem. New York Bot. Gard. 28(2): 1-220. (W. C. Steere and H. Crum) Proposal for the conservation of Pleurozium Mitt. Taxon 26(5/6): 596, 597. (M. R. Crosby and H. Crum) Desmatodon steereanus, a new species of Pottiaceae from Peru. Bryologist 80(4): 638-640. (R. H. = Zander and H. Crum William Campbell Steere. An account of his life and work. Bryologist 80(4): 662-694. 1978 Sphagnum richardsianum, a new species from Mexico. Contr. Univ. ree Herb. 11(5): 281-283. E-ntodon schleicheri new to North America. ema 81(3): 429-432. (W. R. Buck and H. Crum) A re-interpretation of the Fabroniaceae with notes on selected genera. J. a Bot. Lab. 44: 347-369, Buck and H. Crum) Alexander William Evans. /n: C. C. Gillispie (ed. in chief), Dictionary of Scientific Biography 15(Suppl. 1); 148-151. Charles Scribner’s Sons, New York 1979 n expanded description of Sphagnum splendens. Michigan Bot. [8(2): 69-72. a hardsia, anew genus of Amblystegiaceae (Musci). Fieldiana, Bot., n.s. 1: 1-18. (H. Crum and L. Ee. derson Zortula ammonsiana, anew species from West Virginia. Bryologist 82(3): 469-472. (H. Crum and L. E. Anderson) 1980 A guide to the identification of Mexican Sphagna. Contr. Univ. Michigan Herb. 14: 24-52. Comments on Sphagnum henryense. Contr. Univ. Michigan Herb. 14: 53-56. a . North America. [Exsiccati and ne ae | Fase. I-1V. Nos. 1-200. University of Michigan and e University. (H. Crum and L. E. Anderson) 1981 Mosses of eastern North America. Vols. | and 2. 1328 pp. Columbia University Press, New York. (H. Crum and L. E. Anderson) Mosses of North America. eee and oes |] Fase. V-X. Nos. 201-500. University of Michigan and Duke University. (H. ¢ rand L. E. Ande ete An inventory of John Macoun’s aa Te Occ. Pap. Farlow Herb. 36. Taxonomic and nomenclatural notes on Middle American mosses. Rina 84(3): 390-393. Philonotis corticata, new from Mexico. ede 84(3): 398-401. CH. Crum oe Hp ee Ill) Eccremidium, a genus of Ditrichaceae new to the Americas. Bryologist 84(4): 5 Joyce Hedrick Jones (U.S.A.): eer nee Lichenol. Newsl. 14(2): 15, ra 982 Bryophyta. a: S. P. Parker, Svnopsis and classification of living organisms 1; 272. McGraw-Hill, New York. Mosses Musci. Pages 525-527; Liverworts and hornworts Hepaticae and Anthocerotae. Page 527; Lichens Lichenes. ct S32, 533. In: S. J. Wernert (ed.), Reader's Digest American wildlife. cece New Comments on the Mielic ee ferioideae of Central pecs with the description of a new species of Sy see eran Contr. Univ. Michigan Herb. 15: 209-217. (A. J. Shaw and H. Crum 1992 ANDERSON: A TRIBUTE TO HOWARD CRUM 37 Mosses of North America. [Exsiccati and schedae.] Fasc. XI-XIV. Nos. 501-700. University of Michigan d Duke University. (H. Crum and L. E. Anderson) 1983 Grimmia hermannii, new to Canada. Michigan Bot. 22(1): 10 Mosses of the Great Lakes Forest. Ed. 3. University of Michigan Herbarium, Ann Arbor. 417 pp. Diagnoses of the new species [in part]. /n- K. Magdefrau, The eee vegetation of the forests and paramos of Venezuela and Colombia. Nova Hedwigia 38: 5 [Review of] H. Godwin: The archives of peat bogs. Bryologist an a6 Mosses of North America. [Exsiccati and schedae.] Fasc. XV-XX. Nos. a ee University of Michi- gan and Duke University. (H. Crum and L. E. Anderson) 1984 Vegetative reproduction in Platydictya. Michigan Bot. 23(1): 21, 22. Crassicosta, a new genus of Amblystegiaceae from northern Mexico. Bryologist 87(2): 149, 150. (H. Crum and A. J. Sharp) Fossombronia in Michigan. Michigan Bot. 23(4): 157-163. oe G. Miller and H. Crum) Notes on tropical American mosses. Bryologist 87(3): 203 New ee interesting mosses from Costa Rica. J. Bryol. BO : 193- 200. (H. Crum and P. W. Richards) wo new species of Sphagnum from Costa Rica. Cryptogamie, Bee ee 5(3): 293-297. Acaulon casasianum, a new species from Spain. Lindbergia 10(1): 1-3. (M. Brugués and H. Crum) Sphagnopsida. Sphagnaceae. No. Amer. FI. ser. I, 11: 1-180. Peristome homo a ogy in Mielichhoferia ae a taxonomic account of North American species. J. Hattori Bot. Lab. 57: 363-381. (J. Shaw and H. m) Re Se ae a new species of the eee es from Quebec. Bryologist 87(4): 371-373. R. R. Ireland and H. Crum) [Review W. D. = ese: Mosses of the Gulf South. From the Rio Grande to the Apalachicola. Bryologist 87(3): 287, 288 1985 Lewis Anderson, a tribute from a friend and Siete Monogr. Syst. Bot. isso Bot. Gard. 11: 1-27. Two undescribed species of Hygrohypnum from Mexico. sae 88(1): 2 A new genus of Hookeriaceae from Mexico. Prolog eae 2B 24: Notes on Pylaisiadelpha in Mexico. Bryologist 88(1): 2 New Sphagna from Brazil. Cryptogamie, Bryol. Lichen, Co: 181-184. Traditional make-do taxonomy. Bryologist 88(3): 221, Pseudoleskeella papillosa, a rare moss in Quebec. pened 88(4): 357, 358. l A survey of the moss genus Sclerodontium. Hikobia 9(4): 289-295. Fossombronia and the mystery of spore dispersals. nee Bot. 25(3): 120-122. Sea sa In: G. S. Mogensen (ed.), Illustrated moss flora of arctic North America and Greenland. el. Gronland, Biosci. 18(2): 3-61. Aes and nomenclatural addenda to the Mexican moss flora. Bryologist 89(1): 23-27. 1987 A new section and species of eae from Ecuador. Contr. Univ. Michigan Herb. 16: 141-143. Some Latin American mosses new to science. Contr. Univ. Michigan Herb. 16: 135-140. Bestia, Ee aoe and en an pias of ees ee 90(1): 40-42. of Sphagnum from South America. J. Hattori Bot. Lab. 63: 77-% Leratia ind its place in the oui crt Sane, ve New York Bot. Gard. 45: )2—608. 1988 A focus on peatlands and peat mosses. University of Michigan Press, Ann Arbor. xi + 306 pp. (H. Crum in collaboration wit anisek) A new species of Aulac one from Brazil. Bryologist 91(3): 191, 192. 38 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 [Review of] R. R. Ireland and G. Bellolio-Trucco: Mlustrated guide to some hornworts, liverworts and mosses of eastern Canada. Bryologist 91(3): 250. [Review of] E. Nyho as Illustrated flora of Nordic mosses. Fasc. I. Fissidentaceae-Seligeriaceae. Bryologist 910): 2 A contribution to the a (Sphagnaceae) flora of Paraguay. Brittonia 40(2): 188-194. (H. Crum and W. R. Buc 1989 Noteworthy collections. Michigan. Sphagnum obtusum Warnst. (Sphagnaceae). Michigan Bot. 28(1): 38 Notes on South American species of Sphagnum. J. Bryol. 15: 531-536. New tropical American species of Sphagnum. Bryologist ae 98-104. William Campbell Steere (1907-1989). Michigan Bot. 28(2 Long live the lichens. Michigan Nat. Resources Ma New names for some North American mosses. Bivoloeist ma 533. (H. Crum and L. E. Anderson) 1990 An evaluation of familial ie among the genera traditionally aligned with the Thuidiaceae and Leskeaceae. Contr. Univ. Michigan Herb. 17: 55-69, (W. R. Buck and H. Cru Comments on Sphagnum sect. Sphagnum in South America. Contr. Univ. wichiean Herb. 17: 71-81. A new look at ie sect. Acutifolia in South America. Contr. Univ. Michigan Herb. 17: 83-91. Preliminary notes on Sphagnum sect. Subsecunda in South eens Contr. Univ. Michigan Herb. 17: 93-97. Noteworthy collections. Michigan. si ea acai anes (Funariaceae). Michigan Bot. 29(1): 27. Sphagnum Seinen sie Crum sp. Jn: W. R. Buck, Contributions to the moss flora of Guyana. Mem. Nev k Bot. aay 64: 185, My sieges inretortum, a New species in a new section from Bolivia. Bryologist 93(3): 283-285, > mosses of North America north of Mexico. Bryologist 93(4): 448-499. (L. E. Anderson, H. ae ah W. R. Buck) wn pee ay 1991 Two new species of Sphagnum from Brazil. Bryologist 94(3): 301-303. Liverworts and hornworts of southern Michigan. University of Michigan Herbarium, Ann Arbor. vii + A partial clarification of the Lembophyllaceae. J. Hattori Bot. Lab. 69: 313-322. Riccta frost, a liverwort new to Michigan. Michigan Bot. 30(1): 15-21. (M. R. Penskar, N. G. Miller and H. Crum Contr. Univ. Mich. Herb. 18:39—41. 1992. CRUMUSCUS VITALIS GEN. ET SP. NOV. (DITRICHACEAE) William R. Buck New York Botanical Garden Bronx, NY 10458-5126, U.S.A. Jerry A. Snider Department of eros ee University of as Cincinnati, OH 45221-( 006, a S.A. In 1982, Daniel M. Vital of the Instituto de Botanica in Sao Paulo, Brazil, sent around duplicates of a ditrichaceous moss he had collected on Itatiaia five years earlier. After receiving no responses, he sent it to W.R.B. several years later, who also let it languish for a few years before facing up to it. After some discussion on its affinities to Pleuridium, we have decided to describe it as a new genus. Both of us have had close and amiable ties to Howard Crum for many years and are delighted to have this opportunity to describe a distinctive new genus for a good friend and colleague. Crumuscus vitalis Buck & evteleit eo et sp. nov. Fics. 1-10. A Pleuridio folus perict bus a caeteris valde absimilibus, cellulis foli distaliter bistratosis Ae ioc eee necnon calyptris mitriformibus differt. Plants perennial, very small, to ca. 8 mm tall but often leafy only in upper 2-3 mm with old, denuded stems buried in soil. Stems reddish, especially with age, branching from beneath perichaetia and thus obscuring them, ca. 100 wm in diame- ter, in cross-section with ca. 7 layers of small, firm-walled cells surrounding a well developed central strand of very small, thin- and fragile-walled, collenchymatous cells; rhizoids smooth; axillary hairs 28-45 wm long, 2-celled, with a single, short to elongate, brown basal cell and a single, elongate, hyaline apical cell. Leaves when dry erect, often + falcate, obscurely homomallous, when moist remaining erect but neither falcate nor homomallous, becoming larger toward stem apices, oblong- lanceolate, sometimes broadly so, ca. 0.6-0.9 mm long, broadly long-acuminate from an expanded base; margins subentire to crenulate, plane; costa broad and strong, ca. 40-55 wm wide at base, almost filling the acumen, excurrent, roughened above at back from projecting cell ends, in cross-section with 4 guide cells and two bands of stereids; laminal cells bistratose above (except at extreme margins), at shoulders unistratose with bistratose streaks, unistratose below, mostly rectangular, (1-)2-5:1, (9-)11-22 um long, firm-walled, mammillose-prorulose at back above, smooth below, scarcely differentiated at insertion. Paroicous? Perichaetia terminal, leaves strongly and rather abruptly differentiated, erect, lanceolate to broadly lanceolate, to 2.5 mm long, concave, gradually acuminate; margins entire, plane to irregularly narrowly recurved; costa subpercurrent to percurrent, not filling the acumen, roughened at back; cells unistratose except perhaps in extreme acumen, quadrate to short-rectangular above, long-rectangular below, smooth. Setae very short, ca. 0.1 mm long, brittle, smooth, from a naked vaginula ca. 0.45 mm long; capsules immersed, cleistocarpous, eperistomate, spherical to ovoid, stoutly apicu- 39 40) CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 | | Rp Jt) 3 Ww a} O, 7 meen 9 e Wepea ne wtae USOHS CES ¢ inden Y iu y ) \ GSI Ser Bek SSG y, ain) IS Vay ( ) Ons )\) Sie? sien Crumuscus Mus \) vitalis 8 | op) fe WV { \) a QD id ie es5 r a FIGS. 1-10. Crimuscus vitalis. 1. Habit, x8. 2. Vegetative leaves, x50. Laminal cross-sections (from top to bottom) near leaf apex, at leaf shoulder, near leaf base, x 200. 4. Leaf apex, x 200. 5. Upper laminal cells at margin, x 200. 6. Lower laminal cells at margin, 200. 7. Perichaetial leaf, x50. 8, Lower marginal perichaetial leaf cells, x 200. 9. Calyptra, x50. 10. Detached capsule, «50. Figs. 1-6, 9, 0 from the type (NY); figs. 7, 8 from Schdfer-Verwimp & Verwimp 14639 (NY), —_— late, 0.75—0.85 mm long including the 0.15—0.2 mm long apiculus, separating from the seta at base of the capsule; exothecial cells irregularly rectangular, firm-walled, becoming gradually shorter and quadrate toward the apiculus; stomata none. Spores + spherical, tardily separating from tetrads, 17-20 sm in diameter, papillose to verruculose. Calyptrae mitrate, minute, covering scarcely more than the capsular apiculus, 0.25—0.3 mm long, naked, smooth or slightly roughened. Type. Brazit. Rio de Janeiro: Parque Nacional de Itatiaia, ca. 3 km SW of Pico das Agulhas Negras, 23°23'S, 44°38’W, on rocky cliff covered with thin layer of soil, 24 July 1977, D. M. Vital 7435 (NY!, holotype; BUF, CINC!, FLAS, MO, SP!, isotypes). ADDITIONAL SPECIMEN SEEN. BRazit. Minas Gerais: Nationalpark Itatiaia, auf feuchtem, offenen Humus nahe Brejo da Lapa, 2120 m, 7 July 1991, Schdfer-Verwimp & Verwimp 14639 (NY). This new genus of Ditrichaceae is named in honor of Howard Crum on the occasion of his 70th birthday; we are pleased to have come up with a name that is 1992 BUCK & SNIDER: CRUMUSCUS 4] not crummy-sounding! The specific epithet honors the collector, Daniel Moreira Vital, now a vital 67. Crumuscus 1s closely related to Pleurtdium as evidenced by capsule morphology and leaf areolation. It differs from Pleuridium on the basis of the following: the lack of stomata in the capsule wall, the mitrate calyptrae, the bistratose and mammillose- prorulose upper lamina of the vegetative leaves, and the leaf costa showing four guide cells and two stereid bands in transverse section. Crumuscus appears to approach Pleuridium through P. lindigianum (Hampe) Churchill. The latter usually lacks stomata in the capsule wall and has mitrate calyptrae. However, the leafy gametophyte is typically that of Pleuridium in having unistratose, smooth leaf cells and undifferentiated, sub-stereid cells in the costal cross-section. Differentiated guide cells and stereid bands are absent. Pleuridium papillosum Magill, a species endemic to South Africa, is the only known Pleuridium to exhibit papillose leaf cells. In the latter, however, the unistratose laminal cells have several low, blunt papillae over the lumina. Additionally, the capsules contain stomata, and the calyptrae are cucullate. Crumuscus appears to be even more closely aligned to the geographically sympatric Cladastomum, a genus both poorly known and collected. This genus, of two species (and a dubious variety), is confined to the Agulhas Negras region of Itatiaia National Park and Pico da Bandeira of Capara6é National Park, both part of the Serra da Mantequeira in eastern Brazil. The two genera share the following features: mitrate calyptrae, capsules lacking stomata, stems with a well developed central strand, and leaf costae in cross-section showing guide cells surrounded above and below by stereid bands. However, Cladastomum differs from Crumuscus in its overall aspect of julaceous sterile shoots, its ovate-cuspidate leaves appressed to the stems, as well as its production of faintly bipolar spores. ACKNOWLEDGMENTS We are grateful to the collectors of the material, Daniel Vital and Alfons Schafer-Verwimp (really his wife Inge), for making it available to us, and for accompanying the senior author to Parque Nacional Itatiaia, where, in addition to the Schafer-Verwimp collection of Crumuscus, Cladastomum was also collected. Rupert Barneby graciously corrected the Latin diagnosis, and Bobbi Angell prepared the elegant illustration. a aa > jai SS pt pay al) he nce ee -_ - oo so a ba _ 7 Oo y= > _ : ; : _ - 7 : : a eee Contr. Univ. Mich. Herb. 18:43-71. 1992. KEY TO THE MOSS GENERA OF NORTH AMERICA NORTH OF MEXICO Dale H. Vitt Department of Botany University of Alberta Edmonton, Alberta T6G 2E9, Canada William R. Buck New York Botanical Garden Bronx, NY 10458-5126, U.S.A. Traditionally, mosses have been considered by many to present severe difficul- ties in identification of both species and genera. However, often if the genus is known, species identification follows with much less difficulty. Moreover, most regional moss floras present keys to species beginning at either the family or genus rank. We present here a key to the genera of mosses found in North America, north of Mexico. The key is strictly dichotomous and attempts were made to use gametophytic features whenever possible. Mosses (Class Bryopsida or Musci) are the largest group of plants in the Bryophyta, and are the third largest class of green land plants in North America following the Monocots and Dicots. The recently published lists of North American mosses (Anderson er al. 1990 [Andreaeidae and Bryidae]; Anderson 1990 [Sphagni- dae]) tabulate 1320 species in 312 genera. To these, we add the genus Takakia with two species (Murray 1988; Smith et a/. 1990). Within the 313 genera of mosses found in North America, 15 have 15 or more species. Only two genera have more than 50 species—Sphagnum with 72 and Bryum with 66; five genera have 30 or species (Orthotrichum—37, Fissidens—36, Grimmia and Brachythecium each with 34, and Pohlia—30). Other large genera are Tortula (29), Dicranum (27), Racomitrium (24), Hypnum (21), Campylopus (18), Hygrohypnum (18), Encalypta (16), and Poly- trichum (15). These 15 genera contain 476 or 36% of the North American species. Historically, keys to mosses have used growth form (acrocarpy vs. pleurocarpy) as a major dichotomy. We have resisted the use of this character whenever possible, and instead placed more emphasis on microscopic features of the leaves; namely the presence, absence and number of costae found on leaves and on the ornamenta- tion of leaf cells (papillose vs. smooth). Generic concepts depend largely on previous monographic work completed in particular families. Some North American families have had no recent generic revision, while others have newly revised generic concepts in place. We have fol- lowed the generic concepts presented in Anderson ef al. (1990), including the recognition of segregate genera in the Mniaceae, Amblystegiaceae, and Grim- miaceae. These generic concepts differ somewhat from those accepted in the Cana- dian (Ireland et a/. 1987) and European checklists (Corley et a/. 1981). Reprints of this article may be purchased from the American Bryological and Lichenological Society. Send orders to Jerry A. Snider, Dept. of Biological Sciences, University of Cincinnati, Cincinnati, OH 45221-0006, U.S.A. Ad CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 Floristic treatments of North American mosses are available for many areas of the continent, and date back to the I1880’s when Lesquereux and James (1884) published the first treatment of mosses of the continent. Still a valuable source of information and the only flora that treats the southwestern United States is Grout’s Moss Flora of North America (Grout 1931-40). In more recent times (since 1970) several excellent regional floras have been written and form the basis of our understanding of North American mosses. Included are the following manu- als: Eastern North America—Crum and Anderson (1981); the United States Pa- cific Northwest—Lawton (1971); Utah—Flowers (1973); northern Michigan— Crum (1973); Maritime Provinces of Canada—Ireland (1982); the southeastern Gulf Shore—Reese (1984); the Interior Highlands—Redfearn (1983); and north- western North America—-Vitt ef a/. (1988). Definition of terms used in this key follow those given in Crum and Anderson (1981) and in Magill (1990) We have the great privilege to dedicate this key to Howard Crum, our teacher and good friend. His patience and perseverence in performing the unenviable task of teaching both of us the fundamentals of science and bryology are remarkable. Without him, neither of us would have been able to complete the present work. To Howard Crum we give our heartfelt thanks. — . Gametophytes seemingly absent (consisting only of protonemata); sporo- phytes of asymmetric capsules and papillose setae ............. Buxbaumia = 1. Gametophytes present (with obvious leaves); sporophytes various... . 2 2. Leaf cells arranged in a network of narrow, green cells alternating mil ae hyaline cells branches usually in-clusters | Gea %s a8.au4s Sphagnum . Leaf cells of one kind (green), or if of two kinds, branches never in GUISES: Suaw vices coast ie eeu k ta tae ase ee tees 3 3. Leaves attached all around the stem (foliate stems sometimes flattened (SONAL): = nh gavin oe ata saa. ea eee eee ad eon eee eee ee 4. Leaves appearing split at the base, consisting of two vaginant laminae which clasp the stem and base of the leaf above (equitant) .... Fissidens 4. Leaves with expanded bases, not clasping leaf above ............ 5 5; Saves Ccosiaie: DrOIOneIata \UMINOUS® wider Sea ere wes Schistostega 5, Leaves umicostate: protonemata not UMINOUs: 2c: fs 9.44 jas so cect dee 6 6. All leaves with a rough (papillose), linear subula ........... Distichium 6. Vegetative leaves with a smooth mucro; perichaetial leaves with a SIO OOUT SIOUIG.« «ig'ch Aa buds. 1-44.44 etd ba aneaewe Re ORR ae a aS Bryoxiphium 7. Leaves with lamellae or filaments on the adaxial (upper) surface of the COsSta cece PLOpaIUla).” vx axacak Peawned sd nde ees uew ete eeees 8 7. Leaves without lamellae or filaments on the adaxial surface of the costa (Di prop arin SOmenINEeS DEESEDL) © uss veua doce ee nen oe es ste ees 23 8. Leaves with filaments on the adaxial surface of the costa ......... ") 8. Leaves with lamellae on the adaxial surface of the costa ......... 10 9. Leaf margins broadly inrolled (and mostly obscuring the filaments) .... Aloina Pe Menta ie Celene 10 TOVOUNES. <5 tu nicdieneSaae ewes keeer ows Crossidium 10,-Leaves bordered with elomeate cells to thick-wallederoundedtolinear 222. f..22s aan 63 62. Lateral and dorsal leaves differentiated in areolation; stems pinnately DRANG OAR. eka t os ths eee ene tre aes ss eG ated Vesicularia 62. Lateral and dorsal leaves with similar areolation; stems irregularly branched? sa. 253.5 wR eee ee ee goer ea tre a phen ee Hookeria OS: ecavessslendenly lome-decuimem mere ohare ee ee Plagiothecium 63. Leaves not at all decurrent to broadly auriculate ................... 64 64. Plants slender, thread-like; leaves mostly less than | mmlong.... 65 64. Plants larger; leaves more thanl mmlong. ..................... 67 O52 Branch leaves more than 0-5 mmilongee eee tee es os. - Homomallium Gsm ranchileaves less than’ 0). mmimylOme separ es See oe 66. Costa short and single; leaf cells rounded-elliptic ...... Pseudoleskeella 66. Costa none or very short and double; leaf cells rhombic .... Platydictya GYR beaves: Tal Cate=SGCunlld: s 1.4 1, scree ete remanence es eGo he cee, Gia weaves straight). 2:24 .as. 4 ee ee ean bee ae. iG 84 6s: Upper leat cells oblons-rhombiccae si Rees ee 6 Sek ee 69 68, leat cells more or less lincarsmonesthamOse Gr o5 6.52. 70 aD \O . Alar cells numerous, oblate, extending up the margins in many rows ee ee perm or cS 0 cea re ee Leucodon 69. ~] ~~] ~ os) ~l oe) | oa) ~l WS ~~ ~ ~— ~ CO os) (oe) eS) oe) WW CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 Alar cells few, oblong-quadrate, in 1—2 rows along the insertion Pee eet ee eee See eee ee ar ere e ey eee rs ee Sematophyllum 70. Plants regularly and closely pinnate, feather-like; fronds flat, erect to PSCOndINe OOMMNG-BanCuIaL. 25. beediw gee dc ede ee ees Prilium 70. Plants irregularly pinnate to unbranched, not feather-like; stems pros- trate 1O-loossly ascendia® = .) Capsules with a short, inconspicuous neck; costa filling 2 or more of the TAR AS Cee sere h acest Sea tins feather this ae sie OY mr Chet a 186 186. Leaves 3.5—5.0 mm long; upper laminal cells elongate, 75-140 wm lone ibroodsleaves absenin <2... 2 472. 6 nt ae ee one Campylopodiella 186. Leaves 2.0-3.0 mm long; upper laminal cells rectangular, 25-55 wm long; brood leaves usually present, often abundant ......... Brothera Plants pleurocarpous (sporophytes lateral); stems mostly prostrate with lateral branches, often mat-forming, or stems prostrate with erect branches bearing terminal sporophytes (cladocarpous) ............. 188 Plants acrocarpous (sporophytes terminal); stems erect, not branched or occasionally: branchedibeneath milorescences: ~~ .- ii 2 a eee ee os 280 188. Leaves bordered by elongate cells, sharply differentiated from SHrOmerelinnietnCelliswe wren rer a os «eke anya ce a Gr esem ewan ts) 189 188. leaves motnondercd by elongpate cells... | eee 19] Riantsepiphytics borden unisiratOSse —s.,... 6.5 Aes ee ee eee = Daltonia Plamtsraqiiaticn Dongen —S-SiialOSe: inc. atin. fn) ae cea mA ge, ese 190 190. Margins serrate almost to base; leaves ovate-oblong to ovate-lanceo- Inte acuminate: plantsom O1reconummra ser 2552 one a Limbella 190. Margins serrulate; leaves ovate, often broadly so, bluntly acute; plantsiof eastern North AmenCae rc ee Platylomella eaticells papillose, OF PLORUIOSCmmeee ayers oO he i hey eSateCellSssmmOO th: © a.z.5 eee ts rea More, china ate 205 ISPalbcaves micose;and plicatc see wser rye ne ey Rhytidiun 1O2 ke avesmot Tugase. fae lysplicalcime were 822 og ga e 4 193 leeat-cellsiplutapapllOse: 2c: caureae emerge gs errata. gph Ste, Wee een cc 194 leeat.cells, unipapillose OT piOnUlOCcHeen ss sees ee se a 198 194. Papillae arranged in a row over the cell lumina; cells longer than 3:1 194, Papillae randomly arranged over the cells; cells more or less iso- CLAS Tel Opis scar ea ea ep ee 196 Plants stiff, dark green to blackish; leaves ovate, plicate, cordate ee ee ER ne) CS nme ee ae an a a Papillaria 5. Plants soft, bright green (when alive); leaves lanceolate, smooth, not CONG AUS 0 cori, lea Ae ae opm tea cate noe ieee Barbella Great apices fragile (fy nee er te fh. a8: Haplohymenium SG e ceake amices Ot tails | sen near wet es gel eine PS are etek . Leaves strongly inrolled when dry; marginal basal cells elongate, smooth; SpoLrophytes terminal On branchesmp eee oe ee 5), Macromitrium . Leaves loosely erect to contorted; marginal basal cells rounded, papillose=sporophytes laverdlie~ eeweerreweet ie va eee yo ae Anomodon LOSseC@ellseunmipapiilose ovetsthe Iie ete yo es ee 1OSenCelllssprorulose: 4 nacre eee ae ein ety ey ea 20) Upper leaf cells linear-flexuose; uniseriate propagula common in leaf NSTI Sirs «56 Stas wtih os sce 3 ch cat Go Gre ft ec Henicodium Upper leaf cells short, 1-3:1; asexual reproduction absent or by branch- LSS ley Satin) oe tle io ene ele, i een ee a 200 200. Leaves ovate-lanceolate, narrowed to a nonpapillose, pale acumen, SqUAaTLOSe WHEN MOIS wan aes rer d cd) cael gh « Lindbergia 200. Leaves oblong-ovate, bluntly acute, loosely erect and more or less COmiplamate: 55... ak eee ee tials oR Stereophyllum seat cells sHoOnte IS eyes ee ee er ere tone 209. SF oa 2( CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 eet Cee Ile TUN Ek oink vwiew v8 heck abe ee eh os ek 203 202. Leaves appressed when dry, wide-spreading when moist; perichaetial leaves strongly differentiated; capsules immersed Lee Tee ee See eee ee ee eee Tre ee ret ces oe oY eee Cryphaea 202. Leaves little altered when moist; perichaetial leaves scarcely differ- CMS CASTES CXSCIIC kd ey as pee ce ea Lescuraea . Plants epiphytic with inconspicuous, creeping, primary stem and erect Secondary SteMin POsta SUDPEICUTIONE isc vsoueee ees wees Pireella . Plants terrestrial and epiphytic, with primary and secondary stems not differentiated; costa ending well below the apex ................... 204 204. Leaves decurrent; setae rough; capsules inclined; plants largely ter- Re Seis bette dhs lae sateen te cones eee ee Bryhnia 204. Leaves not decurrent; setae smooth; capsules erect; plants epiphytic OTRARICOOUR 4 £4c bp oew ey ands ee nee diie cee ie eee ks Isothecium ; bearapices rounded-obtuse to trumcate evauvuv id depp belteii sehen: ~ eal apices acuminate to DIUNUY aCUte: an dsss cp bacancreaaeecdaees PAC POO GAP ACeS TUNCIIES 4p dgeh iid eyes bddds edd eRdEeEISeS! Neckeropsis 206. ear apices COUNGEE-ODUISe <4 chs xrdnse rae uaeey cous teers 207 ADDS ear CCSICSGUNGNO. xcrteddespusanesyhecee case anes 208 \Jpper leat cells (not apical ones) more than S']. «accede sd ¥o ew ose eeus 209 208. Plants strongly complanate, shiny, little altered when moist (Ler sa eRe nee pauVNaeel eeunia he eee her cee kh eats re Homalia 208. Plants at best weakly complanate, dull, strongly incurved when dry, Srech WGN MOISE 266ecy en as nesueos abe aad pce a eee Leptodon PRIA CONS AM AUO Cu. 5:3- 4 apg cp wr ehiow ed wa ASUS tie ews ew REA ee Es 21 Alar eele tor at AU cs 4-4 ad oy Lanta eee ieee od ya Pete 211 210. Plants strongly julaceous; leaves concave; occurring in or near Sas “yc rics eee eee eds be eee ee Scleropodium 210. Plants with loosely spreading leaves; leaves only slightly concave; COCMMANEAN TONS: a32105 1040144032 ucaun anes eer Calliergon PROS ENMIMCIDE SPINE =. 2) ino avanrda evaded ue overneleu eke Platyhypnidium eta TO PIO eciNne Al APES g555c1eaiuenecxceadh oamieee Hygrohypnum Pion AP perieal CONS ie s5.0e sheave OPENS. CU OKG eet z13 oie. Wpper leat Cle MOTE MAN Sb ecicescdxe ae okes-eiorusune eels: 227 . Sporophytes terminal on erect branches; upper cells rounded ....... 214 Sporophytes lateral; upper cells rhombic to subquadrate .... 0... ... rae 214. Basal marginal cells short, rounded-quadrate ................. 215 Zi abel arena Celle Clonal vope cea Pe eraers ynkew< cane nes 216 . Upper leaf cells strongly bulging; leaves lanceolate; calyptrae hairy, GUS <7 pek ay ons Hosea ke Awa Seay eee OAS ee ss Macrocoma . Upper leaf cells flat; leaves oblong-lanceolate; calyptrae naked, cucullate Let R eG ee Rep eoms Me Amama nies ae fe Bk oe eee, Drummondia 216. Inner basal leaf cells subquadrate to rounded; calyptrae short, Be pido asi eae 2 10d cee ea eaas es Groutiella 216. Inner basal cells linear; calyptrae long, not plicate ...... Schlotheimia . Costa strongly flexuose above; leaf margins with multicellular teeth Ppa Reeth Aiea mie hate we ee ace ok eer eens ee Herpetineuron . Costa straight or faintly flexuose; leaf margins entire to serrulate .... 218 Zo; Costa ending inthe I6AT Apex 2c csc chaedey alle oubeuls'vantuerds Zi 210. Coste ending well below the IEAT ADEs ju jv 2viveeesen xxccadees aoe 1992 AM, 2A9: PENG _ Mp —" im) NW bdo Cd Oo 2 Z NK bM ~I~ tO NO \O 23), VITT & BUCK: KEY TO MOSS GENERA a7 Upper leaf cells thick-walled, rounded-quadrate to elliptic .......... 220 Upper leaf cells firm-walled, short-oblong torhombic .............. 222, 220. Plants with clusters of axillary brood branchlets ........... Leskeella 220 erlantsowithOul DrOoGmranchlets- os. 4, vt ict ieee eee: Alar cells extending to the costa; inner basal cells not differentiated; DeMIStOMen NC p thai Wate. eee 8 oe a ete tee ine eee ane Lindbergia . Alar cells restricted in 5-6 rows along the margins, not reaching the costa; inner basal cells long rectangular; peristome yellow Peer Mee Sot ae ah ee Oe ee me Oe eI re ot Pseudoleskeella DOee Costa relatively narrow, less than 35 4m wide at base; plants terres- Url caterer, eeu net sees ec tye a it Sac euete Pet Amblystegium 222. Costa relatively broad, mostly more than 50 um at base; plants ACU aU Camere Nie SAMI fr ee tc eee eee Hygroamblystegium 224 Plants with secondary.stems erect and’ branched ©2322. 5-82... . Plants without differentiated secondary stems, prostrate ............ 225 224. Leaf apices shortly acuminate; leaves more than | mm long; plants SAS UST Me Meee sete tas he ea rere i eCaGee ee aan Forsstroemia 224. Leaf apices broadly acute; leaves less than 1 mm long; plants west- STAIN de A area hie gir ae ee reas Porotrichum (Bryolawtonia) . Leaves acuminate, often homomallous .................. Pseudoleskeella sBCAVeSuOIUINU yea CUUC iq eck ee oo 8 ak eee en 294 294. Leaves bordered by 2 or more rows of elongate cells .......... 295 294. Leaves unbordered or if bordered then by only a single row of somewhat clonvate cells,-or by short cells: —. 125s) ee ee 309 Wecteniancumsue niin Wr crad alk che os te eter eh sae tenme en ace ey 296 eat margins toothed, sometimes obscurely SO” ......90. 226 ae: - 300 OIG leeal CellselOmeen thai 5 lee wares eee re tetas tt che ee 297 29 Gslee alece) Spleen eee ae tte ahi 0 Ry ec at 298 . Leaf cells in oblique rows; leaves rounded-obtuse, bluntly apiculate Be Fo wh ria hn hots Ts oes igi een een RE Pseudobryum . Leaf cells not in obvious rows; leaves variouS)«.. wwe eee Bryum 298. Rhizoidal (macronematal) initials in longitudinal rows; endostome fusedntoracdOnve™ a,c ealammeunepeneter eee we art deen an mae Cinclidium 298. Rhizoidal initials not in longitudinal rows; endostome segments free NGO a. yh otra a ee me cence Ae sg eben reemticcen Vaca ah? 299 . Leaves bluntly acute; border unistratose; stems blackish; stomates super- TGA lim erect ik ee eo AG io DO ee hes WS Cyrtomnium . Leaves emarginate, rounded-apiculate; border various; stems reddish, StOmalesumnvensed(* Gi. eaetre ye ema teat) fra, ot oe ares Rhizomnium 300. Leaves lanceolate from an expanded base, the base with numerous, aDTUpULy Myallive: COllSt um cwemra Gt Nee tee eek Syrrhopodon 300. Leaves without expanded base, without hyaline cells .......... 301 ealemmarcinsawith: palicad tectlipae eer pete ein Grae bey eke e 302 eal, mancins withesinele-Vce Nemaannmnnr trop eco es Sete re «oa 304 BO2elecatcells stronely DilOm gee eens ee eee ea ne te Trachycystis SU2ealseaticell se flat] as ant paqeeerweey tana et meee hes epigiediast o. 303 Some leaves with low, inconspicuous lamellae; costa in transverse section with two stereid bands; peristome of 32 small teeth attached at tips to a TyTN TIN, = 8s cs ose cae ee eat aaa vrs Atrichum No leaves with lamellae; costa with or without a single stereid band; exostome Of |G free teeth (eee came rere erate tens ttiinaiae Somer ad Mnium 304. Leaves rugose wet or dry; plants restricted to Pacific Northwest be eSte okie Se ek eer rere ee ulin os ee aie! Roellia 304. Leaves flat when wet, contorted but not rugose when dry; plants VIL SS PI AL ie cs ee oo ee TI eee a Aes eatcells;shorter than 221, teers tren cea Plagiomnium 62 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 SUS CACCIO SOP INONe teva ed 4 dw een tai oa sR eae cae s 306 306. Stems rosulate-foliate, occurring erect from a horizontal under- ground stem; sporophytes often clustered .....44 040.605 Rhodobryum 306. Stems foliate throughout, without rhizome-like connections be- Tween erect stems; sporophytes not clustered -..- 44s 2.902 307 307. Leaf cells in oblique rows; leaves rounded-obtuse, apiculate Sam Darn aie wane at PU dekh a Gin dk & OSPR SAE ee EE Pseudobr se 307. Leaf cells not in obvious rows; leaves broadly acute to acuminate 308. Capsules erect and symmetric; endostome with a high basal mem- brane, segments lacking or rudimentary, cilia absent SS ca Pe SMa toe ini ed GBR Ok Soar B46, dhe ash ES Brachymenium 308. Capsules inclined and asymmetric; endostome with a keeled basal membrane, keeled and perforate segments, and usually with cilia ee ee ee eee ee er ee eet er Bryum S02 (Gear Mmorcine With PAIGGTEEIN: yy 02nn da Radstone nie takers hes 310 200. Leatinarcins with single tecth Oriente «2c .sahok @ersasgo sew as a13 310. Leaf bases with abruptly differentiated colorless cells (cancellinae); PrOparula Dome On costa Near apex. .¢c.ida eee ee ees Syrrhopodon 310. Leaf bases without cancellinae; propagula absent ............. a1 Oe ed PCC Me IMOOU 4 ex uvansat en cen erkg eax see ele eet as 312 S11. Leaf cells prorulose or with fine cuticular ridees ©. 2.2.04 A oes Shas 318 Sie eeAves CIDE PO OVATe-CINNKO 15 csc2 bun gee eae a eee tees Mnium $12? Leaves linear lanceolate. .4is4 eae bag erxa Gon eee ees Pyrrhobryum 313. Upper leat cells with cuticular ridges; stems triangular in transverse section RHE os nt 4g ota 04 hah VERS aR Ste Pee Plagiopus 313. Upper leaf cells prorulose; stems round in transverse section ........ 314 314: Leaves crispate when dry 2. ius decade cians d¥edeovecaess Bartramia DiS seavOGRITAICNC WHINY yer eddie ae ee eens cole Philonotis 315. Leaf cells strongly bulging on one or both surfaces, not papillose .... 316 315. Leaf cells flat, smooth or papillose, or if bulging then papillose ...... 318 210; Sal amie iSIPAlOSe 5.25 5-84 x dna e Pax dae ee waned Timmiella Pie eal JAI AS UMISICAUOSE 5 yy apace ew 6 prawn Bae etek aa atals Ly ol. ae WISP eA DASeS. 2.2 ¥en vacated uae Rie eadaeeen Zimmia o1/. Plants without sheathing leat bases ..iccce bs. cyy a caea wes Bryobrittonia 318. Leaves with abruptly expanded, sheathing leaf bases .......... 319 318. Leaves with leaf bases not or only gradually expanded, rarely sheath- Cee re ed re eee re ee ee ee ere ae 326 Sido, eae cells papillose (at least on sheath) or prorulose ............... 320 Oey COE COMscSINOOLN - “24254489 ceengamues keae aw eneeen te twee ere SZ Oo eat Celle papiloseOver (hE ANMINA 4'.ncu\apeageutee ccake he ZTimmua S2Us ear Cells rOmose?, vo00 ne hay Ga4-4cean came cee ereine + Wes Oe Bartramia Sc ly AIpper ea pCe le QUAdEALe: ~ .464s ndcen oe Saas open oem eens ORE eS a Ol. A Ppemieal Cells Sorereciqneulat © s1¢.10..o6svccw meee eee PO mre ie 323 322. Capsules inclined and asymmetric, strumose; plants widespread ee eee ee ore ei eet, eee rot tee Oncophorus 322. Capsules erect, symmetric, not strumose; plants restricted to south- Werle: ght tune e dsm che easier Ry Symblepharis 323. Awns roughened throughout by projecting cellends ........... Trichodon 2295 AWS SODEMOrOnIVTOUPN at ENEx x4 .aded eae eee ee es 324. Capsules with neck as long as or longer than the urn”... Treniaiocon 224; Capsules with neck much shorter than the tim? .0io4.cy4082 56589 325 Wd Oo Gd G © © 341. 341. VITT & BUCK: KEY TO MOSS GENERA 63 5. Capsules erect, cylindric, smooth; peristome teeth irregularly perforate or deeply cleft into terete, sometimes filiform divisions ........ Ditrichum . Capsules inclined, oblong, smooth or furrowed, if erect then furrowed; peristome teeth flat, split % way down, vertically pitted ........ Dicranella o7 Oa Nalgccl |Sienlanecdmcoloned Or inflated: 2. ue ery ye 2 S20 NlamcellSiscance Waite nem tated rs cox) ances sateen Rate a 333 . Upper and median leaf cells with coarse, irregular cuticular ridges, resem- bling papillae in transverse section; capsules cylindric, smooth, erect Es ee Mn oa a eg ine ye hE Olona chan ye ae ON mame SEAT 328 . Leaf cells without cuticular ridges; capsules various ................ 529 328. Leaves with clusters of spherical propagula at leaf apex; leaf mar- PANS MO IS ETAGOS Cx area seed fT Ri 10 ON ONO ten ee ge RRS Grimmia 328. Leaves without propagula; leaf margins unistratose .... Dicranoweisia Be COS) WItMeSLCheLOP DANO S ee a2. 5. got at av mani sa eran ey ements e Dicranum FeO OST VDI OURS UC TOT Lie Sg te oh aa eh ye ata ng ese oe 330 330. Capsules curved, strumose; plants on alpine rocks ........... Kiaeria 330. Capsules erect, not strumose; plants widespread .............. 331 . Capsules cylindric; plants usually terrestrial or on tree trunks, rarely on TOG) cA aes atest Ss sd AR ey ree ee 2 Ko Rene en Dicranum . Capsules short, obovoid to pyriform; plants on rocks .............. 332 332. Capsules ribbed when dry; peristome vertically pitted-striolate be- LOW Aare ihren relies, 4 Gols rere ere ae Pra EE A ccicay ic. 'a 2a Sl RS Ree a Arctoa 332. Capsules smooth when dry; peristome papillose ............. Blindia mlSCatcellssimOOthy 62a. ere Sosy ern at. 334 mlbealecellsepapllose- Or PLOnul OSG emer eee son he eee ae, 398 334. Asexual propagula borne on the leaf surfaces ......... Rhachithecium 334. Asexual propagula absent, axillary or on leaf apices or specialized SS UTEUN GTS Si ots tet asa eae RS if, mete Burt Sea 535 . Leaves ovate to obovate, broadly obtuse, with the costa ending below the LS ATRAD ENS Seiler A007 ete ee eR PR Str talc ah ao Nas ol 2 . Leaves linear, ovate-lanceolate, ligulate to ovate, acuminate to acute or awned, with costa ending below the apex to excurrent ............. 342 Son Wealcapices uct Ale aaa ee eee ae Globulinella DO Oneal APICeo Ol CUCU Aiea mmae tree men muni ieetn ta te, ces e One or eet 537 . Plants with large, multicellular propagula in leaf axils; leaves obovate, withvelliarvan the-basal Maneins een eee, | ee Oedipodium . Plants without axillary propagula; leaves ovate to spatulate, eciliate .. 338 338. oe small and bulbiform, on dry soil; leaves concave and imbri- BAS ERT ce eer cere 1 cis A Ree ee Stegonia 338. nae small to medium-sized, never bulbiform, on moist soil; leaves flat tOxcOncave smoOsty NOt ImMbRICalCe fen ou = See cas 339 . Leaf cells short-rectangular, lax, thin-walled; capsules erect ......... 340 . Leaf cells long-hexagonal, firm-walled; capsules inclined to horizontal Se tag RI erie eens Ce ee a ee ee 34] 340. Apical marginal leaf cells short-rhombic, oblong-hexagonal inter- nally; capsules without a distinct neck; plants of southern United SEATS sa So ae ae, else Splachnobryum 340. Apical marginal leaf cells laxly rectangular, similar to internal ones; capsules with well differentiated neck; plants northern ...... Tayloria Plants whitish-green, terete; upper leaf cells long, 7-9:1 .... Anomobryum Plants reddish or green, not terete; upper cells shorter, 2-6:1 ..... Bryum 64 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 Sa2 eaves lone Jineulate TO OVEIC’ «a; denis tetee ee tats weenees 343 Sad SVedves lanceolate IO NNCAl ° cb:nede dada sGnage ns ere ee ees 368 343. Leaf cells rounded to quadrate, more or less isodiametric ........... 344 e435. Dear cells suoperectancular to linear (2212. euevereserenes eer 352 344. Leaf bases with abruptly differentiated, hyaline cells (cancellinae); leaf apices often with propasila o..14 2107 dees eae es Calymperes 344. Leaf bases without cancellinae although sometimes gradually hyaline; leat apices without propapula | ics. ca es aes eee os 345 345. Costa excurrent to ending in the cusp; leaf apex cuspidate to piliferous 345. Costa subpercurrent; leaf apex broadly acute or obtuse to rarely mucro- 0 ne re ae ee ene re ee ee ee ene er ee ee re SHO eat apices PIMCIOUS. 4.25 iucses costs ees Desmatodon 240; edt apiece CUstIdale. 14 ci cy haerre veins MOOG uate ees 347 347. Plants with rhizoidal tubers, known only from Louisiana; capsules unknown WA EQA Rae ee See haan ba eS Oe o8 SE OL ee Tortula (Chenia) 347. Plants lacking tubers, widespread in the north; capsules exserted .... 348 348. Capsules 0.4—0.8 mm long, obovoid; spores more than 20 ~m in diameter; peristome absent or rudimentary; plants boreal- FOIE MO sobs s Siacee eee ernie. es. ede ks Pottia 348. Capsules 2.5—3.5 mm long, cylindric; spores less than 20 wm in diameter; peristome well developed with twisted teeth from a basal Membrane, Plants OL NIMOla padre hia raha eRe ea eee es Crumia 349. Peristome teeth four; propagula cups borne on apices of sterile shoots ee ee ee ee re ee eer ee ree Sr en ees ye. see eee 2 Tetraphis ooo): Penstome tseth 1G; propacuiacups aDSenl: 44) ..eane noir ak ee es 350 350. Leaves homomallous, broadly oblong to oblong-ovate, subacute to OWINSE: witeet tac tassdyie otid epie-s8 Rate dees Aulacomnium 350. Leaves not homomallous, oblong to lanceolate, mucronate to POUNe psy Cuneta ene G es ReSeET EE ee ea es ] 351. Leaves broadly to narrowly lanceolate; peristome double ......... Meesia 351. Leaves broadly oblong to spatulate; peristome single .............. og 352, Costa. in transverse section With 2 Steicia Daas 4 ease cases Barbula 352. Costa in transverse section with 1 stereid band ......... Scopelophila 353. Plants filiform, julaceous (if costa not excurrent see also Anomobryum) Ee er eee ee ee ee ere ee re ee ree te ee Aongstroemia S55; Fiaie COdtech. MOL MaCEOUS: 114.402 95 45245005 chee eek 4 ekg ee re Soe, (Capsiies DOmZOntal TO DENGUIOUS:.. 2ikicna ve Pea eee bea 525 Soe eeu leseleel-” occ eexwag a ture ee hoy Wee era oer eed 360 D> Na DSUeSCUIVeU aSVININEIe 24.25 ou crete tara eae ene pe eee ees 356 Sys) CApsiics salen SVINIMCLUIC fx ~ shay dy hdd oe Meee ba ba ees B57 356. Exostome shorter than and alternate with endostome ... Plagiobryum 356. Exostome longer than and opposite to endostome .......... Funaria 357. Capsules ovoid, less than 1 mm long; stomates none; endostome fused to SxOstOnie, CIA NONEY 4uisdeare caved cuba pee see een aes sas Discelium 357. Capsules cylindric to ovoid, usually greater than | mm long; stomates present; endostome free of exostome, cilia mostly present .......... 358 Soe, deeat CEISIN GDNGUETOWS un.0idvadu eink gue eeeee or ene DUaeIOOrUn Sos edt Cells NOt OOVviCls OUNGUGTOWS ¢425.0%9040 ee hdox cae Steen 359 cab eat celle AS Orless. ooh cues ieee peeteensah oye Grad tee oae ees Bryum 1992 361. 3: Nn ww 369: 369. SAk St 3 | oa) VITT & BUCK: KEY TO MOSS GENERA 65 pele altace! Seo On MON CC lay sae hee ee a ae oe es Gta at Pohlia SGU MPCTIStGMe a DSC Miter) ght eres ia ae eg, Porn Morn aces. ss 361 BOOMREDISLOMeN NCSC Min se ales shea pee eee ee eg eae eee 363 . Calyptrae 4-angled, sheathing the entire capsule until after dehiscence Ne ere a Ree Re AOA Talay drei aS CG: oc A, ye eo eR em Pyramidula Calyptrae not angled, not sheathing or persistent .................. 362 362. Capsules subcylindric or narrowly pyriform; annuli none; exothecial cells oblong to oblong-linear; calyptrae inflated-cucullate ...... 362. Capsules urceolate to broadly pyriform; annuli present; exothecial cells irregularly hexagonal; calyptrae inflated-mitrate Ea see ert APG Ae esse Sh oe eae ae Physcomitrium . Peristome of endostome only, without a center line on the outer surface Sa a tee Besant et acto artes Ck Es ohh ee ae a Mielichhoferia . Peristome double or of exostome only, with a center line on the outer SUITRACEE (ate a ah ee are Ae Cw Dn ecb CREO ae cae 364. Capsules with scarcely differentiated neck ................... 365 364. Capsules with well-differentiated neck (apophysis) ............ 366 Blantsotianctic tum ase eee, oor, eee ee: Funaria Plants of British Columbia (50° N) and south ............... Entosthodon 366. Urns black, sometimes with yellow apophyses .......... Tetraplodon 866) Wins and: apophyses Crcem (O DIO WN myers oes faded oe ee 367 . Setae pale greenish-white, slender; costa filling the acumen, plants of eastenne NOnuieAImenica:. , aera eee cre ee ead Splachnum . Setae brownish, stout to slender; costa ending below the leaf apex; plants Ob western and northern NonthgAmencay 22... 233 G Soot ness. ayloria S68: eaves COnsplcuoUSly 4-1anked) aie a Conostomum 368. Leaves not conspicuously ranked (if 3-ranked see Meesia) ...... 369 eat cells:short, 1(—2) sl rounded toquadrate ait ts 370 Leaf cells long, (3—)4:1 or longer, rectangular to long-hexagonal .... 386 370. Perichaetial leaves strongly differentiated, with an awn as long as the laminas leaves bistratosemums cee. oe ee Diphyscium 370. Perichaetial leaves slightly or not differentiated, never long awned; [Saves Uni- tOmmUltiSthalOSc merece Soa ees 371 Capsules distinctly 8 or 16 ribbed and furrowed ...............--.. 372 Capsules smooth or indistrictly furrowed when dry but without distinct Gil SMMR Ss Grete ee Sc icine Sem re ree ets Chetan We et Oe 6/2 4Calyptrac Mitlate’ > aimee ime peer nete teh eae eeavi ta ahh S76 Slee @alypinaercucuiilate, jatar nee eminent raat on) es Rega e oa ve S75 Galyotacheiny: 2h, Savi scskei de tenw edd bese ees Aulacomnium 408. Plants with or without various means of asexual reproduction but these never borne on stalks from the stem apices .............. 409 Plants occurring on tree trunks or bare rock, with immersed to shortly EXSCLICC MONE MIDDEG@GANSUIES . «x s.an-cx won hada eem arose ae eee Plants usually occurring on soil, if on rocks or trees then capsules long exserted and mostly unribbed (or plants sterile). ..54.0.4-2) das Shands 410. Calyptrae cucullate, not plicate, naked; plants occurring on wet rocks Fb 6 Pruter aecdlehy gesinh tind 44 8% v4 x GH 0-4 4 ae Amphidium 410. Calyptrae mitrate, plicate, usually hairy; plants occurring on tree MiUnks ane Athy TOCKS 2 .0rs nten ese ache cote eee ae es | . Leaves usually crispate to contorted when dry; basal marginal cells with thickened transverse walls; stomates superficial; capsules shortly CASOUCOS -$41R yh ide uid ens a baie eos ores Mee ee oe ota . Leaves usually little altered when dry; basal marginal cells not differenti- ated; stomates immersed or superficial; capsules immersed to shortly Cet eds ih 4 at sens ba dutenne eed ccs eee eee rthotrichum 412. Costa in transverse-section more or less homogeneous, without dif- TOVCDNGtOd SICIOIIS: -yhasawseyweaw anaes aOR ere we eees 413 412. Costa in transverse-section with differentiated stereid bands .... 417 . Leaf cells with C-shaped papillae; leaves broadly spatulate Sis Sok fete ol is atic: hugs & fp Gerd ba a kag el Reet Og Gymnostomiella . Leaf cells with simple, conic papillae; leaves various but not broadly SOG. 445-45 G58 Hae oda shoes eee WRU ees ne meres 414. Upper leaf cells with 4—7 small, conic papillae ............ Zygodon 414. Upper leaf cells with 3 or fewer, simple to branched papillae... 415 . Leaves less than 0.6 mm long, ligulate, rounded-obtuse; rare plants of calcareous rocks; annuli of 2—3 rows of well-differentiated cells eS eee eee ern ee ee ee Cae aie eres ie ee etry ee en ee Gyroweisia 5. Leaves more than (0.7—)1.5 mm long, lanceolate to ovate, acuminate to Obiusesannull POOL y dilterentiated ...-.2 54 ds usaa4 Gee ee Rete oe a 416 416. Leaves typically crispate to contorted when dry; basal marginal cells with thickened transverse walls; capsules shortly exserted; calyptrae ely? ite SS oGF ieee hit G ee eye ka neers 6 eure oe Ulota 416. Leaves usually little altered when dry; basal marginal cells not differ- entiated; capsules immersed to shortly exserted; calyptrae naked (species with contorted leaves) to hairy (species with unaltered ISAS)? 754114 a oddy booed eke beds see A Orthotrichum Costa witha single, dorsal stereid band juie cedar ed eeds4 ud bases Costa with both dorsal and ventral stereid bands .................. 429 418. Calyptrae campanulate-mitrate, covering entire capsule, often lobed at base; basal leaf cells with thickened transverse walls Dis stalu acite PW Sh aoa nize tee ais A ad PE ee ee Encalypta 418. Calyptrae cucullate, covering only operculum and capsule apex, un- lobed at base; basal leaf cells usually without thickened transverse Male Sie bucsusyn Sees one ows Pune s ee Ree rek ae roa 419 1992 419. 4 — \O 421. ADB: 4 bo eS) 427. 4 NO ~ 429, 429. VITT & BUCK: KEY TO MOSS GENERA 69 Leaves linear-lanceolate to lanceolate, never with hair points; leaf mar- gins recurved to plane; leaf cells with papillae conic, clavate or branched, ab eI EAS) oe OX Aree eee ern oe rN rs Teenie Sarees tO rs Persea 420 . Leaves broadly lanceolate (from revolute leaf margins), lingulate or oblong-ovate, sometimes with hair points; leaf margins recurved to strongly revolute; leaf cells with papillae stellate from a stipitate base to (CAS) ary oforal games Clee rate, ey or) OMe ear arama errr tree Say ROP oer tte 2 eee 424 420. Leaf cells with 4-7 widely spaced, small, conic papillae; elliptic propagula present in leaf axils; peristome double; plants usually OCeUmngeonatheess marely On TOCK (2254. t4 ae a eee Zygodon 420. Leaf cells with 1—3(—4) closely set, simple to branched papillae; propagula absent; peristome single or absent; plants occurring on S SO Pane OCKSe ow. stewed ghia s cc. neen ay eee en oes 42] . Leaf margins serrulate above; capsules erect to inclined, often furrowed and strumose; peristome vertically pitted-striolate .......... Cynodontium Leaf margins entire; capsules erect, never furrowed or strumose; peri- STOMem Me MED ESeM Ur DAP OSe” a aixkh ead egat een Roa amma tae « 422 422. Basal leaf cells lax and hyaline; upper leaf margins bistratose ChHTOUCH OUT PETISLOMME PLESCDU > ic. j.ek eee ead shear Trichostomopsis 422. Basal leaf cells firm-walled, pale but not hyaline; upper leaf margins unistratose or irregularly bistratose; peristome absent .......... 423 Stem monopodially branched; archegonia on short, lateral branches; stem transverse section rounded-triangular) ............... Anoectangium . Stem sympodially branched; archegonia terminal on main stem; stem (aN SViCTSe Se CLIOM: TOU = sugar meer ner eee ts St pene Gymnostomum 424. Leaves with an intramarginal band of enlarged, smooth, often WTAMCENCCIIS: “yas ine ae eer eer men eee ha Crumia 424. Leaves without differentiated intramarginal cells but sometimes with smooth marginal cells, these not enlarged... ... 1... 425 . Leaves narrowly lanceolate; leaf margins erect and bistratose Trichostomopsis . Leaves oblong, lingulate to ovate; leaf margins plane to revolute, uni- stratose or if bistratose then always revolute ....................04. 426 426. Upper leaf margins broadly revolute to spirally revolute, with cells often more strongly chlorophyllose than median cells .......... 426. Upper leaf margins plane to revolute, with cells undifferentiated or paler than mie dianscell simran eer Shs eae ees cco: 427 Leaves unbordered; peristome none or rudimentary ............... Pottia . Leaves often bordered by smooth cells; peristome present .......... 428 428. Peristome teeth united in a high or rarely low, tubular basal mem- brane, spirally twisted above; cells of adaxial (upper) surface of costa similar to or smaller than laminal cells in transverse section Ref eehfo ieiacth AOA Se tee ne ad Bhi ys Tortula 428. Peristome teeth free or rarely united in a low basal membrane, erect to slightly twisted above; cells of adaxial surface of costa mostly larger than or otherwise differentiated from laminal cells in trans- VETSE SCCELOM i okie ere er ee ee Sesh ae A Bh Desmatodon Leaf margins abruptly serrate at the shoulder ............... Eucladium / Keatimarcins entite at ShOUldeTSHrmeemrte ree ts acs aes es 430 431. 431. 437. 44 4 ' og os) o 3. Peristome usually present; leaves recurved on both sides ........... CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 430. Leaf cells with numerous (more than 7), minute, elliptic papillae Pine wales as well asinine lumina: sacdy.ceeu eens + Amphidium 430. Leaf cells with fewer than 7, larger, round papillae only over the PUY % Sgeatcp cite tne tea igs siete ayo Sea ep aes a how ene eee aes 43] eal Mareme TOGINEC BOVE, oy . — 4 “ oe oe Contr. Univ. Mich. Herb. 18:99-103. 1992. WHAT IS LEJEUNEA TRIGONA? (Studies on Lejeuneaceae subfam. Ptychanthoideae XXII) S. Rob Gradstein Institute of Systematic Botany 3584 CS Utrecht, The Netherlands Lejeunea trigona was described by Montagne and Nees (1836) based on a liverwort specimen collected in Peru by the French traveller and palaeontologist Alcide d’Orbigny. The material consists of only a few stems, which according to the original description were taken from a lichen (“in Parmelia speciosa parasita”). Although the material is fertile and has both antheridia and perianths, there has been some confusion about the identity of Lejeunea trigona, presumably due to the poor condition of the type specimen. The authors of the Synopsis Hepaticarum (Gottsche, Lindenberg and Nees 1844-1847) considered the species to be most closely related to Lejeunea chrysophylla (Lehm.) Gott. et al., an African taxon now placed in the genus Acanthocoleus (Kruijt 1988). Stephani (1911), however, treated Lejeunea trigona as a member of the genus Archilejeunea. More recently, Lejeunea trigona was studied by Kruijt (1985) who considered the species a synonym of Lejeunea phyllorhiza Nees 1833. The latter is a widespread neotropical species, which was usually placed in Dicranolejeunea but was transferred to Brachiolejeunea by Kruijt and Gradstein (1986). In the course of my work on neotropical Lejeuneaceae for Flora Neotropica, | have recently had a chance to reexamine the material of Lejeunea trigona. It ap- pears that the species has affinity neither to Archilejeunea nor to Brachiolejeunea phyllorhiza. The rather flat leaves with reduced leaf lobules and the smooth perianths without or with broadly rounded keels of Lejeunea trigona would rule out B. phyllorhiza, which has concave and more or less squarrose leaves with large, unreduced lobules, and sharply keeled, ciliate perianths surrounded by winged female bracts. Another important characteristic of Lejeunea trigona is its thin stem, which has a distinct hyaloderm and a ventral merophyte which is mostly only two cells across. Both Brachiolejeunea and Archilejeunea have broader ventral mero- phytes, being four or more cells wide. In Archilejeunea, moreover, a hyaloderm is lacking. Because of its narrow ventral merophyte and other morphological features, Lejeunea trigona undoubtedly bears closest resemblance to members of the genus Acanthocoleus (subfam. Ptychanthoideae). This genus was recently segregated from Dicranolejeunea by Schuster (1970) and was subsequently monographed by Kruijt (1985, 1988) who recognized seven species, two of which occur in tropical America. Diagnostic characters of Acanthocoleus are its rather unspecialized stems, lacking a subepidermis and _ bulging epidermis characteristic of Dicranolejeunea, its untoothed female involucres, its creeping habit and its brown- ish color. A comparison of Lejeunea trigona with the species recognized in Acanthocoleus shows that it is different from all of them. Its most striking feature is the strongly inflated perianths which are barely keeled and completely smooth, lacking any trace of cilia. None of the species of Acanthocoleus has such perianths. Furthermore, the antheridia of L. trigona are borne singly in the axils of leaves just 99 100 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8& FIG. 1. Acanthocoleus trigonus (Nees & Mont.) Gradst. below the perianth, and the species should thus be considered paroicous. Within Acanthocoleus a paroicous sex distribution is characteristic for A. juddii Kruijt from the Greater Antilles and Mexico, and A. chrysophylla (Lehm.) Kruijt from Africa. These two species are very different from Lejeunea trigona, however, by their keeled perianths with ciliate margins and their apiculate leaves. Lejeunea trigona thus appears to be a distinct species of Acanthocoleus and the following new combi- nation may be proposed: 1992 GRADSTEIN: LEJEUNEA 101 Acanthocoleus trigonus (Nees & Mont.) Gradst., comb. nov. Lejeunea trigona Nees & Mont. in Montagne & Nees, Ann. Sci. Nat. Bot., Ser. 2,5: 61. 1836; Archilejeunea trigona (Nees & Mont.) Steph., Spec. Hep. 4: eles LONE Fic. 1. DISTRIBUTION The distribution of Acanthocoleus trigonus has been remarkably poorly known and for more than a hundred years only the scanty type specimen from Peru has been available. As a happy coincidence, however, I have recently been able to rediscover the species in southern Bolivia (Dept. Tarija), during a collecting trip made in the company of my wife and the bryologist Marko Lewis, a curator at the National Herbarium of Bolivia in La Paz. Acanthocoleus trigonus proved to be quite common in humid Podocarpus forest remnants in mountain valleys, in areas with a prolonged dry season and at elevations between about 1600-2600 m. (Fic. 2). The species was always growing in thin mats on boulders along streams. In the field, the species could be easily recognized with a handlens by its peculiar perianths, which were often completely terete and devoid of keels. Searching through the unidentified liverwort materials of the herbarium in La Paz, I subse- quently found further specimens of A. trigonus collected by Marko Lewis in the same general area. In addition, two further collections of A. trigonus have recently become available: one from southern Brazil, collected by Alfons Schafer- Verwimp and his wife, and an older collection from northern Argentina (leg. Hosseus), described by Herzog as Archilejeunea argentinica Herz. (Feddes Repert. 55: 12. 1952). The latter should be considered a synonym of Acanthocoleus trigonus. 3. 2. Remnants of Podocarpus forest in mountain valleys at about 2000 m near Entre Rios, Dept. Tarija, southern Bolivia. Acanthocoleus trigonus occurs on rocks along streams in these forests. 102 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 It thus appears that Acanthocoleus trigonus occurs in relatively dry, subtropical mountain areas of South America (Peru, Bolivia, southern Brazil, northern Argen- tina), in regions with a seasonally dry climate. The Lejeuneaceae flora of this region is rather poor and most of the taxa reported are widespread, neotropical species. Among the Ptychanthoideae, Acanthocoleus trigonus is the only species endemic to this region. An explanation for the poor Lejeuneaceae flora of subtropi- cal South America is probably the rarity in these areas of well-developed evergreen rain forests, which are the main habitat for the species of this family. Support for this assumption comes from a comparison between the Lejeuneaceae flora of South America and Australia (Thiers 1990, Gradstein 1991). As demonstrated by Thiers, the subtropical east coast of Queensland has a rich Lejeuneaceae flora with several endemic species. This high diversity is due to the occurrence in this region of rain forests, distributed in patches from north to south along the coasts of Queensland and New South Wales and ranging well into the temperate zone. These forests offer a great variety of habitats for inhabitation by Lejeuneaceae. A similar latitudinal gradient of rain forest is lacking in the New World, however, and this, then, may well account for the poverty of Lejeuneaceae in the subtropical regions of South America. KEY TO THE NEOTROPICAL SPECIES OF ACANTHOCOLEUS — Lobule large, 2-4 lobe length, rarely reduced, first tooth of the lobule 3-6 cells long; leaves orbicular; paroicous; Greater Antilles, Mexico, ca. 0005010 ae ee ee ee ane cee ee Sea Sees aT A, jJuddii Kruiyt . Lobule smaller, %43-*4 lobe length, often reduced, first tooth 0-2 cells long; leaves ovate; autoicous or paroicous. 2. Paroicous; perianth terete or very bluntly keeled, smooth; leaf apex rounded, entire; S Brazil, Peru, Bolivia, northern Argentina, ca. LODO OO ciis. en keene at eo ces A. trigonus (Nees & Mont.) Gradst. 2. Autoicous; perianth sharply keeled, ciliate; leaf apex pointed, rarely rounded, toothed or entire; throughout the neotropics, 150-2600 m. eee Sate ie ee hea A. aberrans (Lindenb. & Gott.) Kruijt — A full description of Acanthocoleus trigonus will appear in my forthcoming monograph of the neotropical Ptychanthoideae in Flora Neotropica. Specimens examined. Peru. LAGUNA: Santa Cruz, “in Parmelia speciosa”, D’Orbigny s.n., is of Lejeunea trigona Nees & Mont. (STR, holotype; PC-Mont., isotype). BRAZIL. SANTA CATARINA: Serra do Corvo Branco, road Urubici-Crao Para, just below the pass, Dec 1990, Schdfer- “Ver ae 13496 (hb. : y. BoLivia. CHUQUISACA: Rio Jatum Mayu 14 km NW of Zudanez, Lewis 84-0489 (LPB, U). Tarua: along road Tarija-Bermejo, ca. 10 km N of La Marmora, Bate: er 7/21 (LEB, U); headwaters of Rio Los Pinos 20 km NW of Padcaya, Lewis 84-2910 (L ; headwaters Rio Posta along road Tarija-Entre Rios 1950 m, ela 7648 (LPB, U); een Rio Huayco oe road Narvaez-San Josecito, Gradstein 7687 (LPB, U); along Rio Tambo ca. 5 km S of Narvaez, Lew 84-2675 (LPB), Gradstein 7051 (LPB, U); Cerro Sarzo, 4 km NW of Canaletas, Lewis 84- 2510 (LPB). ARGENTINA. CORDOBA: Ongamira, Hosseus s.n., type of Archilejeunea aie Herz. (JE, holotype). & mR Be fe) zé& Sa = =I 3 e ACKNOWLEDGMENTS express my gratitude to Mariette Aptroot-Teeuwen for preparing the drawing of Acanthocoleus trigonus and to the curators of the herbaria cited for the loan of specimens. 1992 GRADSTEIN: LEJEUNEA 103 LITERATURE CITED eo is M., J. B. Lindenberg and C. G. Nees von Esenbeck. 1844-1847. Synopsis Hepaticarum. urg. ee a R. ree ie and distribution of Asian Lejeuneaceae subfamily Ptychanthoideae. Bryol. 4: Kruijt, R. 1985. A ee revision of the genus Dicranolejeunea (Spruce) Schiffn. Beih. Nova Hedwigia 80: 155-163. 1988. A monograph of the genera Dicranolejeunea and Acanthocoleus. Bryophyt. Biblioth. 36: 1-135. and S. R. Gradstein. 1986. On Brachiolejeunea phyllorhiza (Nees) Kruijt & Gradstein comb. nov. Nova Hedwigia 43: 299-309. ee J. P. F.C. and C. G. Nees von Esenbeck. 1836. Jungermanniearum herbarii Montagneani Spec ae Sci. Nat. Bot., Sér. 2, 5: 52-72. Schuster, R M. 1970. ie on Hepaticae XLIX—LIII. New Lejeuneaceae from Dominica and Ja- maica. Bull. ae Bot. Club 97: 336-352. Stephani, F. 1911. Archilejeunea. In: Species Hepaticarum 4: 703-736. Geneva. Thiers, B. 1990. An overview of the Lejeuneaceae in Australia. Trop. Bryol. 2: 273-283. Contr. Univ. Mich. Herb. 18:105. 1992. MOSSES AS USEFUL PLANTS A. J. Sharp Department of Botany University of Tennessee Knoxville, TN 37996-1100, U.S.A. When the uses of mosses are mentioned, the ones usually thought of are the peat mosses (Sphagnum spp.). These have been used for centuries in the form of peat which when dried can be used as fuel furnishing heat in their habitations. It also was used as a dressing on open wounds in war, and also in forestry accidents. It contains sphagnol (an antiseptic phenolic compound), and when dried it is also very absorptive. This also led to its use as menstrual pads in the past. Mosses are also collected for ornamental purposes in florist shops, and also are used in terraria which has led to the decimation of forest mosses in some places in the Southern Appalachians. In the streets and on store fronts of the villages in Mexico and Guatemala, pendant forms are frequently used for decorations during their festivities. In nature, mosses growing on logs and decaying forest residue tend to hold moisture which facilitates the return of woody plants, and acts as fertilizer for the next plant crop. Less well-known is the use of mosses by animals for food and other purposes. | call attention to the following in order that we may have a more thorough and scientific knowledge of another aspect of the significance of mosses in the natural environment. Twice in the Southern Appalachians I have seen mice eating the sporangia of Polytrichum and Pogonatum. Unfortunately, I made no record of the exact locality and date. Several times in Alaska I have encountered lemming pathways which had been tunnels under the snow. Most of the mosses in these runways were devoid of capsules, and in some instances, the setae were missing. Again, I failed to record the name of the mosses, the locality, or the date. I am sure that one of the mosses was a Ceratodon. This indicates that mosses can be of importance to some of our furbearers, foxes, wolves, and cats, that feed on rodents. Thus, a woman could be wearing a furpiece derived indirectly from mosses In another instance in Alaska I encountered a wasp nest in a relatively dry hummock of peat moss (Sphagnum). They pointedly let me know that it was their territory. Thus we have a moss home for wasps on which some predators feed. Again, I kept no locality or date. I write this as a challenge for bryologists to record data that we need to under- stand the full importance of mosses in the environment. I also write this as a salute to Dr. Howard Crum who was not afraid to tackle a big problem when he first worked on his Ph.D. dissertation concerning the mosses of Mexico. 105 Contr. Univ. Mich. Herb. 18: 107-111. 1992. STUDIES ON GEOCALYCACEAE VII. SUBSPECIFIC DIFFERENTIATION OF CHILOSCYPHUS SEMITERES TOGETHER WITH FURTHER REFINEMENTS IN CHILOSCYPHUS (S. LAT.) John J. Engel Department of Botany Field Museum of Natural History Chicago, IL 60605, U.S.A. Chiloscyphus semiteres is a pan-south temperate species occurring in the Aus- tralasian, American and African sectors (Fic. 1). The species occurs in variable frequencies in different parts of its range. It is very rare in the Falkland Islands (Engel, 1990), probably lacking in the Magellanian zone of southern South Amer- ica (I did not record it for the Brunswick Peninsula [Engel, 1978], and the few other Magellanian records are suspect), and somewhat common in the Valdivian zone and on Juan Fernandez. In the African sector the species is restricted to Cape Province, South Africa, where apparently it is rather common judging from the number of specimen citations in Grolle (1959). The species is of variable frequency within Australasia: it is rather common in New Zealand, but in Tasmania and Australia C. semiteres is one of the most common or perhaps the most common hepatic. The species also occurs in the oceanic Isles of Scilly (Paton, 1965) and Scotland (Long, 1982). Chiloscyphus semiteres is characterized by a) undivided, entire leaves that are broadly rounded to truncate at the apex; b) conspicuous, deeply divided un- derleaves that frequently are asymmetrically inserted and have the laminal margin on each side with a process grading from dentiform to lobuliform; and c) a variable gynoecial position: some to many on short lateral-intercalary branches lacking normal leaves, while others are on terminal branches and main shoots. The species exhibits subspecific differentiation only in the Australasian sector, where it may be separated into two varieties as follows: KEY TO VARIETIES OF CHILOSCYPHUS SEMITERES — . Underleaf disc lateral armature well-developed: mostly laciniate to lobu- late; trigones minute to small, only occasionally medium and straight- sided; underleaves often 1.5—3 x stem width; plants translucent te et ee a ae ee eae nl eich ee C. semiteres var. semiteres . Underleaf disc lateral armature reduced: mostly dentiform to ciliiform or lacking, only sporadically small-laciniiform; trigones normally large and bulging to knotlike (in shade forms medium and straight-sided to weakly bulging); underleaves normally 1-1.5x stem width; plants dull-opaque C. semiteres var. canaliculatus — Chiloscyphus semiteres (Lehm.) Lehm. & Lindenb. var. semiteres Jungermannia semiteres Lehm., Linnaea 4: 363. 1829; Chiloscyphus semiteres (Lehm.) Lehm. & Lindenb. in G. L. & N., Syn. Hep. 190. 1845; 107 Chiloscyphus semiteres (Lehm .) Lehm & Lindenb. var. canaliculatus (Hook. f. & Tayl.) Engel A 108 FIG. 1. Distribution of the two varieties of Chiloscyphus semiteres. 1992 ENGEL: CHILOSCYPHUS 109 Lophocolea semiteres (Lehm.) Mitt., J. Linn. Soc., Bot. 16: 188. 1877. ORIGINAL MATERIAL. SOUTH AFRICA: Cape Prov., (Cap. Bonae Spei), “la- tere orientali montis Teufelsberg”, Ecklon (G!). Plants translucent. Trigones minute to small, occasionally medium and straight- sided, 2.4-4.8 um, the ratio of trigone diameter to length of intervening thin wall ~ between trigones 1:2-5.6. Underleaf disc often 1.5-2x stem width; the lateral armature well developed: mostly laciniuform, at times lobuliform. Distribution: Pan-south temperate (Fic. 1). Chiloscyphus semiteres var. canaliculatus (G. L. & N.) Engel, comb. & stat. nov. Chiloscyphus canaliculatus G. L. & N., Syn. Hep. 710. 1847; Jungermannia canaliculata Hook. f. & Tayl., London J. Bot. 3: 563. 1844, hom. illeg., non J. canaliculata Nees, Hep. Javanicae 10. 1830; Lophocolea heterophylloides var. canaliculata (G. L. & N.) Carring. & Pears., Pap. & Proc. Roy. Soc. Tasmania 1887: 51. 1888; Lophocolea canaliculata (G. L. & N.) Steph., Bull. Herb. Boissier 6(9): 786. 1906 (= Spec. Hep. 3: 86). LECTOTYPE (nov.). NEW ZEALAND: “Falls of Waitangi”, Hooker s. n. (FH!—c. per. + Plants dull-opaque. Trigones normally large and bulging to knotlike, in shade forms medium and straight-sided to weakly bulging, (4.8—)6—7.2(—8.4) um, the ratio of trigone diameter to length of intervening thin wall between trigones vari- able: 1:(0.9—)1.2—2.6(—3.2). Underleaf disc usually 1-1.5 stem width; the lateral armature reduced: mostly dentiform to ciliiform or lacking, only sporadically small-lacinuform. DIsTRIBUTION: New Zealand, Tasmania and southeast Australia (Fic. 1). FURTHER REFINEMENTS IN CHILOSCYPHUS 1) SUBSPECIFIC DIFFERENTIATION OF CHILOSCYPHUS NOVAE-ZEELANDIAE This polymorphic species occurs in New Zealand, Tasmania and southeast Aus- tralia. It may be subdivided into three units, as follows: a) Chiloscyphus novae-zeelandiae var. novae-zeelandiae Jungermannia novae-zeelandiae Lehm. & Lindenb. in Lehmann, Nov. Min. Cogn. Stirp. Pug. 6: 33. 1834; Lophocolea novae-zeelandiae (Lehm. & Lindenb.) Nees in G. L. & N., Syn. Hep. 168. 1845; Chiloscyphus novae- zeelandiae (Lehm. & Lindenb.) Engel & Schust., Nova Hedwigia 39: 420. 1984 [1985]. ORIGINAL MATERIAL. NEW ZEALAND: South Is., Dusky Bay, s. coll. [Menzies] (BM!, G!, S![2], ex hb. Hooker—c. per.). b) per novae-zeelandiae var. meridionalis (Steph.) Engel, comb. & stat. OV. Lophocole meridionalis Steph., Bull. Herb. Boissier 6(10): 888. 1906 (= Spec. ~ ORIGINAL MATERIAL. NEW ZEALAND: “Insula meridionalis”, Waume (Canterbury), May 1901, Beckett 2737 (G!-c. ¢&). 110 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 c) Chiloscyphus novae-zeelandiae var. grandistipulus (Schiffn.) Engel, comb. & Stat. nov. Lophocolea grandistipula Schiffn. in Naumann, Forschungsr. Gazelle 4: 12, pl. De 32. 1890. ORIGINAL MATERIAL. NEW ZEALAND: Auckland, 8 Nov 1875, Naumann (FH!, sub L. stipularis, nom. hb.). 2) CHILOSCYPHUS PERPUSILLUS COMB. NOV. Chiloscyphus perpusillus (Hook. f. & Tayl.) Engel, comb. nov. Jungermannia perpusilla Hook. ft. & Tayl., London J. Bot. 3: 380. 1844; Lophocolea perpusilla (Hook. f. & Tayl.) G. L. & N., Syn. Hep. 163. 1845. OrIGINAL MATERIAL. NEw ZEALAND: Campbell Is., Hooker (BM![4], FH!). 3) THe TAXONOMIC POSITION OF CHILOSCYPHUS STRIATELLUS Mass. This species occurs in both the Valdivian and Magellanian zones of southern South America. It has elongate, longitudinally oriented lamellae on the stem, and I am unaware of any other member of Jungermanniales with such processes. It belongs in a section of its own, as follows: Chiloscyphus sect. Striatelli Engel, sect. nov. Folia caulina et amphigastria caulina profunde bilobata, superficiebus levibis. Caulis lamellis parallelis, longitudinalibus, latitudine 2—5 cellularibus ornata. Type: Chiloscyphus striatellus Mass. Leaves and underleaves deeply bifid, smooth. Stems on all sides with parallel, longitudinal lamellae 2—5 cells high. 4) THE TAXONOMIC POSITION OF CHILOSCYPHUS APOSINENSIS P1IPPO eee late aposinensis of China is similar to the pan-tropical C. muricatus (Lehm.) Engel & Schust. of sect. Microlophocolea Spruce in having spinose leaf surfaces and cor divided leaves and underleaves, but differs from that species in 1) the form of the leaf spines (the lumen of each protuberance is confluent with that of the subtending laminal cell and not separated from it by a cross wall [fig. 4 in Chang & Gao, 1984]); 2) the placement of leaf spines only on the dorsal surface of the leaf; and 3) the dense clusters of spines at the dorsal base of each leaf (fig. 4 in Chang & Gao, 1984). Chiloscyphus aposinensis belongs in an independent section, as follows: Chiloscyphus sect. Aposinenses Engel, sect. nov. Folia caulina pagina dorsali spinosa; spinae dispersae (cellula quaque spina unica ornata), sparsae ad basin ventralem, dense fasciculatae ad basin dorsalem, digitiformes, luminibus confluentibus cum luminibus laminae; folia caulina pagina levi instructa. Type. Lophocolea chinensis Gao & Chang, Bull. Bot. Res. 4: 87. 1984 = Chiloscyphus aposinensis Pippo (cf. Pippo, 1990). Dorsal leaf surface spines scattered, 1 each per cell, sparse at ventral base, in dense clusters at postical base, fingerlike, the lumina confluent with that of lamina; ventral leaf surface smooth. 1992 ENGEL: CHILOSCYPHUS it ACKNOWLEDGMENTS Iam grateful to Dr. Rolf Singer for assistance with the Latin diagnoses. LITERATURE CITED Engel, J. J. 1978. A taxonomic and phytogeographic study of Brunswick Peninsula (Strait of Magellan) Hepaticae and Anthocerotae. Fieldiana, Bot. 41: i-vin, . 1990. Falkland Islands (Islas Malvinas) Hepaticae and Anthocerotophyta: A taxonomic and Pyocceraere study. Fieldiana, Bot. N. S. 25: 1-vili, 1-209 Chang, Kuang-chu and Chien Gao. 1984. Plantae novae hepaticarum sinarum. Bull. Bot. Res. 4: 83-99 Grolle, R. 1959. Beitrag zur Kenntnis der afrikanischen Lophocoleen. Trans. Brit. Bryol. Soc. 3: 582- 598, meas = G. 1982. Lophocolea semiteres (Lehm.) Mitt. established in Argyll, Scotland. J. Bryol. 12 ie : ic eee eon en semueres (Lehm.) Mitt. and Telaranea murphyae sp. nov. established on Tresco. Trans. Brit. Bryol. > 775-779, f. 1. Pippo, oe 1990. ener tare of Chinese Hepaticae and Anthocerotae. J. Hattori Bot. Lab. 68: 1-192. 4S BY aide erm pens me Pe ae -—= * - - oe - we ' 2 - 7 7 7 a ier =| 5 : 7 =~ 8-5, = f= yeh wo _ - a ° > A 7 / 7 7 7 - ; - 7 > : : - ; - a _ » 7 - : _ a 7 ’ , - ar om fad ci 7 , 7 > = se, 2 -_-r-._ ~~ '6 oe - 7 : - sy 2) 7 7 , 7 7 7 — - : _ ‘4 : _ ' a : . ' a : | ; ssa be * ip ne Contr. Univ. Mich. Herb. 18:113-—130. 1992. A REVISION OF OCHROBRYUM (LEUCOBRYACEAE) Bruce Allen Missouri Botanical Garden P.O. Box 299 St. Louis, MO 63166-0299, U.S.A. DEDICATION Not so long ago I began to learn moss taxonomy with a copy of Howard Crum’s Mosses of the Great Lakes Forest, a gift from Ronald Pursell. My continued interest in mosses, to a great extent, is due to my beginning with that special book. I am, therefore, especially grateful to be able to contribute the following paper to this Festschrift honoring Howard Crum. INTRODUCTION Ochrobryum was established by Mitten (1869) for two leucobryaceous mosses with short, immersed capsules. The genus has a bizarre sporophytic complex that is unique in the Leucobryaceae: a short seta, a hemispherical, eperistomate capsule, and a short, symmetrical, rostrate operculum tightly enclosed by an exceedingly long (to three times the length of the operculum), narrowly subulate, mitrate calyptra. In form the capsule and the operculum are so similar that the former has been described as “an inverted operculum .. .” (Robinson 1990). The calyptra, which may or may not be fringed at base by long unicellular hairs, is generally longer than the combined operculum, capsule and seta. At capsule dehiscence the oper- culum commonly remains inside the calyptra. The resulting operculum/calyptra complex, although longer, has the same form as the deoperculate sporophyte. The leaves of Ochrobryum have the alternating leucocyst/chlorocyst arrange- ment typical for the Leucobryaceae. Unlike many members of the family the leaves at base are nearly always three layered: one dorsal leucocyst layer, a central chlorocyst layer and one ventral leucocyst layer. Occasionally, the unistratose pat- tern in the dorsal layer is disrupted by the subdivision of one or two dorsal leucocysts. But, this is a non-typical event and leaves that are three-layered at base represent an essential feature of Ochrobryum. Three-layered leaves are also found in a few Leucobryum species, and in the neotropics Ochrobryum is frequently misidentified as L. martianum (Hornsch.) Hampe, a three-layered leaf species that otherwise has little resemblance to Ochrobryum. In general, Ochrobryum leaves have ventral leucocysts positioned opposite the dorsal leucocysts. As a result the chlorocysts are four-sided. The chlorocysts proxi- mally are isocentric, i.e., as close to the dorsal surface as to the ventral surface. However, as noted by Cardot (1900), the chlorocysts distally are hypocentric, 1.e., closer to the dorsal surface than to the ventral surface. This hypocentric chlorocyst pattern is caused in two ways: all Ochrobryum species have enlarged upper ventral leucocysts, and in some species the upper ventral leucocysts are multilayered. The degree of chlorocyst hypocentricity is greater in these latter species than in the species with only enlarged leucocysts. Commonly, the species with multilayered 113 114 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 upper ventral leucocysts have fleshy or swollen leaf apices that are obtusely triangu- lar in cross-section. In past treatments of Ochrobryum leaf apex form has been used as a primary taxonomic character at the species level. This character, though useful, is astound- ingly variable within single collections of some species. Consequently, it is best regarded as a secondary taxonomic character. The greatest variability in apical leaf form is found in those species that have multilayered upper ventral leucocysts (O. gardneri (C. Mull.) Mitt. and O. Aurzianum Hampe in Besch.). In these species most leaves have fleshy, broadly acute to obtuse leaf apices. However, nearly all collections have some sharply acute leaves. Commonly, in these species the lower leaves are sharply to broadly acute while the upper leaves are fleshy and obtuse. For the species having only enlarged upper ventral leucocysts or a multi-layered pattern for only a single cell width (O. subulatum Hampe in Besch., O. sakalavum Card. & Par., and O. sessile Allen) the leaf apices are generally more sharply acute (obtuse in O. sakalavum) and not fleshy or swollen. Since all species of Ochro- bryum have at least a few leaves with some type of acute apex, the use of leaf apical form as a distinguishing feature must always be accompanied by subjective qualifi- ers such as “most” or “some”. An interesting and characteristic feature of the leaves in Ochrobryum is the appearance of the upper chlorocysts. Distally the chlorocysts are short, and form chains of rounded to quadrate cells. This feature is well-developed in all Ochro- bryum species and its presence is of critical importance in recognizing the genus when sporophytes are absent. Unfortunately, the feature is not unique to Ochro- bryum, Cardot (1900) noted it ina number of Leucobryum species and considered the Leucobryaceae to commonly have chlorocysts that are wider toward the apex than toward the base. In this regard it should be noted that Leucobryum in- curvifolium C. Mull. (one of the few Leucobryum species with erect capsules) has this chlorocyst feature as equally well-developed as any species of Ochrobryum. lobose, budlike propagula were not attributed to Ochrobryum until 1901 (Geheeb 1901). Robinson (1990) considered these propagula to be a feature that uniquely defined the genus within the Leucobryaceae. All species in the genus, except O. sakalavum which is known only from a scanty type collection, possess propagula of this type. Generally the propagula end in a single blunt cell, however some species have propagula that end in a sharp cell. But, even in these latter species a few propagula with blunt terminal cells can also be found. The globose propagula are attached at their proximal ends to a central, reddish-brown stalk of variable length by an clongated cell. The position at which the propagula occur on the plant is a character of primary taxonomic significance at the species level. The propagula may be borne on the upper dorsal surface of the leaves, in globose masses at the end of a specialized, apical stalk, or clustered in the leaf axils. Although Ochrobryum is largely defined by its sporophytic complex, the com- plex within the genus is nearly uniform. The only character in the complex that varies greatly is the degree and type of fringing of the calyptral base. In three species, O. subulatum (Asia, South America, Africa), O. kurzianum (Asia), and O. sakalavum (Madagascar) the base of the calyptra is densely beset with long, single-celled fringe cells. In the other species, O. gardneri (Central and South America, Africa) and O. sessile (Central and South America), the calyptra base may be weakly fringed or simply laciniate. In O. gardneri both fringed and laciniate calyptrae can be observed in the same collection. Long, single-celled, fringed calyptrae are widely and sporadically found within 1992 ALLEN: OCHROBRY UM 1s) the Dicranaceae, Leucobryaceae, Hookeriaceae and Daltoniaceae. The unsuitabil- ity of this character as an indicator of natural relationships outside of a generic context is obvious from the fact that the calyptral fringe cells of Ochrobryum are identical in morphology and position to those found scattered in many genera in these families. Even the variable nature of the calyptral fringing in Ochrobryum has a counterpart in some species of Campylopus where fringed and un-fringed calyptrae can be found within single collections. Robinson (1990) considered fringed calyptrae in the Leucobryaceae (Schistomitrium and Ochrobryum) to be a localized apomorphy because of its sporadic occurrence within the family. The erect capsules and mitrate calyptrae of Ochrobryum have led to the view that the genus is closely related to Schistomitrium (Mitten 1859, Andrews 1947). Further gametophytic support for this view is found in the hypocentric upper chlorocysts of Ochrobryum. As noted above, some species of Ochrobryum have leaf apices with enlarged, multilayered leucocysts. These leaf apices are essentially identical to the leaf apices found in species of Schistomitrium where this type of leat apex represents a generic feature. Robinson (1990) considered the generic subgroup of Schistomitrium, Cladopo- danthus and Holomitriopsis to be the only Leucobryaceae-subgroup marked by more than one synapomorphy—mitrate calyptrae, hypocentric chlorocysts, and erect capsules. Ochrobryum has all three of these character states and clearly belongs within the complex. Indeed it appears better placed there than Holomitriop- sis which has a cucullate calyptra. Robinson (1990) did not include Ochrobryum in the complex because of its unusual sporophyte and its globose vegetative propagu- la. As treated here these features are considered indicators of generic rather than subfamilial rank. Ochrobryum was revised by Bescherelle (1897). This revision added fourteen species and one variety to the genus, many of which were sporophytically unknown. Cardot (1900) considered Ochrobryum and Leucobryum gametophytically insepara- ble and so transferred or excluded Bescherelle’s species known only from gameto- phytic material. Cardot recognized nine species of Ochrobryum distributed in south- ern Asia and South America. Subsequently, the genus has been discovered in Central America and Africa. Wijk et al. (1964) attributed 20 taxa to the genus. Robinson (1965) added another species, but since he also transferred a Sri Lankan species of Ochrobryum to Leucobryum, 20 species are currently recognized in the genus. TAXONOMIC TREATMENT Ochrobryum Mitt., J. Linn. Soc., Bot 12: 107. 1869.—TyPE: Leucophanes gardneri C. Mill., Bot. Zeit. 2: 741. 1844. Plants small, 3-20 mm high, pale-green to whitish. Stems red to reddish-brown, erect, simple or sparingly branched, commonly with numerous arrested branch primordia along all sides; in cross-section cells thin- to firm-walled, epidermis undif- ferentiated, central strand absent. Rhizoids of two types: stout, smooth, sparsely and irregularly branched, dark red to red-brown ones distributed on all sides of the stems at base: and slender, smooth to lightly roughened, at times pinnately branched, pale reddish-brown ones on stems above and below leaf insertions and at times from the dorsal sides of the leaf apices. Pseudoparaphyllia absent. Paraphyllia 116 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 absent. Leaves erect-incurved, erect-spreading to flexuose, channeled; margins en- tire; lamina narrow, restricted to the lower half of the leaf: in cross-section at base with a median row of quadrate chlorocysts and single layers of dorsal and ventral leucocysts, at leaf apex chlorocysts hypocentric, and having one or several layers of enlarged ventral leucocysts; chlorocysts short in the upper region of the leaf, form- ing chains of rounded or quadrate cells; alar cells not differentiated. Axillary hairs delicate, 4—6 cells: basal cell short rectangular, hyaline to gray, upper cells long rectangular, hyaline, terminal cells rounded at the apex. Asexual reproduction by means of globose propagula clustered on short stalks: 1) on dorsal surface of leaf towards the apex, 2) on apex of specialized terminal branch or stem extension, 3) in leaf axils, and 4) on slender rhizoids at base of stems. Dioicous. Perigonia terminal on short branches at apex of stem. Perichaetia terminal on stem, archegonia 800 wm long, with slender, elongate neck 550 wm long. Setae short, smooth, vaginula well developed; capsules immersed to shortly exserted, hemispheric to obconic, wide- mouthed when dry; annulus non-revoluble, consisting of several rows of thick- walled irregularly shaped cells at operculum and capsule mouth and two rows of long rectangular, yellowish, thin-walled cells at site of separation; stomata absent; opercula long-rostrate; peristome absent. Calyptra long-rostrate, the rostrum more than twice the operculum length, mitrate (occasionally splitting irregularly and appearing cucullate), laciniate or ciliate-fringed at base. Etymology. A combination of the Greek ochros (pale, sallow, pale-yellow) and bryon (moss). Distribution, CENTRAL AMERICA: Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, and Panama. SouTH America: Colombia, Venezuela, Brazil, and Bo- livia. AFRICA: Guinea, Zaire, and Madagascar. Asia: India, Nepal, Burma, That- land, Cambodia, and Malaysia. na KEY TO THE SPECIES OF OCHROBRYUM — . Leaves linear to setaceous, gradually narrowed from base to apex; apices mostly sharply acute, usually a few apices broadly acute ............. |. Leaves lanceolate, narrowly ovate at base; apices broadly acute or obtuse, DOMME Pes 1) Same COUCCHION 455 .2xsecka ks cea) Hee ee ea kee oe 2. Propagula clustered in leaf axils; apical ventral leucocysts frequently two-layered in the middle for a single cell width .......... 4. O. sessile. 2. Propagula clustered at tip of propaguliferous branch or stem exten- sion which has reduced leaves; apical ventral leucocysts in a single PAIS Bei sr a ace ace Sahay 34. baw apo adc Ge ae 5. O. subulatum. . Leaves with apical ventral leucocysts two-layered in the middle for a single cell width; leaf lamina narrow at base, 2—4 cells wide ..... - 3. O. sakalavum. . Leaves with apical ventral leucocysts bi- to multi-layered in a broad band; leaf lamina broad at base, 4-10 cells wide 2.2.2... 4. Propagula on upper, dorsal leaf surface; basal leaf laminae with 4— 6 layers of long-rectangular cells; plants of the neotropics and Af- eo) ww TAG elie ts ss ay heh ae ee ead a AS oo . O. gardneri. 4. Propagula clustered in leaf axils; basal leaf laminae with 6-10 layers of long-rectangular cells; plants of Asia ................ 2, O. kurzianum. 1. Ochrobryum gardneri (C. Mull.) Mitt., J. Linn. Soc., Bot. 12: 108. 1869, “gard- nerianum” see nomenclatural note 1. Leucophanes gardneri C. Mill., Bot. 1992 ALLEN: OCHROBRYUM 117 Zeit. 2: 741. 1844. Schistomitrium gardnerianum (C. Mill.) Mitt., J. Proc. Linn. Soc., Bot. Suppl. 1: 26. 1859. Type. BRaziL. Gardner 94 (BM, H, NY isotypes). Leucophanes gardnerianum C. Mill., Syn. Musc. Frond. 1: 85. 1849, ortho. var. Leucobryum obtusifolium C. Mill., Bot. Zeit. 15: 577. 1857. Schistomitrium obtusifolium (C. Mull.) C. Mull., Ann. Sci. Nat. Bot., Sér. 5, 3: 339 : eee obtusifolium (C. Mull.) Mitt., J. Linn. Soc., Bot. 12: 108. 1869. Type. CoLtomsiA. Santa Martha, Minca, Funck & Schlim in coll. Linden 913 (BM, BR, H, NY isotypes), syn. nov. Ochrobryum phyllanthum Lindb. in Besch., J. Bot. (Morot) 11: 141. 1897, nom. nud. in synon. Based on BraziL, environs de Caldas, juillet 1854, Lindberg (BM). Ochrobryum paulense Broth. & Geh. in Geh., Rev. Bryol. 28: 11. 1901. ‘Type. BraziL. Sao Paulo: near Apiahy, Puiggari 56/ (H isotype), syn. nov. Ochrobryum normandii Card. & Par. in Par., Rev. Bryol. 29: 65. 1902. TyPEs. Guinea. Fouta Djallon: prés Donhol Neera, 29 mars; entre Dango & Bouna, 30 mars; Bourrougal Tope, 2 avril; pres riv. Kaligourou, 29 avril; all collected in 1901 by Normand (types not seen, see nomenclatural note 2), syn, NOV Oe onal dendeliae Broth. & Par. in Par., Rev. Bryol. 31: 117. 1904. Type. Guinea. Ad arbores secus amnem Dendeli, 4 mart 1904, Pobeguin (BM, H, NY isotypes), sya. nov Ochrobryum crumi Robins., Bryologist 68: 92. 1965. Types. Mexico. Jalisco: a Cumbre, 10 mi SW of Autlan, Crum 1247 (US holotype; F, NY isotypes). Paratypes. Mexico. Jalisco, Crum 438a (MICH, not seen), /334 (US), syn. nov. Nomenclatural notes. Wijk et al. (1964) incorrectly attributed the combination O. eee to Lindberg (1872) rather than Mitten (1869). This error can be attributed to Miiller’s (1849) use of an orthographic variant Shihan for a listing of his previously desc weed ee aie. gardneri. Miller never in- Be een to be a new species as is evidenced by his direct reference in the species citation to his earlier paper. Mitten (1869) unfortunately used Miller’s orthographic variant when transferring the species to Ochrobryum. In accordance with Article 75.3 (Greuter 1988) the epithet should be automatically corrected. In fact, it would seem that Lindberg (1872) was not effecting a transfer of the species but simply rectifying an orthographic error as can be seen from his citation of the species: O. gardneri (C. Mull.) Mitt. The types of O. normandii were not available for study. Nevertheless, it is synonymized with O. gardneri on the basis of Cardot’s statement (in Paris 1904) that the type of O. maclaudii is a mixture of that species (the fertile plants) and O. normandii (the sterile plants). My examination of this type material reveals a mixture of O. subulatum (fertile plants) and O. gardneri (sterile plants). Plants in compact tufts or cushions, greenish-white, becoming whitish-brown with age. Stems 3-13(-20) mm high. Leaves 2-5 mm long, 0.3-0.8 mm broad, closely and densely erect-patent or stiffly patent-spreading, narrowly ovate- lanceolate to lanceolate, shortly acuminate, bluntly acute or rounded obtuse, at times cucullate, frequently apiculate; margins erect; leaf lamina narrow, consisting of 2-4 rows of long-rectangular, variously porose cells at midleaf and 4—6 rows of rectangular, firm-walled cells at base; costa with a single dorsal leucocyst-layer and a single ventral leucocyst-layer throughout most of the leaf; apical ventral leucocysts multi-layered. Asexual reproduction by brown, multicellular, globose propagula 118 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 5 Ceeseoa: ROOD 5) (gees ) as m SS ORO AEA = o : f Ft gp Pipe ea ee XK ODOC ye 9 10} FIGS. 1-12. Ochrobryum gardnert. | & 2. Leaf apices. 3, 5,9, 10. Leaves. 4. Leaf cross-section at apex. 6-8. Globose propagula. 11. Basal leaf lamina. 12. Leaf cross-section at base. Scales in mm: 1,2 & 5 = 0.1; 3 = 0.2; 4 = 1.0. Figs. 1, 2, 4, 8, 11, 12 from Gardner 94; figs. 3, 5, 6, from Crum 1247; figs. 7, 9, 10 from Linden 913. 1992 ALLEN: OCHROBRYUM 119 clustered on short stalks on the upper dorsal leaf surface. Setae smooth, 1.0—1.5 mm long; capsules immersed to emergent, 0.5-1.0 mm long; opercula 1.5 mm long. Calyptrae 4-5 mm long, ciliate or laciniate at base. Spores spherical, smooth or lightly roughened, 12-18 um. Distribution. Central America (Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, and Panama); South America (Colombia, Brazil, and Bolivia); Africa (Guinea). Illustrations. Bescherelle (1897, figs. 1, 5); Cardot (1900, plate 8, fig. 37); Bartram (1949, fig. 31, A-D); Robinson (1965, figs. 15-21); Yano (1975, fig. 28, a— g); Fics. 1-12. Habitat. On rotting logs or tree trunks, and on bark at base of trees; 1OS—2200 m. Specimens examined. CENTRAL AMERICA. Mexico. Chiapas: above oR Sharp 4409 (US); Colima: Alzada, Orcutt 5359 (FH); Jalisco: La Cumbre, Crum 1247 (F, NY, US). GUATEMALA. Alta Verapaz: Coban, Turckheim (NY); Baja Verapaz: near Jicaro, Sharp sor (FH). Honpburas. Olancho: 14°54'N; 85°56" W, Olson 83-7b (MO). NICARAGUA. Jinotega: near Jinotega, Standley 9566 (F). Costa Rica. San José: Santa Maria de Dota, Standley & Valerio 43142 (FH, NY, US), Standley 41628 (FH, H, US), Standley 41657 (FH, US). PANAMA. Chiriqui: 8°35’N, 81°45’W, Allen 5218 (MO), Santa Clara, Mori & Bolten 7169 (MO), 2 mi S of Boquete, Crosby 3961 (MO, PMA), Cerro Pando, 8°52'N 82°45'W, sate S al, 811 (MO), 2 mi E of Boquete, Nee 10804 (MO, NY, PMA, US RICA. Cotomsia. Magdalena: Santa Marta, Linden 9/3 (BM). Braziv. Bahia: 11°39'S, 41°07'W, ne 8038 (MO); Golds: Serra Geral do Parana, Irwin et al. 31982 p.p. (FH, H, NY), Irwin et al. 33185 (FH, H, NY), Irwin 33208a (NY); Minas Gerais: Caldas, Mosén (H), Lagoa Santa, Warming (BM); Sao Paulo: Faxima, ee 561 (H). Botivia. La Paz: Santa Anna, Apolo ae Tle 1861 (BM, FH, H, NY); San ay Velasco Province, Rio Blanco, Herzog (H), Prov. Velasco, 15°58’S, 61°05’ W, Lewis 85- ae AFRIC Guinea. Féfiné River, ae p.p. (BM); Lambeira, Pobequin (BM, NY); Dombiagni, Pobeguin (H, NY); Dendeli, Pobeguin (H); Diendieu, Pobeguin (NY). This species and O. kurzianum form a close pair marked by the presence of bluntly acute to obtuse leaves that are distinctly swollen or thickened at the extreme apex due to their enlarged and multi-layered upper ventral leucocysts. Ochrobryum kurzianum differs in having propagula clustered in the leaf axils and by its better developed leaf laminae (6—10 layers of long rectangular cells). The two species are geographically distinct: O. gardneri Central America, South America and Africa; O. kurzianum Asia. The leaf apices in O. gardneri are remarkably variable, even within single collections. At times nearly all leaves are obtuse and apiculate, conversely some collections have mostly broadly acute leaves. All collections have leaves with thick- ened apices and at least some obtuse leaves. Globose propagula borne on the upper dorsal leaf surface are a constant, diagnostic feature of O. gardneri. This species is similar to O. sessile which is sympatric with it in Central America and northern South America. Ochrobryum sessile has moderately stout leaves with leaf apices that approach those of some plants of O. gardneri. In addition, the leaf apices frequently have ventral leucocysts in a two-layered pattern for a single cell width in the middle of the leaf. Ochrobryum sessile differs not only by its lack of an extensive apical region of multilayered ventral leucocysts, but also by having propagula clustered in the leaf axils and smoothly tapered, narrower, more sharply acute leaves. Ochrobryum subulatum frequently occurs mixed with O. gardneri. It is a more slender plant with narrowly acute leaves, a constant uni-layered ventral leucocyst 120 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 pattern, and has globose leaf propagula on specialized, terminal branches or stem extensions. Vegetatively this species resembles Leucobryum polakowskyi (C. Mull.) Card. or L. incurvifolium. But, these two species proximally have multi-layered dorsal and ventral leucocysts, quadrate basal laminal cells, and they reproduce asexually by means of terminal clusters of reduced leaves. Holomitriopsis laevifolia (Broth.) H. Robins., a Venezuelan species with ob- tuse, apiculate leaves and a single dorsal and ventral leucocyst-pattern, may also approximate this species. Holomitriopsis laevifolia is a more robust plant, lacks rounded to quadrate apical chlorocysts and leaf propagula, has a cucullate calyptra, and a peristomate capsule 2. Ochrobryum kurzianum Hampe i7 Besch., J. Bot. (Morot) 11: 145. 1897. Types. Burma. Moulmein, Stoliczka [Kurz] 4434 (BM lectotype; BM, BR, H isolectotypes), Stoliczka | Kurz] 4438 (BM, H syntypes); Pegu- Yomah, Kurz 2890 (BM, H, NY syntypes); Martaban, Kurz 2842 (BM, H, NY syntypes). Ochrobryum kurzianum Hampe in F. Mason, Burma 2: 49. 1883, nom. nud. Ochrobryum nepalense Besch., J. Bot. (Morot) 11: 145. 1897. Type. NEPAL. Wallich 30 (BM holotype; BM isotypes), sya. nov. Ochrobryum parishti Besch., J. Bot. (Morot) 11: 142. 1897. Type. Burma. Moulmein, Parish 2 (BM holotype; NY isotype), svi. nov. Nomenclatural notes. . The above three species have equal ee The epithet Aurzianum is used for the combined taxon since it is the name in common use in . The lectotype of O. kurzianum (the ee wpe material in the Bescherelle herbarium) was cited as “Stoliczka, S. Kurz 44347 (Bescherelle 1897), but the specimen in Hampe’s herbarium was evidently collected solely by Stohezka. Hampe’s specimen has one of nee printed labels on which Stoliczka’s name has been hand-written and Kurz’s name crossed 0 The type of O. parishit is Parish 2, but specimens at BM and NY wit ee the number oe may also be type material. The error may have occurred when Mitten @ 856) listed Parish 2 as “22. Schistomi- (rim garderianum”, in the same paper Parish 22 is a hepatic. The holotype has no collection number. In the Mitten herbarium there is one specimen of Parish 2. All material listed as No. 22, Parish (BM, NY) is considered possible type material. i) Ww Plants in compact tufts or cushions, greenish-white, becoming whitish-brown with age. Stems 5-15 mm high. Leaves 2-4 mm long, 0.5—0.8 mm broad, closely and densely erect-patent or stiffly patent-spreading, narrowly ovate-lanceolate to lanceolate, shortly acuminate, bluntly acute or rounded obtuse; swollen or fleshy at the apex; margins erect; leaf lamina narrow, consisting of 6-8 rows of long- rectangular, variously porose cells at midleaf and 6—10 rows of elongate-rectangular, firm-walled cells at base; costa with a single dorsal leucocyst-layer and a single ventral leucocyst-layer throughout most of the leaf; ventral leucocysts multi-layered at the extreme apex. Asexual reproduction by brown, multicellular, globose propagula on very short stalks clustered in the leat axils. Setae smooth, 1.0—1.7 mm long; capsules immersed to emergent, 0.5—0.75 mm long; opercula 1.0 mm long. Calyptrae 4-5 mm long, strongly ciliate at base. Spores spherical, lightly rough- ened, 10-17 um. Distribution. Asia (India, Nepal, Burma, Thailand, and Cambodia). Mlustrations. Bescherelle (1897, figs. 2, 3, 4); Cardot (1900, plate 8, figs. 39, 40); Gangulee (1969, fig. 193); Eddy (1990, fig. 185); Fics. 13-26. Habitat. On rotting logs or tree trunks, and soil over rock; 50-1600 m. 1992 ALLEN: OCHROBRYUM 121 O ay 0) OS A 7 OSS SCS IC SESE Ss FIGS. 13-26. Ochrobryum kurzianum. 13. Leaf cross-section at apex. 14 & 15. Leaf apices. 16. Calyptra. 17. Operculum. 18. Capsule, seta and vaginula. 19 & 20. Leaves. 21. Basal leaf lamina. 22. Calyptral hairs. 23 & 24. Globose propagula. 25. Spores. 26. Leaf cross-section at base. Scales in mm: 1 &2=0.1;3&4= 1.0;5& 6 = 0.2. Figs. 13 & 26 from Touw 9450; figs. 14-21 from Kurz 2890; figs, 22- 25 from Beddome 804. 122 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 Specimens examined. INDIA. Assam: Garo Hills, Marten 5702 (H, NY); Orissa: Eastern Ghats, Jeypore Talug, Walker 529, 579 (BM), Russelconda Hills, 19°50'N; 84°35'E, Beddome 804 (BM, NY). Nepac. Wallich 30 (BM). Burma. Pegu: Pegu-Yomah, Kurz 2890 (BM, H, NY): Tenasserim: Moulmein, Parish 2 (BM, NY), Parish 112 (BM, H), Stoliczka 4434, 4438 (BM), Martaban, Kurz 2842 (BM, H, Y). THAILAND. Chanthaburi: Kao Sabah, Kerr /08 (BM); Kanchanaburi: Ta Kanun, Kerr 145 (BM); Krabi: Krabi, Kerr 503 (BM); Payap: 98°35'E, 18°30'N, Touw 9450 (BM, MO, NY); Trat: Koh ce long Munsé, Bea 20 (BM, H); Adang: Salut, Kerr 245 (BM). Camsopia. Kandal: near To Bong Poilane (BM, H, NY). In one paper, Bescherelle (1897) described three Ochrobryum species—treated here as one species—based on variations in plant size, leaf apex shape and the number of cell rows in the basal lamina. These distinguishing features are extremely variable and not constantly expressed even within single collections. A nearly identi- cal range of variation is exhibited by O. gardneri for the same characters. Ochrobryum kurzianum and O. gardneri were treated by Mitten (1859) as a single species. Bescherelle (1897) separated them on the basis of their allopatric distributions, and the presence in O. gardneri of shorter, flatter, narrower, acuminate-acute leaves that had a narrower basal leaf lamina (4 vs. 10 rows of cells). Most of these distinctions are not valid, but these two species do differ in three ways. The basal leaf lamina of O. gardneri is generally narrower, its propagula are borne on the upper dorsal surface of the leaves rather than in the leaf axils, and its calyptrae tend to be laciniate rather than strongly ciliate at base. The significance of their different geographic distributions is minimized by the occurrence of O. subulatum in Asia and South America. Globose propagula clustered in the leaf axils are also a feature of O. sessile, but the two species do not appear closely related. Ochrobryum sessile has linear leaves, slenderly acute leaf apices and apical ventral leucocysts that are uni- or bi-layered for only a single cell width. a CO oe sakalavum Card. & Par. in Card., Bull. Herb. Boissier, Sér. 2, 4: YPE. MADAGASCAR. Territoire Sakalave: cercle de Maintirano, secteur de Rakobe, poste d@ampoza, prés du village de Tahutofoly, sur bois pourri; récolté par un tirailleur sakalave (PC). Ochrobryum sakalavum Card. & Par. in Par., Rev. Bryol. 29. 1902, nom. nud. Plants in compact tufts, yellowish-white. Stems 5 mm high. Leaves 2-4 mm long, 0.3—0.4 mm broad, closely erect-patent or subflexuose, narrowly ovate-lanceolate, shortly acuminate, obtuse to subacute and apiculate, not particularly swollen or fleshy at the apex; margins erect; leaf lamina narrow, consisting of 2—4 rows of elongate-rectangular cells at base; costa with a single dorsal leucocyst-layer and a single ventral leucocyst-layer throughout most of the leaf; apical ventral leucocysts in two layers for single cell width at the middle of the leaf. Asexual propagula not seen. Setae smooth, 1.2 mm long (with vaginula); capsules immersed, 0.3—0.5 mm long: operculum and calyptra not seen. [Cardot ages eae convex-conical to | mm, calyptra long subulate, fimbriate at base, 2.5-2.75 mm long.] Distribution. Madagascar. Illustrations. Renauld and Cardot (1915, pl. 155, figs. 2a—2s); Fics. 27-34. Habitat. On rotting wood. Specimen examined. MADAGASCAR. Territoire Sakalave: cercle de Maintirano, without collector or number (PC). 1992 ALLEN: OCHROBRYUM 13 FIGS. 27-34. Ochrobryum sakalavum. 27. Capsule, seta and vaginula. 28. Leaf apex. 29 & 30. Leaf cross section at apex. 31. Basal leaf lamina. 32 & 33. Leaves. 34. Leaf cross-section at base. Scales inmm: 1 & 4 = 1.0; 2 = 0.1; 3 = 0.2. All figures from the type. 124 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 Ochrobryum sakalavum has narrowly ovate-lanceolate leaves that are com- monly obtuse and apiculate above, narrow leaf laminae, and immersed capsules. This species occupies an intermediate position in the genus. The other four species can be divided into two groups: gardneri—kurzianum and subulatum —sessile. Its ovate-lanceolate, obtuse, occasionally apiculate leaves are found in the gardneri- Aurzianum group, but unlike that group the leaf apices are not distinctly fleshy and the apical ventral leucocysts are bi-layered only for a single cell width at the very tip of the leaves. These deviate features are found in the swbulatum—sessile group, but that group has longer, linear, sharply acute leaves. It differs from all species of Ochrobryum by its relatively narrow leaf laminae that are composed of broader, less attenuate cells. Globose propagula were neither reported by Cardot (1904) nor Renauld and Cardot (1915), nor observed in the only collection available for study. This collection had only a few plants and it was not possible to adequately deter- mine if the species produces propagula. 4. Ochrobryum sessile Allen, sp. nov. Ab O. gardnerit (C. Mull.) Mitt. cellulis hyalinis distalibus ventralibus saepis- sime unistratis gemmisque sessilibus in axillis foliorium superiorum differt. Plants in compact tufts or cushions, dark greenish-white, becoming yellowish- green with age. Stems 3-10 mm high. Leaves 4-5 mm long, 0.2-0.5 mm broad, erect-patent to falcate, narrowly lanceolate, acuminate, ending in a single sharp, hyaline cell, flattened above and commonly spirally twisted when dry; margins erect to incurved below; leaf laminae narrow, consisting of 2-3 rows of long-rectangular, variously porose cells at midleaf and 4—5 rows of clongate-rectangular cells at base; costa with a single dorsal leucocyst-layer and a single ventral leucocyst-layer throughout most of the leaf, apical ventral leucocysts occasionally two-layered for a single cell width. Asexual reproduction by hyaline or brown, multicellular, globose propagula on short or long stalks usually clustered in the axils of the upper leaves, at times also in the lower leaf axils or on slender rhizoids at base of stem. Setae smooth, 1.0-1.5 mm long; capsules immersed to emergent, 0.7-1.0 mm long; opercula 1.5—2.0 mm long. nee 3-6 mm long, weakly ciliate at base. Spores spherical, lightly papillose, 15-18 Type. CoLomBiA. Meta: in Remiita forest west of Cordillera Oriental, Sreere 7054 (MO holotype; H, NY isotypes). Distribution. Central and South America (Costa Rica, Panama, Colombia, and Venezuela). Mlustrations. Fics. 35-46. Habitat. On rotting logs or bark at base of trees; 700-2000 m. —_ ules examined, CENTRAL AMERICA. Costa Rica. San José: El General, Skutch 2209 (FH, MO, US); Puntarenas: 5 ie Sof S. Vito, © hile 2658 (MO). PANAMA. Darién: Pirre Massif, W of Cana, Allen 8849, S897 (M MA); Panama: Cerro Azul, Castillo 112C (MO. SOUTH AMERIC ie 2 OLOMBIA. Meta: Remija west of Villavicencio, Steere 7654 (H, MO, NY). VENEZUELA. Aragua: Parque Nacional Henri Pittier, Guzman Ma-761a (US); Carabobo: between Valen- cia and Maracay, Alston 6/52 (BM, FH, US). This species resembles O. subulatum in leaf shape, but is a larger plant that grades into the more narrow-leafed expressions of O. gardneri. It differs from QO. subulatum not only by its larger, more frequently spirally twisted leaves but also by its more strongly papillose spores, and propagula clustered in the leaf axils—a 1992 ALLEN: OCHROBRYUM 125 OCH \es OF BGO 38 CORD: aA BA = —— == = Sole —— _———— ae a Gn a a FIGS. 35-46. Ochrobryum sessile. 35. Operculum. 36. Calyptra. 37. Leaf apex. 38. Leaf cross- section at apex. 39. Spores. 40. Capsule, seta and vaginula. 41 & 42. Globose propagula. 43 & 44. Leaves. 45. Basal leaf lamina. 46. Leaf cross-section at base. Scales in mm: | & 4 = 1.0; 2,3 & 6 = 0.155 = 0.2. All figures from the type. 126 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 feature it shares with O. kurzianum. In O. gardneri and O. kurzianum the leaf tips are thickened, fleshy and have an extensive multi-layered ventral leucocyst pattern. In addition, O. gardneri has propagula borne on the upper dorsal surface of the leaves. The leaf tips in O. sessile are identical to those of O. sakalavum in having the apical ventral leucocysts two-layered for a single cell width at the middle of the leaf. Ochrobryum sakalavum has obtuse leat tips. 5. Ochrobryum subulatum Hampe in Besch., J. Bot. (Morot) 11: 150. 1897. Type. Burma. Plumadoe, Kurz 2833 (BM holotype; H, NY isotypes). Ochrobryum subulatum Hampe in F. Mason, Burma 2: 49. 1883, nom. nud. Ochrobryum stenophyllum Besch., J. Bot. (Morot) 11: 150. 1897. Type. Bra- zIL. Spruce 73 p.p. (BM holotype; BR isotype; Spruce 73b BM, H isotypes), see Homenclaniral note 2, syn. nov. pane maclaudiu Card. & Par. in Par., Rev. Bryol. 30: 101. 1903. Type. Guinea. Ad ligna semiputrida secus amnem Féfiné, 20 mart. 1903, Maclaud (H, NY isotypes), see nomenclatural note 3, syn. nov. Ochrobryum normandi var. fragilifolium Dix. & Thér., Rev. Bryol. Lichénol. 12: 69. 1942. Type. ZarRE. Lulua, octob. 1923, Overlaet (BM isotype), syn. nov. Ochrobryum microphyllum Card. ex Herz., Hedwigia 61: 295. 1919, nom. illeg., see nomenclatural note 4. Based on Mataysia. Malakka obes “Batang Padang-Tal”, Stresemann 56 (H). Non O. microphyllum (Besch.) ard. Ochrobryum herzogianum Mohamed & Tan, Bryologist 91: 29. 1988, nom. nov. for O. microphyllum Card. ex Herz., nom. illeg. Nomenclatural no . Ochrobryum Tia and O. pe have equal priority. The epithet swbulanum was chosen for the combined species because its type material is better. The type of O. stenophytlum was ee from a specimen in PC (Spruce 73) issued by Spruce in his Musci Amazonici et Andini exsiccatl. Previously, Spruce 73 in BM had been recognized as a mixed collection, divided, and given the numbers Spruce 73 and Spruce 73b. Spruce 73b (BM) is identical to the holotype of O. Cian. Spruce 73 (BM) is identical to the holotype of O. gardneri var. microphyllum. Cardot and Paris ‘ig to cite type material for Ochrobryum maclaudii. The type was later desig- nated by Paris (19 . Oc Roker ceive te Card. ex Herz. was never validly described, furthermore neither Herzog (1919) nor Dixon (1926) intented to describe a new species for this taxon. Likewise O. herzogianum is also invalid. The specimen on which the name is based is O. subulatum. bo wW a Plants slender, in compact or loose tufts. Stems 2—4(—7) mm long. Leaves 2.0- 4.5 mm long, erect-patent to secund, linear, evenly tapered from the base to a setaceous or narrowly acuminate apex, not apiculate, but frequently ending in a single, sharp, clear cell; margins erect below; leaf lamina restricted to 5—6 rows of linear or elongate-rectangular cells at basal margins; costa in cross-section with a single ventral leucocyst-layer and a single dorsal leucocyst-layer throughout the leaf. Vegetative reproduction by globose, multicellular propagula arising on short stalks and occurring in terminal clusters on specialized branches or stem extensions; propaguliferous branches to 2 mm long, red-brown, with short, ovate leaves. Setae 1.0 mm long; capsules hemispherical to cupulate, 0.5 mm long; opercula 1.5 mm long. Calyptrae 3-4 mm long, ciliate at base. Spores smooth to lightly roughened, 17-20 wm 1992 ALLEN: OCHROBRYUM 127, | EES 47-48-49 50-57 51-54-55-56-58 52-53 FIGS. 47-58. Ochrobryum subulatum. 47. Operculum. 48. Capsule, seta and vaginula. 49. Calyptra. 50. Leaf apex. 51. Leaf cross-section at apex. 52 & 53. Leaves. 54 & 55. Globose propagula. 56. Spores. 57. Basal leaf lamina. 58. Leaf cross-section at base. Scales in mm: 1 & 4 = 1.0;2&3= 0.1. Figs. 47-49, 51, 58 from Touw 10290; figs. SO, 52, 53, 57 from Kurz 2883; figs. 54-56 from Fife er al. 4247, 128 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 Distribution. Asia (Burma, Thailand, and Malaysia); South America (Venezu- ela, Brazil, and Bolivia); Africa (Guinea and Zaire). Illustrations. Bescherelle (1897, fig. 7); Cardot (1900, pl. 8, figs. 38, 42); Fics. 47-58. Habitat. On rotting, decorticated logs and on bark at base of trees; LO0—1650 m. Specimens examined. ASIA. Burma. Plumadoe, Kurz 2833 (BM, H, NY). THAILaANnb. 18°35'N, 98°30'E, Touw 10290 (BM, MO, NY). Macaysia. Batang Padang-Tal, Stresemann 86 (H). SOUTH AMERICA. VENEZUELA. Bolivar: 4°25'N, 61°20'W, Steyermark & Dunsterville 112253 (MO); Mérida: 7°41'N, 71°28'W, Liesner & Gonzales 9371 (MO). Brazit. Acre: 33 km S of Rio Branco along road to Brasiléia, Reese & McPherson 13263 (MO, NY); Alagoas: 33 km N of Macei6, Boom & Mori 1024 (NY); Goias: 3 km S of Sao Joao de Alianga, /rwin et al. 31982 p.p. (FH, H, NY); Mato Grosso: Veu de Noiva, Chapada dos Guimaraes, Prance et al. 19015, 19016, 19149, 19398 (all NY); Minas Gerais: near Franca, Wacker 1171, 1172 (H):; Rondonia: 11°S, 64°W, Reese 13488 (MO, NY, US), 15 km E of Santa Barbara, Prance & Ramos 6910 (FH, NY), 9° 10'S, 63°07'W, Fife etal. 4247 (MO), 128 km SW of Ariquaes, McFarland 138 (MO, NY). Bouivia. Beni: vicinity of Guayaramerin, Reese 12910, 13130 (MO, NY). AFRICA, Guinea. Koulaire, Maclaud (H), Casamonce-Boubounda, Maclaud (H), along Feéfiné River, Maclaud (H, NY). Zaire. Lulua, Overlaet (BM). This is the smallest species in the genus and the only one with propagula in terminal clusters on specialized branches or stem extensions. The propaguliferous branches can be recognized, even without propagula, by their terminal position, striking red-brown color and short, ovate leaves. Ochrobryum subulatum and O. sessile form a pair united by their narrow leaves that smoothly taper from base to a setaceous or long-acuminate apex. Unlike QO. Aurzianum, O. gardneri, and O. sakalavum the apex is not obtuse or apiculate. However, the leaves frequently end in a single, sharp, clear cell, a condition also seen in O. sessile and some leaves of O. gardneri. Ochrobryum kurzianum and O. gardneri also have thickened or swollen leaf apices. Ochrobryum sessile is a larger plant with falcate leaves that are distinctly twisted when dry. It has propagula clustered in the leaf axils and calyptrae that are weakly ciliate at base. In South America and Africa O. subulatum often occurs mixed with O. gard- nert. Ochrobryum gardneri has larger, stouter leaves, some obtuse, swollen leaf apices, and leaf propagula on the upper dorsal surface of the leaves. There is a small, sterile Leucobryum-like moss in Brazil (Buck 2336 NY, 2749 NY, 3081 MO, NY) that closely mimics O. subulatum in size and aspect. This taxon also has leaves with a single layer of dorsal and ventral leucocysts. It differs from O. subulatum in its virtual absence of a basal leaf lamina, lack of globose propagula and the presence of rectangular chlorocysts in the upper parts of its leaves. TYPES NOT SEEN Ochrobryum mittenii C. Mull. ex Besch., J. Bot. (Morot) 11: 148. 1897. Type. CEYLON. 1862, Nietner (in Herb. C. Miull.). Ochrobryum nietneri C. Mull. ex Besch., J. Bot. (Morot) 11: 151. 1897. Type. CEYLON. 1862, Nietner (in Herb. C. Miull.). Cardot (1900) provisionally excluded these species from Ochrobryum since their sporophytes were unknown. From their descriptions it is not possible to deter- mine if they belong in Ochrobryum. 1992 ALLEN: OCHROBRYUM 129 EXCLUDED SPECIES Leucobryum incurvifolium C. Mull. Ochrobryum obtusissimum Dix., J. Bot. 76: 225. 1938. Types. TANZANIA. Mufindi, Balbo 96 (BM holotype); NyasaALANnp [MaLawi]. Bandowe, Chinteche, Little 2539 (BM paratype), Nchese Mt., Davy 1312 (BM paratype), sym. nov. The sporophytes of O. obtusissimum are unknown, but the presence of multi- layered leucocysts in the leaf bases, quadrate lower laminal cells and masses of brood leaves at the stem apices are features indicative of Leucobryum rather than Ochrobryum. In these gametophytic features, as well as its leaf shape, cucullate, fleshy leaf apices, and rounded apical chlorocysts it is a perfect match for Leucobryum incurvifolium. The inclusion of this taxon in L. incurvifolium gives that species a bi-centric distribution: Central America (Mexico to Honduras, see Robinson 1965, and Nicaragua [Standley 9561, F, FH])—Africa (Angola [Goss- weiler 11448, BM], Malawi, Tanzania). Leucobryum subobtusifolium (Broth.) Allen, comb. nov. Ochrobryum subobtusifolium Broth., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 26 Afd. 3(7): 10. 1900. Type. Brazi_. Mato Grosso: Serra da Chapada, Lindman 402 (H holotype; H isotype Ochrobryum parvulum Besch., J. Bot. (Morot) iL: 147. 1897. Non Leuco- bryum parvulum Card. Type. BRaziL. Ad flumen Amazone, Spruce 73 p.p. (BR isotype), syn. nov. Ochrobryum gardneri var. microphyllum Besch., J. Bot. (Morot) 11: 142. 1897. Ochrobryum microphyllum (Besch.) Card., Mém. Soc. Sci. Nat. Cher- bourg 32: 28. 1900. Type. BraziL. Sur les troncs d’arbres le long du fleuve Amazone, Spruce 73 p.p. (BM holotype; BR isotype) syn. nov. The oldest name for this taxon in Leucobryum is preoccupied by L. parvulum Card. The epithets subobtusifolium and microphyllum have the same year of prior- ity (both distributed as off-prints at earlier dates than the journals in which they appeared). Ochrobryum subobtusifolium appeared sometime between 14 March and 31 December 1900; O. microphyllum sometime between 31 May and 31 Decem- ber 1900. I have not determined the exact dates of issue for these off-prints but have chosen the epithet subobtusifolium as the name for the combined species because of its potentially earlier date. The sporophytes of this species are unknown and so its generic position 1s uncertain. Unlike Ochrobryum it reproduces asexually by means of apical clusters of brood leaves rather than globose propagula. It is similar to Leucobryum in- curvifolium and like that species and Ochrobryum species has short, rounded apical leaf chlorocysts. It differs from L. incurvifolium in having its dorsal and ventral leucocysts in a single layer throughout the leaf. Leucobryum ceylanicum (Besch.) Card., Mém. Soc. Sci. Nat. Cherbourg 32: 15. 1900. Ochrobryum ceylanicum Besch., J. Bot. (Morot) 11: 148. 1897. Type. CEYLON. Thwaites 81 (BM holotype; H, NY isotypes). The sporophytes of this species are unknown. Cardot (1900) transferred it to Leucobryum on the basis of its multilayered, basal leucocysts. 130 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 Leucobryum humillimum Card., Mém. Soc. Sci. Nat. Cherbourg 32: 15. 1900, nom. nov. pro Ochrobryum wightii Besch., J. Bot. (Morot) 11: 149. 1897, non Leucobryum wightti Mitt. Type. INDIA. Madras, Wight (BM holotype; BM, NY isotypes). eas: propaguliferum Dix., Notes Roy. Bot. Gard. Edinburgh 19: 281. . Leucobryum propaguliferum (Dix.) Robins., Bryologist 68: 91. 1965. oe CEYLON [SRI Lanka]. Hunasgiriya, Alston 1895 (BM holotype). PARATYPES. CEYLON [SRI LANKA]. Hatton near Hambantotuoya, Alston 2607 (BM); Wight (E, not seen), sy. nov. Dixon unwittingly suggested the above synonymy by his inclusion of an isotype of O. wightti in the protologue of Ochrobryum propaguliferum. Robinson (1965) placed O. propaguliferum in Leucobryum on the basis of its terminal clusters of deciduous leaves. As in Ochrobryum this species has a single layer of dorsal and ventral leucocysts throughout the leaf. ACKNOWLEDGMENTS thank BM, BR, H, F, FH. NY, PC and US for the loan of material. | am especially grateful to Gordon McPherson for help with French translation. LITERATURE CITED Andrews, A. L. 1947. Taxonomic notes VI. The pecan: nie 50: 319-326. Bartram, E. B. 1949. Mosses of Guatemala. Fieldiana, Bot. Bescherelle, E. 1897. Révision du genre Ochrobryum. J. ee aos aa —153 Cardot, J. 1900— ee Fee Recherches anatomiques sur les ee Mém. Soc. Sci. Nat. Cherbour 1904. 1 ie “Lescobrocts de pis as et des autres Iles Austro-Africaines de L’Océan fadien: Bull. . Boissier, Sér. 2, 4: 97-118 Dixon, H. N. Se) - ae st of the mosses of the Malay See Gard. Bull. Straits Settlem. 4: 1-46. Eddy, - [1990]. A Handbook of Malesian Mosses. Vol. 2. Leucobryaceae to Buxbaumiaceae. cue History Museum Publications, Londo Gangulee, H. C. 1969. Mosses of a India and Adjacent Regions. Fascicle 1. Sphagnidae, An- dreaidae & | ae ee Calcutta Geheeb, A. 1901. Révision des mousses Shack en Brésil dans la province de San-Paulo par M. Juan J. 77-188 je) anna pendant les années 18 . Rev. Bryol. 28: 9-11 Greuter, W. (ed.). 1988. International code of botanical po enciarars: Regnum Veg. 118: 1—xiv, 1-328 Herzog, T. 1919. Die Laubmoose der zweiten Freiburger Molukkenexpedition. Hedwigia 61: 286- 299, Lindberg, S. O. ee eee ad floram cryptogamam Asiae Boreali-Orientalis. Acta Soc. Sci. Fenn. 10: 223- Mitten, W. 1856. A ae of some mosses and Hepaticae, collected by the Rev. Charles Parish at Moulmein, and communicated to Sir W. J. Hooker. Hooker's J. Bot. Kew Gard. Misc. 8: 353-357. . 1859. Musci Indiae eee an enumeration of the mosses of the East Indies. J. Proc. Linn. Soe. Bot. Suppl. | 69. Musci Aree aan J. Linn. Soc., Bot. 12: 1-659. Miiller, . 1849. Synopsis Muscorum opoian, 7. Foerstner, Berlin Paris, [E. G.]. 1904. Ochrobryum maclaudii et normandi. Rev. Bryol. 31: 14. Renauld, F. and J. Cardot. 1915. Mousses. Jn: A. & G. Grandidier (eds.), Histoire, physique, naturelle et politique Madagascar 39: 1-5 Robinson, H. 1965. Notes on Leucobryaceae in Central America. ee 68: 89-93. aX funciona evolution of the Leucobryaceae. Trop. Bryol. 2: 223-237. Wijk, R. van der, W. wi argadant and P. A, cece 1964: Index reer Vol. HI (A/ypnum-O). Regnum Veg. 33: 1-529. Yano, O. 1975. Leuc ee aceae (Musci) do Estado de Sao Paulo. M.S. Dissertation, Area de Productos Naturais, Escola Paulista de Medicina, Sao Paulo, Brazil. Contr. Univ. Mich. Herb. 18:131-—140. 1992. REPRODUCTIVE BIOLOGY OF THE RARE “COPPER MOSS” MIELICHHOFERIA MIELICHHOFERIANA A. Jonathan Shaw, N. J. Niguidula and T. M. Wilson Department of Biology Ithaca College Ithaca, NY 14850, U.S.A. INTRODUCTION The “copper mosses” are unique among land plants. These plants, which actually include both mosses and liverworts, have extremely broad geographic ranges that typically span several continents. Broad intercontinental distributions are not, of course, especially remarkable for bryophytes, as most species of mosses and liver- worts have wide ranges compared to seed plants. It is the highly restricted and disjunct occurrences with each site hundreds or even thousands of miles from the nearest station, a general rarity of sexual reproduction over much of their ranges, and at least some association with deposits of copper and/or other metals, that sets the copper mosses apart from other plants. The fact that several species of copper moss often co-occur at precisely the same widely separated stations makes their geographic distributions and ecological characteristics unique and even more bizarre. There are different kinds of rare plant species, each with different biological causes. Rarity resulting from habitat specificity is one of three basic types recog- nized by Rabinowitz (1981) and others interested in explaining the biological basis of rarity. These species are thought to be sporadic in occurrence primarily because their habitats are discontinuous and uncommon. Rarity resulting from edaphic specificity has been most thoroughly studied with regard to serpentine endemism (Kruckeberg 1984), but species restricted to deposits enriched with heavy metals such as copper and zinc are also known (Antonovics et al. 1971). Some highly restricted taxa are endemic to deposits at one or a few metal mines and appear to have originated in situ from widespread, more ecologically generalized species. For example, Mimulus cupriphilus Macnair (Macnair 1989) is known only from two copper mines in the foothills of the Sierra Nevada Mountains of California and appears to be a derivative of the widespread species, Mimulus guttatus Fischer ex DC. Perhaps a comparable example among the mosses is Ditrichum plumbicola Crundw., described from lead mine wastes in Great Britain and clearly related to the more widespread species, D. lineare Sw. (Crundwell 1976). Species known as copper mosses, however, are very different biogeographically and ecologically from these narrow endemics, and certainly represent a different form of rarity reflecting different ecological limitations and evolutionary histories. The widely distributed, disjunctive copper mosses pose biological questions that are without parallel in any other plant group. The copper mosses themselves are ecologically and taxonomically heteroge- neous. They include about ten species of mosses and liverworts representing eight families. For most species the relationship with copper appears to be facultative, and typical substrate copper levels vary from one species to another (Shaw 1990). Mielichhoferia mielichhoferiana (Funck) Loeske was one of the first plants to be 131 132 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 recognized as a copper moss. The species was described on the basis of collections made by Malthias Mielichhofer in 1815 on copper mine tailings in the Salzburg area of what is now Austria, and numerous subsequent authors have reported a correla- tion between occurrences of M. mielichhoferiana and copper deposits (reviews in Persson 1956; Shacklette 1967; Shaw 1990). The genus Mielichhoferia is comprised of about 25 species in South America, Africa, and some of the Pacific Islands, and is represented by 3—4 species in North America and northern Eurasia. Mielichhoferia mielichhoferiana, with scattered occurrences across North America, Europe, and Asia (south to Japan), is the most widespread of the northern taxa. In contrast, M. tehamensts Showers is found only around Mt. Lassen, a volcanic peak in north- central California, and M. macrocarpa (Hook. ex Drumm.) Bruch & Schimp. ex Jaeg. is known from about ten disjunct sites in Canada and the United States and at one extremely isolated site in central Asia. Research described in this paper deals with certain factors that might control the distribution and abundance of Mielichhoferia mielichhoferiana. The species is exceedingly rare throughout its range. For example, in the northeastern United States, it has been found at only one site each in Maine and Michigan, and at two sites separated by over 300 km in New York. From these sites in the northeastern and north-central United States, the species is disjunct at about five proximate localities (i.e., within a few km) in the southern Appalachian Mountains along the border of Tennessee and North Carolina (Schofield 1959). Additional disjunct popu- lations occur in the western United States, boreal and arctic Canada, and Alaska. It has not been demonstrated that the presence of higher than normal concentra- tions of copper is the critical factor limiting the abundance of M. mielichhoferiana (or, in fact, any other copper moss). Although substrate copper concentrations reported in the literature for this species tend to be elevated relative to most soils (Persson 1956; Shacklette 1967; Shaw 1990), a considerable number of populations grow on soils or rocks with only trace amounts of copper (Shaw 1990, unpublished data). The species may show an association with sulfur (Schatz 1955) rather than copper (most copper deposits also contain high sulfur), but little data on substrate sulfur concentrations are available. Substrates of M. mielichhoferiana are exception- ally acidic and this ecological characteristic is in fact more consistent across the range of the species than is enrichment with copper: 21 North American soils samples had the remarkably low mean pH of 3.6 (unpublished data). Experimental studies of mineral nutrition in M. mielichhoferiana will be reported in a later paper. This paper reports studies on the reproductive biology of M. mielichhoferiana. Previous discussions of factors limiting the distribution and abundance of copper mosses have focussed on substrate chemistry, but the rarity of spore formation suggests that these species may have limited potential for dispersal and establish- ment of plants in nature. Virtually nothing is known about reproductive processes in natural populations of the copper mosses. It has been suggested that plants rare because of extreme habitat specificity might be genetically uniform (Huenneke 1991). This uniformity could result from founder events and population bottlenecks that occur during colonization of disjunct localities, and inbreeding due to small population sizes. This might be especially true of the copper mosses such as M. muielichhoferiana, in which sexual reproduction appears to occur infrequently. A series of questions were addressed in this study. What factor(s) limit sporophyte formation in natural populations? When sporophytes are formed, do they contain viable spores? How do germination percentages vary among sporophytes and popu- lations? Do germinating spores produce viable gametophytes? Do populations dif- 1992 SHAW, NIGUIDULA & WILSON: MIELICHHOFERIA 138 fer in gametophytic growth rates? Are gametophytic plants capable of vegetative regeneration following prolonged dormancy and does this capability vary among populations? MATERIALS AND METHODS Plants from widely disjunct populations were included in the experimental work (Fic. 1). These five populations represent five states spanning a range from the Atlantic to the Pacific coasts of the United States. In New York, M. mielich- hoferiana grows on moist, somewhat crumbly metamorphic rock along Cattaraugus Creek and several of its tributaries in Cattaraugus and Erie Co. (no. | in Fic. 1). The species occurs at several localities within a few kilometers of each other. Copper occurs at trace concentrations only at these sites. Collections of M. mielichhoferiana were originally made from Cattaraugus Creek in 1941. It presently FIG. 1. North American map showing the distribution of collections used in the experimental studies on Mielichhoferia mielichhoferiana described in this paper. 134 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 occurs as numerous colonies of varying sizes on vertical or nearly vertical rock. The only Michigan population of M. muelichhoferiana (Fic. 1, no. 2) grows on moist rocks along the south shore of Lake Superior in Pictured Rocks National Lake- shore. Professor G. E. Nichols collected M. mielichhoferiana there originally in August, 1933, the first record for this species in North America (Andrews 1933). Numerous collections of M. mielichhoferiana have been made subsequently from Pictured Rocks, but the population has never been marked and it is not clear to what extent all collections derive from precisely the same population. The present collection, made during October, 1990, originated at a north-facing site subject to harsh winds and severe weather from Lake Superior. Field work at several of the other accessible areas along Lake Superior did not reveal additional sites for the species. Shacklette (1967) reported elevated copper levels in substrates of M. mielichhoferiana trom Pictured Rocks. In fact, the Upper Peninsula of Michigan is famous for extensive copper deposites that have been mined for hundreds of years. In Tennessee (FIG. 1, no. 3; actually on the North Carolina border), M. mielichhoferiana occurs at several sites within a few kilometers of each other in the Great Smoky Mountain National Park (Schofield 1959). The experimental popula- tion grows at “Charlie’s Bunyan”, a large outcrop of schist amidst a mixed hardwood-coniter forest at approximately 5000 ft. elevation. The moss is abundant in protected recesses on the rock In Colorado, M. mielichhoferiana occurs at scattered localities and has been found at a number of abandoned mines in Ouray Co. At one mine, M. mielich- hofertana occurs with M. macrocarpa. The site is one of only three areas that M. macrocarpa has been found in the United States outside of Alaska. The experimen- tal population grew on mine tailings at approximately 10,000 ft elevation just north of the summit of Red Mountain Pass (Fic. 1, no. 4). The last experimental popula- tion, from California (Fic. 1, no. 5), occurred on decomposing metamorphic rock along a roadside 1n the Sierra Nevada Mountains. The moss is abundant at the site, but only over a distance of about 25 meters along the road. Field work did not reveal any additional sites in the area. The reproductive behavior of gametophytic plants was investigated by inten- sively sampling individuals from the New York population. Preliminary work showed that plants form gametangia in the spring and sporophytes mature in late summer and autumn. During June of 1990, 317 plants were collected and scored for the presence of archegonia or antheridia. Plants occurred on the gorge walls as discrete colonies that were sampled independently in order to assess small scale between-colony differences in reproductive behavior. This sampling scheme was repeated in 1991, when a total of 419 plants were collected. During the second year of sampling, the sizes of individual colonies were estimated in order to determine if there was a relationship between colony size and sexual expression. For each col- ony, the longest two dimensions were measured in centimeters. Spore germination percentages were estimated for 30 sporophytes from the Michigan population, and ten each from the New York and California populations in the following manner. Sporophytes were surface sterilized by soaking in full strength bleach for 8 minutes, rinsed in sterile distilled water, and the spore con- tents were emptied into 4 ml of distilled water. 0.5 ml of the spore suspension was then pipetted onto a Knopp’s nutrient medium (recipe in Basile and Basile 1988) solidified with 1.2% agar. Three replicate cultures were initiated from each sporophyte and the petri dishes were completely randomized on a lighted shelf with a 14 hour photoperiod. 1992 SHAW, NIGUIDULA & WILSON: MIELICHHOFERIA 135 Another experiment was initiated to determine if germinating spores produce vigorous sporelings, and whether sporeling vigor varies among natural populations. Spores from five sporophytes each from the Michigan, New York, Tennessee, and California populations were inoculated on Knopp’s medium. Only one sporophyte was included from the Colorado population because no others were available. Two days after the spores germinated, the lengths of ten randomly selected sporelings were measured under a compound microscope. Three replicate cultures of each sporophyte were inoculated and the dishes were completely randomized as in the previous experiment. The viability of gametophytes following a prolonged period of dormancy was assessed by incubating stem fragments in liquid Knopp’s medium. Gametophytic plants were collected from a series of Colorado populations during the summer of 1990. These plants were then stored as herbarium specimens until the spring of 1991. In April, 1991, 25 stems from each population were placed in Knopp’s me- dium in 5 ml sterile tissue culture tubes (one stem per tube). After one month, the presence or absence of renewed gametophytic growth was scored in each tube. This growth was most commonly in the form of elongation and branching of the original fragment. In some cases, protonemata and/or rhizoids were also produced. A sec- ond and third replicate block of 25 stems from each population were initiated during the spring and summer of 1991, and each was scored in a similar manner following one month of incubation in Knopp’s medium. RESULTS AND DISCUSSION Few sporophytes were observed in the New York population during the 1990 and 1991 seasons. This is consistent with observations made on populations else- where in the United States during the last three years in which relatively few sporophytes were observed (unpublished data), and with literature reports in which absence or scarcity of sporophytes has been noted (e.g., Hartman 1969; Schofield 1959). Repeated visits to a population in Colorado showed that successful sporo- phyte formation varies tremendously from year to year (Hartman 1969). The popu- lation at Pictured Rocks, Michigan, on the other hand, has been visited repeatedly since it was discovered in 1933, and numerous herbarium specimens have been collected with abundant sporophytes over the years. Without yearly visits and systematic sampling of populations, however, it is impossible to determine what factors limit sporophyte formation during any given year. At the New York site, over 90% of the plants collected during both the 1990 and 1991 seasons lacked gametangia of either sex (TABLE 1). In 1990, only six out of 317 plants bore antheridia, and only 20 bore archegonia. In 1991, the percentage of sterile plants was exactly the same as in the previous year (92%), but the propor- tions of plants bearing archegonia versus antheridia was reversed (TABLE 1). Since the combined abundance of reproductive plants was so low in both years, the difference between male and female reproductive expression in 1990 and 1991 may well be the result of sampling error. On the other hand, the scarcity of reproductive plants in general is clearly at least a proximate cause for the rarity of sporophytes in this population. Successful sporophyte formation is dependent not only on the formation of archegonia and antheridia, but on their proximity, since gamete dispersal is limited by the ability of free-swimming sperm to reach eggs. Plants of M. mielichhoferiana 136 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME I8 TABLE 1. Sexual expression in gametophytes from the Cattaraugas Co., New York population of Mielichhoferta mielichhoferiana. 1990 199] Total Plants 317 419 Male Plants 6 (2%) 27 (6%) Female Plants 20 (6%) 7 (2%) Sterile Plants 291 (92%) 385 (92%) Total Colonies 1s 45 Male Colonies 4 (22%) 12 (27%) Female Colonies 4 (22%) 0) Bisexual Colonies 1 (6%) 1 (2.2%) Sterile Colonies 9 (50%) (71%) at the New York site occur in more or less discrete colonies, and when these are separated by more than a few centimeters it 1s likely that successful fertilization will occur primarily or exclusively between plants within individual colonies. During both 1990 and 1991, only one colony each year was observed with both male and female reproductive plants (TABLE 1). It is not surprising that these colonies were the only ones that bore sporophytes each year. Approximately one fourth of the colonies sampled each year bore male but no female gametangia. During 1990, four colonies contained female but no male gametangia, but in 1991 no such exclusively female colonies were sampled (TABLE 1). The total number of fertile colonies was too low to determine if there was a relationship between colony size and sexual expression. In 1990, the only colony with both males and females was among the smallest (< 25 cm’). In 1991, the bisexual colony was by far the largest, occupying several meters’ of the rock wall. Studies of the factors limiting sporophyte formation in moss populations have shown that plants bearing antheridia are often scarce in spite of an abundance of plants with archegonia (review in Longton & Schuster 1983). In some dioicous mosses, spatial separation of male and female plants may severely limit fertilization frequency. At the extreme, male and female plants of some mosses appear to be separated by hundreds or even thousands of miles. For example, North American populations of Jortula pagorum contain only female plants, European populations appear to be exclusively male, and sporophytes have been observed only in Austra- lia where both males and females occur (Anderson 1943, Stone 1971). In the New York population of M. mielichhoferiana, successful sexual reproduction appears to be limited by the infrequent expression of sexuality in spite of the occurrence of both male and female plants. The percentage of spores that germinate varied greatly among sporophytes both within and between the three populations for which data are available (Fic. 2). On average, sporophytes from the California population had the lowest germina- tion (mean 42.5% + 11.2 SE) while the two others had almost identical, higher average percentages (New York, 64.9 + 9.1%; Michigan, 64.8 + 5.7%). All three populations contained individual sporophytes with very low or zero germination. In fact, about 30% of the sporophytes from the California and Michigan populations contained spore progeny exhibiting lower than 5% germination. Nevertheless, a 1992 SHAW, NIGUIDULA & WILSON: MIELICHHOFERIA ie PICTURED ROCKS, MI 1.0-> 0.9 - 0.8 - 0.7 - 0.6 - 0.5 - 0.4 - 0.3 - 0.2 - 0.1 = INnNOD PROPORTION PER STANDARD UNIT PER CENT GERMINATION G. 2. Germination percentages in families from the Michigan population of M. mielich- hoferiana. Family means are based on three replicate petri dishes; 35 spores were scored in each dish. e distribution of germination percentages in the California ancl New York populations were similar, and are not shown. See text for additional discussion. huge number of viable spores were produced in each of these populations, and numerous studies of other mosses, both rare and common, have shown similar, highly variable germination percentages. Differences in spore viability between sporophytes may have both genetic and nongenetic determinants. Although “common garden” experiments, including the present study of germination rates, suggest genetic differences, strong environmen- tal effects are also possible, if not likely. In an analogous situation, for example, it has been demonstrated that gametophytic (pollen) vigor can be affected by the nutrient status of the parental sporophyte (Young & Stanton 1990). Statistically significant differences in sporeling vigor also occurred among fami- lies within populations, and among the populations (TABLE 2). On average, plants from the Colorado and Michigan populations grew three times as large as those from California and New York over the same time interval. Tennessee plants were almostly exactly intermediate between these two extremes (TABLE 2). The presence of significant variation in vigor between families of gametophytic plants derived from different sporophytes adds support to the interpretation that family differ- ences have a genetic component. Nevertheless, effects of parental sporophyte nutri- tion and health on progeny performance is still possible, especially as sporeling growth was measured just 48 hours subsequent to germination. The early growth of sporelings may represent what is essentially an extended germination phase, af- fected by resources obtained by the spore while still in the sporangium. 138 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 BLE 2. Early sporeling growth in five ae tions of Mielichhoferia mielichhoferiana on Knopp’s medium. Mean values (+ standard errors) are based on five families (capsules) per population. Spores from each family were aoanleted on three replicate petri plates and 10 sporelings were measured on each plate. The analysis of variance for these data is shown below. POPULATION SPORELING LENGTH (um) California 2 Oia. Tea Colorado 60.6 + 3.5 Michigan 62.9 + 5.3 New York 20.3 + 2.3 Tennessee 426+ 8.5 SOURCE df. SS MS F Populations 3 2416.5 805.5 16.43** Families (within pops. ) 10 1372.1 137.2 2.80* Error 27 $130.0 190.0 t=P=001,"*=P= =~ ).0001 Taste 3. Gametophytic regeneration following 9-14 months of dormancy in Colorado populations of M. mielichhoferiana. Each block consisted of 25 gametophytic fragments per population. Integers represent percentages of growing stems out of the 25 for each population. POPULATION BLOCK | BLOCK 2 BLOCK 3 Eureka Mine 52 44 4 Goth 55 44 13 Red eee e Pass South 40 40 8 Red Mountain Pass North 33 40 Telluride 16 28 21 Yankee Boy Basin 85 72 25 Differences between the populations were also evident in studies of gameto- phytic regeneration following nine to fourteen months of dormancy (TABLE 3). When the first block of fragments was initiated the plants had been dried for approximately 9 months, the second block was initiated when the plants had been dried for 12 months, and the last block when the plants has been dried for 14 months. It is clear that plants from all populations lost viability during this period. Indeed, the decline in regeneration was drastic between 12 and 14 months of dormancy. Nevertheless, some consistent population differences were evident. Plants from Yankee Boy Basin, for example, showed the highest percentage regen- eration in every experimental block. Conversely, the Telluride population showed the lowest regeneration during both of the first two blocks, improving relative to others in the third block because viability in other populations had declined so greatly. It is noteworthy that the two populations from Red Mountain Pass re- sponded very similarly in every block. As with the previous experiments, caution should be exercised in assuming that population differences in regeneration result solely from genetic differentiation rather than environmental effects on plants while they were growing in nature. Nevertheless, consistent population differences are definitely suggestive of genetic variance for tolerance of prolonged desiccation and the ability to resume active 1992 SHAW, NIGUIDULA & WILSON: MIELICHHOFERIA 139 growth upon hydration. Similarity between the two Red Mountain Pass popula- tions, in contrast, may reflect underlying genetic similarity. These two populations originated at abandoned copper mines within a few kilometers of each other. The experimental set-up employed in this study is obviously quite unlike conditions in nature, yet variability revealed by this approach may suggest analogous variation in the propensity for asexual dispersal and regeneration under field conditions. CONCLUSIONS Understanding the reasons for rarity is fundamental to conservation science and constitutes one of the most fascinating problems in organismic biology. Prereq- uisite to understanding why some species of plants are rare is the recognition that there are different kinds of rarity (Rabinowitz 1981). The copper mosses constitute a class of plants that have no counterpart among other organisms. They have the odd combination of being extremely widespread but with highly disjunctive and sporadic occurrences. The copper moss designation for these species is a misnomer not only because some of the species are liverworts and not mosses, but also because their associa- tion with copper is inconsistent and sometimes contradictory. A dependence on elevated copper concentrations in the soil is insufficient to explain the rarity of these species. This study assesses the ability of one of the copper mosses, Mielichhoferia mielichhoferiana, to reproduce and therefore maintain viable popula- tions, by determining the factors that limit sexual reproduction and by evaluating the viability and vigor of spores and sporelings. In the New York population of M. mielichhoferiana, sporophyte formation appears to be limited by the formation of gametangia. However, those spores that are produced are largely viable, and pro- duce relatively vigorous offspring that are capable of sustained gametophytic growth. Although gametophytic growth was measured in this study just days subse- quent to germination, these plants have now formed extensive protonema and gametophores in our laboratory. Population comparisons show that viable spores are the rule and not the excep- tion, and that a significant amount of phenotypic variability occurs within the species. This variability is manifest as both gametophytic growth and the ability to regenerate after prolonged dormancy, and almost certainly reflects some degree of underlying genetic variability. These observations do not suggest that M. mielich- hoferiana as a species is genetically depauperate. Nevertheless, small populations of limited size increase the risk that chance events combined with habitat destruction pose a significant risk to the long term survival of this rare species. ACKNOWLEDGEMENT Funding for this research was provided by NSF grant no. BSR—9020126. LITERATURE CITED Andrews, A. L. 1933. be Mielichhoferia of northern North America. Bryologist 35: 38-41. Anderson, L. E. 1943. The distribution of Tortula pagorum (Milde) De Not. in North America. Bryologist ne ee Antonovics, J., A. D. Bradshaw and R. G. Turner. 1971. Heavy metal tolerance in plants. Adv. Ecol. Res. 7: | 140 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 Basile, D. V. and M. R. Basile. 1988. Procedures used for the axenic culture and experimental treatment of ae Pp. 1-16 in Glime, J. M. (ed.), Methods in bryology. Proc. Bryol. Meth. Workshop, Mai — meet ae Cc ae Ditrichum plumbicola, a new species from lead-mine waste. J. Bryol. 9: 167-169. Hartman, E. L. 1969. The ecology of the “copper moss” Mielichhoferia mielichhoferi in Colorado. Bryologist 72: oe Huenneke, L. F. 1991. Se ealaviael implications of genetic variation in plant populations. Pp. 31—44 in alk, D. A. and K. E. bi oizinger (eds.), Genetics and conservation of rare plants. Oxford Univ. Press, v York & Oxt eae ” R. 1984. ee serpentines. Univ. Calif. Press, Berkeley. Longton, R. E. and R. M. Schuster. 1983. Reproductive ae In Schuster, R. M. (ed.), New manual of on ed L: 386-462. Hattori Botanical Laboratory, Nichir Macnair, M. R. 1989. A new species of Mimulus endemic to copper mines in California. Bot. J. Linn. Soc. 100: 1-14. Persson, H. ay Studies in “copper mosses”. J. Hattori Bot. Lab. 17: 1-18. Rabinowitz, D. 1981. Seven forms of rarity. Pp. 205-218 in H. = ee The biological aspects of rare ant conservation. Wiley, New York. Sarees A. 1955. Speculations on fe ecology and photosynthesis of the “copper mosses.” Bryologist 58: 113-120. Sr eae W. B. 1959. Mielichhoferia mielichhoferiana in the Southern Appalachians. Bryologist 62: 250. Shacklete, H. T. 1967. Copper mosses as indicators of metal concentration. Geol. Surv. Bull. 1198G: Shaw, A. [ 1990. Metal tolerance in bryophytes. Pp. 133-152 in Shaw, A. J. (ed.), Heavy metal tolerance in ae evolutionary aspects. CRC Press, Boca Rat Stone, I. G. 1971. The sporophyte of Jortula pagorum (Milde) De Not. Trans. Brit. Bryol. Soc. 6: 270-277 Young. H. J. and M. L. Stanton. 1990. Influence of environmental quality on pollen competitive ability in wild radish. Science 248: 1631-1633. Contr. Univ. Mich. Herb. 18:141-147. 1992. BEEVERIA (HOOKERIACEAE), A NEW GENUS FROM NEW ZEALAND Allan J. Fife Botany Institute DSIR Land Resources Private Bag Christchurch, New Zealand INTRODUCTION In the course of revising the New Zealand representatives of the Hookeriaceae sensu Brotherus for a moss flora of that country, it was concluded that the New Zealand endemic species currently known as Prerygophyllum distichophylloides Broth. & Dix. in Dix. constitutes an undescribed genus. Given the long-term nature of the New Zealand moss flora project and the recent high level of interest in the Hookeriales (vide Buck 1987; Tan and Robinson 1990; Crosby 1974; Allen and Crosby 1986), it is opportune to discuss this species here and to describe a new genus to accommodate it. A monotypic genus, named Beeveria, is described with notes on its relationships, synonymy, geographic distribution, and ecology. BEEVERIA, GEN. NOV. ee distichophylloides was described by Brotherus and Dixon (Dixon 1915) o e basis of a collection by Donald Petrie from near Auckland. The species has ee little subsequent taxonomic attention, probably due to its relative rarity. Treatments by Brotherus (1925) and Dixon (1927) have added little new information. Sainsbury (1955, p. 402) differentiated it from its congeners (in Pterygophyllum) by a combination of coloration, and leaf cell and margin char- acterictics. He also (p. 406) discussed the leaf areolation and the gemmae-bearing pseudopodia and used them as adjunct species characteristics. Sainsbury was the first, and seemingly the only, collector to find this species with capsules. He de- scribed the peristome in considerable detail, albeit with some internal inconsisten- cies (e.g., with respect to exostomial lamellae). While Sainsbury was impressed with the aggregation of peculiar features in P. distichophylloides, there is no sugges- tion that he considered it misplaced within Prerygophyllum. Vitt and Crosby (1972) proposed Achrophyllum, a new generic name to replace the illegitimate Prerygophyllum Brid., and made appropriate combinations for two New Zealand species. They also stated the unelaborated opinion that P. disticho- phylloides should not be transferred to Achrophyllum. Neither Vitt nor Crosby has returned to the problem of the status of P. distichophylloides. Numerous gametophytic and sporophytic features, including the presence of non-homologous vegetative reproductive structures, preclude the transfer of Preryg- ophyllum distichophylloides to Achrophyllum Vitt & Crosby (TABLE |). There is no genus in the Hookeriaceae sensu Brotherus with the combination of gametophytic and sporophytic characteristics found in P. distichophylloides and thus a new genus is proposed. 142 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 Beeveria Fife, gen. nov. Plantae vivae citrino-virides, siccae eiusdem coloris vel atrovirides. Caules prostrati, saepe in pseudopodia nuda, 2-3 mm longa, gemmarum capitulo termi- nata desinentes. Folia haud vel leniter asymmetrica, illa in seriebus dorsalibus ventralibusque ab illis in lateralibus vix divergentia, sicca valde crispata, elliptica, acuta vel apice infirme cuspidata, integra vel crenulata, limbo nullo. Costa singu- laris, simplex. Cellulae superiores laminales laeves, hexagono-isodiametricae, in- firme vel modice collenchymatosae; cellulae marginales diversitatem nullam prae- bentes. Gemmae in capitulis terminalibus dispositae, anguste fusiformes, 160-240 wm longae, septis transversis 5-9 provisae. Dioeciae. Setae erectae, laeves. Capsulae 1.0-1.5 mm longae. Exostomatis dentes leniter sulcati, ca. 400 2m longi: endostoma membrana basali humili segmentisque perforatis praeditum. Sporae 10— 15 um, virides, subtiliter papillosae. Beeveria distichophylloides (Broth. & Dix. in Dix.) Fife, comb. nov. Fic. | Pterygophyllum distichophylloides Broth. & Dix. in Dix., Bull. Torrey Bot. Club 42: 106, pl. 9, fig. 11. 1915. Type. NEw ZEALAND: near Auckland, 1892, D. Petrie 800 (BM-Dixon!, WELT’). Pterygophyllum colensoi Broth. ex Dix., Trans. New Zealand Inst. 3(5): 290. 1927, nom. nud. BASED ON: Sine loc., Colenso 3115 (BM ex K). The generic name honors Jessica E. Beever, whose collections and taxonomic work have greatly increased our knowledge of the New Zealand moss flora, espe- cially that of the northern parts of the North Island, where Beeveria distichophyl- loides is most frequently collected. Plants medium-sized, oily yellow-green when fresh, unchanged or dark-green when dry. Stems prostrate, sparsely to moderately branched, pale brown to yellow, to 4.5 cm, ca. 3 mm wide (including leaves), often ending in naked pseudopodia 2— 3 mm long and terminated by a capitulum of gemmae; in cross-section with a distinct central strand, lacking a hyalodermis, with 2—4 cortical cell layers with moderately thickened walls; rhizoids on lower portion of stem dense, pale brown, + smooth, much-branched; pseudoparaphyllia foliose, lanceolate, ca. 300-400 xm long. Leaves inserted in 6 ranks, imbricate, complanate, nearly symmetric, those in dorsal and ventral ranks scarcely differentiated from those in lateral ranks, strongly crisped when dry, elliptic, tapered to acute or weakly cuspidate apices, plane, not concave, unbordered, entire (often somewhat crenulate due to collapsed walls of marginal cells in dry material), (1.0—)1.3—2.5(—2.8) x 0.5-1.3 mm; costa single, ill- defined, ca. 35-60 «m wide at midleaf, markedly dilated in lowest third or more, unbranched, % to more than %4 the length of the leaf, in cross-section biconvex, acking stereids; upper laminal cells smooth, thin-walled (ca. 2 wm at thinnest point), weakly to moderately thickened at corners, hexagonal-isodiametric, 24—42 wm diam., gradually becoming + oblong and non-collenchymatous toward base; marginal cells and alar cells not differentiated. Gemmae borne in terminal capitula, narrowly fusiform, 160-240 ~m long, with 5—9 transverse septa. Dioicous; peri- gonia not seen (nor did Sainsbury, 1955, p. 406). Perichaetial leaves ovate- lanceolate, ecostate, ca. 1.2 mm long. Setae lateral, ca. (7—)9-11 mm long, erect, smooth, twisting weakly to the left throughout, red, ca. 150 wm diam., in cross- section lacking a hyalodermis, with 1-2 layers of thick-walled cortical cells and an ill-defined central strand; capsules pendent, ovoid from a short neck, 1.0—-1.5 mm — 1992 FIFE: BEEVERIA ene Messe ti rakes OS™ SSPE I ai ei RO ! FIG. 1. Beeveria distichophylloides. a. Leaf apex. b. Margin at midleaf. c. Upper laminal cells. d— h. Leaf outlines. 1. Gemmae. j. Moist stem. k. Dry stem. |. Exothecial cells. m. Operculum. n. Portion of peristome, showing outer and innter surfaces of exostome tooth, endostomial membrane and seg- ments. 0. Moist stem with pseudopodia. p. Moist stem with sporophytes. q, r. Perichaetial leaves. Scale = 165 um for a—c, 1000 um for d-h, q, r; 2300 wm for j, k, 0; 100 um fori, 1, n; 3500 wm for p; 1400 wm m form. From Fife 6068 and Sainsbury 4388 (both CHR). 144 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 = “ 2 i IG. 2. Total distribution for Beeveria distichophylloides. Distribution dots centered on NZMS 260 1:50,000 topographic map sheets. long, weakly constricted below the mouth and smooth when dry, reddish-brown; exothecial cells oblong to isodiametric, firm-walled, rather weakly thickened at corners, 12-18 sm in greatest diam.; annulus well developed, falling with the operculum; operculum long-rostrate from a conic base, ca. 0.8 mm long; exostome teeth yellow, lanceolate with a rather distinct shoulder, bordered, weakly furrowed (gap between portions of tooth less than 6 wm, extending to less than % the length of the tooth, not visible with stereoscope), ca. 400 xm long, finely cross-striate below, baculate above, with adaxial lamellae projecting as weak trabeculae (6-9 sm long near base of tooth); endostome pale yellow, with basal membrane 100-120 wm high, segments ca. 260 ~m long, nearly equaling the teeth, keeled and weakly perforate, papillose, cilia absent. Spores (10-—)12-15 um, green, very finely papillose. Calyptra mitrate, smooth, lobed at base [description from Sainsbury (1955, p. 405)]. Distribution. Beeveria distichophylloides is known only from New Zealand. There, it has a scattered distribution between latitudes 35° and ca. 42°19’S on the two main islands. It is recorded from North Auckland, Gisborne, Hawke’s Bay, Nelson, and Westland land districts (Fic. 2). It is not known from any offshore island except D’Urville Island (Nelson Land District). It displays a distribution similar to that of nikau palm (Rhopalostylis sapida) and several mosses of northern affinities, e.g., Braithwaitea sulcata, Catharomnion ciliatum, Ctenidium pubescens, and Tayloria callophylla. cology. Beeveria distichophyloides occurs on moist, shaded rock (usually, if not always, limestone) and clay or marl soil, often in association with stream margins or beds of intermittent watercourses. Distichophyllum microcarpum and Achrophyllum dentatum are frequent associates. It ranges in altitude from ca. 90- 1992 FIFE: BEEVERIA TABLE 1. Comparison of character states in Beeveria and Achrophyllum. Character State in Beeveria State in Achrophyllum Dry coloration Shape of gemmae Location of gemmae Nature of exostome tooth furrow Yellowish green Fusiform, transversely septate Terminal on pseudopodia Less than 6 wm wide; not visible with stereoscope Brown or black L- or T-shaped, trans- versely septate Epiphyllous on intra- marginal cells Greater than 30 wm wide; visible with stereoscope (40-50 x Length of exostome trabeculae 6-9 wm 18-24 um Height of endostomial basal ca. 100-200 wm ca. 300 wm membrane 300 m on the North Island and from 20—600 m in the northwest portion of the South Islan Discussion. The generic name Prerygophyllum Brid. (Mant. Musc. 149. 1819) is illegitimate; a proposal to conserve this name against Achrophyllum Vitt & Crosby (Margadant et al. 1972) was rejected. A combination in Achrophyllum for P. distichophylloides has never been published. TABLE 1 summarizes the gametophytic and sporophytic features which distinguish Beeveria and Achrophyllum. In Beeveria, gemmae occur in terminal capitula atop short pseudopodia and are fusiform and transversely septate. The pseudopodia of Beeveria are reminiscent of the unrelated genus Aulacomnium. Sporophytically, the features that separate Beeveria from Achrophyllum are quantitative rather than qualitative (TABLE 1), but in aggregate they support generic segregation. In Beeveria, exostome teeth are weakly furrowed, short, and weakly trabeculate. The furrow is less than 6 4m wide at base, extends ca. %4 the tooth length, and is not visible under stereoscope; the teeth are ca. 400 xm high, with lower trabeculae 6-9 um long. In Achrophyllum quadrifarium (the generitype) the teeth are strongly furrowed, longer, and strongly trabeculate. The furrow is 30-36 pm wide at base, extends %4 the length of the tooth, and 1s clearly visible under the stereoscope; the teeth are more than 700 um high, with lower trabeculae 18—24 wm long. In Beeveria, endostomial segments are perforate and arise from a basal mem- brane ca. 100-120 wm high, while in the generitype of Achrophyllum segments are non-perforate and arise from a basal membrane ca. 300 «wm high. Buck (1987) synthesized the large amount of recent information and opinion (vide Tan and Robinson 1990; Crosby 1974; Allen and Crosby 1986) concerning the classification of the Hookeriales and proposed a reorganization of the order into five families. Buck’s familial concepts place considerable emphasis upon gametophytic features and, despite some criticisms which have been leveled at them (cf. Tan and Robinson 1990), they are accepted here. Buck reduced the Hookeriaceae to only six genera (Achrophyllum Vitt & Crosby, Cyathophorella (Broth.) Fleisch., Cyathoph- orum P.-Beauv., Dendrocyathophorum Dix., Hookeria J. E. Sm., and Schimpero- 146 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 18 bryum Marg.); all other genera in Brotherus’ Hookeriaceae are placed in four segregate families. He placed Hookeria, Achrophyllum and Schimperobryum in one “lineage” within his restricted Hookeriaceae. If Buck’s evolutionary hypothesis is accepted, the lack of a central strand, the large, unbordered leaves, the short, hexagonal and porose leaf cells, the presence of pseudoparaphylhia, and other fea- tures in Beeveria dictate its inclusion in both the restricted family and in the lineage “centered around Hookeria.” Within the “lineage” the nature of the leaf cells and the single, forked costae (interpreted here as synapomorphies) of Achrophyllum and Beeveria differentiate them collectively from Hookeria and Schimperobryum. By this interpretation, Beeveria and Achrophyllum are sister taxa, forming a clade whose relationship to Hookeria and Schimperobryum ts not resolved. he presence of pseudopodia and the fusiform, apical gemmae of Beeveria and the L- or T-shaped, intramarginal gemmae of Achrophyllum are apomorphies which differentiated the two allied genera. They suggest that Beeveria was not the ancestor of Achrophyllum, and the acceptance of Beeveria does not render Ah paraphyletic. erences in coloration of dried plants provide further distinction between eae and Achrophyllum. The relatively weakly collenchymatous thickenings of the leaf cells, remarked upon by Sainsbury (1955, pp. 402, 406), are not considered significant as this feature varies markedly among various species of Achrophyllum. Sainsbury’s statements (1955, p. 406) that the exostome teeth of Beeveria (sub eee ani aude aC aed lack both (adaxial) lamellae and median fur- rows, and that median divisural line is more conspicuous relative to other species which a ei in Prerygophyllum, are incorrect. Both adaxial (ventral) lamellae and a median furrow are present (but the latter is not visible under a stereoscope) and the “zig-zag” divisural line is no more conspicuous than in Achrophyllum quadrifarium. o other genus in the Hookeriales bears gemmae in the manner of Beeveria. The one genus in the order with structures vaguely similar, the monotypic American-African genus Adelothecium (placed in a monotypic family by Buck [1987]), bears gemmae on axillary stalks on specialized microphyllous branches. The thick-walled, porose leaf cells, brownish pigmentation, lack of a central strand, and ascendent habit of Adelothecium are among many features which preclude the placement of B. districhophylloides in, or even close to, that genus, ACKNOWLEDGMENTS Ilustrations were prepared by Sabrina Malcolm. Janie Glasson and Mary-Ann te provided help with preparation of the distribution map and Elizabeth Edgar translated n scription into Latin. Phil Garnock-Jones and Bryony Macmillan provided criticism that allowed sae ca of the manuscript. »ward Crum has provided guidance for many years and it is a pleasure to dedicate this contribu tions to him on the occasion of his 70th birthday. pe ~) LITERATURE CITED Allen, B. H. and M. R. Crosby. 1986. A revision the genera Pilotrichidium and Diploneuron (Musci: Hookeriaceae). J. Hattori Bot. Lab. 61: 45— Brotherus, V. F. 1924-1925. Musci. In: A. oat and K. Prantl (eds.), Die nattirlichen Pflanzen- familien, ed. 2, Bande 10-11. Baceinnan: Leipzig. 1992 FIFE: BEEVERIA 147 Buck, W. R. 1987. Taxonomic and nomenclatural rearrangement in the Hookeriales with notes on West Indian taxa. Brittonia 39: 210-224. Crosby, M. R. 1974. Toward a revised classification of the Hookeriaceae 38: 129-14 e Dixon, H. N. 19 ey ae rare Australasian mosses, mostly from Mitten’s herbarium. Bull. Torrey Bot. Club i 93- . 1927. Studies in ih bryology of New Zone with special reference to the herbarium of Robert Brown. Part 5. =) . New Zealand Inst. 3: 239-298. Margadant, W. D., ve Miller and C. M. Matteri. 1972. Proposal for the conservation of the generic name Persgophiin Brid. 1819 in Musci. Taxon 21: 536. Sainsbury, G. O. K. 1955. A handbook of the New Zealand mosses. Bull. Roy. Soc. New Zealand 5: Musci). J. Hattori Bot. Lab. — 1-490. Tan, = and H. See 1990. A review of Philippine hookeriaceous taxa (Musc1). Smithsonian Contr. . 75: i-iv, 1- Vitt, . ia and M. = ae 1972. Achrophyllum—a new name for a genus of mosses. Bryologist 75: 174, 175.