C:\WAW\CRYPTMS\Mosses2005.wpd

Revised: March 19, 2005

(a Working Document)

Note: the pagination changes as we make revisions. The families are alphabetically arranged. For genera that have been moved to other families, consult the Index and Catalog. Portions of the manuscript that are in need of amplification are indicated by *** .

Preface

Introduction

Historical collectors of Colorado bryophytes

Habitats of bryophytes

Geography of Colorado bryophytes

Little-recognized facts about bryophyte distributions

Some thoughts about ecology, evolution, and taxonomy

Geobotany of bryophytes

Key to the genera

Family, genus and species treatments

Index and catalog of Colorado Mosses

Index by specific epithets

Derivations of the generic names

Meanings of the specific epithets

Sources for Illustrations

Key to the genera

Family, Genus, and species treatments

Index and Catalog of Colorado Hepatics

Index by specific epithets

Derivation of the generic names

Meanings of the specific epithets

Sources of Illustrations

Authors of Bryophyte taxa

Glossary of bryological terms

Historical references

1. Leo Lesquereux (1884)

2. The Colorado Collections of A. J. Grout (1914)

3. The type collections of T. S. Brandegee

4. Colorado Collections reported by T. P. James in Wheeler Expedition Report, 1878

5. F. J. Hermann. Mosses of Rocky Mountain National Park

----------------. Additions paper, 1976

6. Original descriptions of mosses described from Colorado

7. Seville Flowers' Mosses: Utah and The West

8. Phytogeographical comparisons

9. Colorado Bryological Hot Spots

Boulder Mountain Park

Mount Evans

High Creek Fen

Hall Ranch ***

Acknowledgments

Bibliography

"His work will not end until we have traced each delicate web that ties us with all the life of the planet. His greatest worry was that we will finish the planet before we finish that work. The world and all its beauty belongs to all of us, not to politicians and mere ideologies. That our weapons could eradicate all life - plants, insects, mammals as much as man - was a terrible fear for him, and he worked hard, contributing hours and money, against that fear." Eric Hermann in Vos & Reznicek (1988).

I (senior author) first learned about mosses when I was about twelve years old. When I was in High School I built a small terrarium with mosses and lichens in it, and entered it in a contest (this was before the days of Science Fairs) at the American Museum of Natural History. With the $25.00 I won as first prize, I was urged by my biology teacher to buy a small book by A. J. Grout, Mosses with a Hand Lens. Thus, I acquired my first moss textbook on Christmas Day, 1934, and still find it useful eighty years later!

Mr. Grout was a high school teacher on Staten Island, and an associate of the New York Botanical Garden. He spent a lifetime becoming a specialist on mosses, and eventually produced a classic three-volume work entitled Moss Flora of North America, written with the aid of as many specialists as he could muster. Mosses with a Hand Lens was written from the standpoint that Grout felt that, with proper attention and guidance, a person could learn to recognize most of the common mosses of the New York City Region with the use of only a hand lens and his book. This book was so popular that people began to be fairly proficient but needed more. To meet the demand, Grout then wrote Mosses with Hand Lens and Microscope. This required more diligent and critical study, and the area covered was greater. Most professional biologists of my generation in America owe their starts to this remarkable man.

Grout's little moss books, now sadly out of print, were extremely useful in the Eastern United States, and have been followed by many useful books currently available there. One of the very successful ones based on Grout's philosophy was How to Know The Mosses, by Henry S. Conard 1956), one of the most beloved of teachers of budding bryologists. He was professor at Grinnell College, Iowa. Unfortunately, Conard's book was also designed for use in the eastern U. S., and did not help me at all in 1944 when I was collecting mosses as a hobby in the Columbia River Gorge. However, this fact was not too tragic, because it resulted in my corresponding with Dr. Conard, who helped me identify my mosses. Perhaps it was through helping people like me that he realized that he had to publish an expanded version that covered the entire United States. The latest edition is co-authored by one of Conard's protegés, Paul Redfearn, who has just now retired.

One of Grout's disciples was Seville ("Bill"), or to most of his students, "Doc") , a long-time professor of all sorts of biology and botany at the University of Utah. Bill Flowers, over the years, emulated Grout in producing mimeographed outlines of keys to mosses, liverworts, and ferns, of Utah. He amassed a large collection as well, and painstakingly produced marvelous pen-and-ink illustrations for a book on the mosses of Utah. He did not live to see this published, but after his death, Howard Crum, bryologist at the University of Michigan, edited it and it was published under the name Mosses: Utah and the West. For Colorado bryologists this book has excellent descriptions and illustrations of nearly all of our Colorado species. It also has one of the best introductions to moss morphology ever written in America. An excellent account of Flowers' life and work has been published by W. H. Behle (1984).

There are now several excellent moss floras available, but few of them are especially pertinent to the Rocky Mountain region. The Pacific Northwest is well served by Elva Lawton's Moss Flora of the Pacific Northwest; Crum and Anderson have a two-volume Mosses of Eastern North America, and William D. Reese has recently published a Mosses of the Gulf South. Sharp, Crum, and Eckel have published a two volume Moss Flora of Mexico. If anyone is looking for new worlds to conquer, let them plan a Moss Flora of Arizona and New Mexico!

This guide to the Colorado mosses is patterned after the excellent series of little books written by the late Prof. Helmut Gams of Innsbruck, Austria, entitled Kleine Kryptogamenflora von Mitteleuropa. They are "Excursion-floras" - field guides to mosses, lichens, ferns, fungi and algae of Central Europe, and their author was one of the keenest field men in Europe. I remember following him in the field and listening and looking as he pointed out (sotto voce to himself) all of the various species he saw as he passed by. This is only one of the many ways one learns the lower plants.

As a compromise owing to the desirability of having an easily carried field manual, an Excursion Flora contains only keys and ecological notes. For more detailed descriptions and illustrations they presume the availability of more comprehensive literature. They also expect a certain amount of general biology background on the part of the user.

This second edition still does not accomplish my original objective of eventually providing an instructional introduction, but available books and the planned Moss Flora of North America fill that need. Much remains to be done in Colorado in refining our concepts of habitats and distribution. The first edition spawned several local amateurs who now are adept in recognizing mosses, and perhaps more will be encouraged to enter the field with this revision.

One who constructs diagnostic keys can usually make a key that satisfies himself; that is, he can distinguish his species under the best of circumstances, when all of the necessary things (including sporophytes) are present. It is quite another thing to produce a key that works for his students, and that has to "work" even if vital parts are missing. This is the great problem with mosses, especially in an arid area like this one -- that most of the species are rarely or never found in fruiting condition. Thus, the sporophytic information, which often makes identification surer and much easier, cannot be included in a key. The writing of the key thus becomes a difficult art, with inevitably equivocal results. Furthermore, without sporophytes it is impossible to write a satisfactory key to the larger groups -- families and genera -- so that a different method has to be tried. Mrs. Patricia Nelson, my student, worked out a very useful polyclave key on IBM cards called Random Access Key to the Genera of Colorado Mosses, which I consider to be an indispensable adjunct to the guide. Unfortunately there are no copies left!

All of the species listed in the Guide are documented by herbarium specimens in the herbarium (COLO) of the University of Colorado Museum, which now contains over 112,000 specimens of bryophytes. I welcome comments and criticism, and am always interested in adding to the collections through contributions from collectors.

Although this distillation represents more than fifty years of sporadic research, the work is now only well begun in earnest. The stones are laid, but the building has not assumed its final shape. We think we know what grows here, but there will be additions as soon as active field work by many students begins. We have some notions of habitats, but they are primitive and need refinement. Every observer comes to the field with a different pair of eyes and a different core of experience, and there have been too few of us up to now. I shall be satisfied if this guide will serve to bring bryology within the competence of students and ecological researchers, and serve as a guide to visiting bryologists from abroad, until such time as a more sophisticated work is made possible.

This manuscript is a guide book in progress. We are making it available on this web site to make the sections parts available as they near final form. Many of the smaller families are ready for use, but some of the larger or more difficult ones are being worked on continuously. It is hoped that this version might be useful to amateurs, students, amd professional ecologists. This manuscript is an identification guide with more detailed descriptions than are usually given in keys, supplemented by notes on field aspect, ecological niche, and other interesting observations. A collaborative production contributed by a host of professional bryologists is in preparation, which will provide detailed descriptions with illustrations of all the species. Tentative treatments are available on the New York Botanical Garden web site <www.nybg.org/bsci/bfna>.

This is not an introduction to the study of mosses and liverworts. In fact, it is not a primer for beginners, but a source book and guide for persons already literate in the subject, especially those who come to Colorado from previous experience with bryophytes from other parts of North America and abroad. Elementary books on the life histories, morphology, and ecology of bryophytes are available elsewhere. This book concentrates on the taxonomy and ecology, and the history and evidence for the occurrence of the entire moss and hepatic flora of Colorado, from the time of the historical expeditions of the nineteenth century to the present.

Our intention has been to see if we can bring the knowledge of occurrence of the species up to the point of diminishing returns. We are convinced that there is more to discover, and that our understanding of geographical distribution and of habitats is still very imperfect. We are especially interested in drawing attention to the precise microhabitats of the species, for field work in bryology is only successful when the mind is prepared. We can be grateful to whatever creative force there is on our planet that bryophytes, no less than other plants, are not distributed at random! If they were, our task would be hopeless.

We are also aware that there are ways by which we can train our eyes to see distinctive characters of bryophytes that enable us to recognize the different taxa in the field. These are what the ornithologists call "flash characters". Such characters are not always easy to describe nor are they infallible, but as one gains experience, they become indispensable for field identification and can prevent excessive accumulation of collections of species that we are not particularly interested in cramming into our limited space in home or laboratory. Being successful in finding ways to recognize Ceratodon purpureus, that most pervasive herbarium weed, becomes a valuable attribute in the field botanist.

Learning to recognize habitat is very important in developing an understanding of the relative fidelity of bryophyte species to acidic, calcareous, mineralized, acidic or basic substrates,. We still know very little about this and wonder whether our local situation agrees with that of the same species in Scandinavia or Australia.

Colorado does not appear at first glance to be a very interesting place for bryophytes -- much too dry and steep. One doesn't see many mosses beside the superhighways, and then only a few very conspicuous and common ones on seeping road-cuts! Also, the pace taken by hikers does not lend itself to our subject. While a vascular plant collector may cover twenty miles in a day, the lowly bryologist may not stray more than a few hundred yards along the trail before being forced to stop for an hour or so on hands and knees beside an initially unlikely-looking site. Thus, if we were to plot on a map the locations of collections of bryophytes in Colorado, the dots would follow the roads.

We make no apology for the technical difficulty of identification of bryophytes. Anyone who ha a hand lens can, with some guidance in the field, learn many of the common and conspicuous species, For this purpose we have produced a primer for those foresters and wetland managers who need to be able to do this to a limited degree.. For amateurs, being able to recognize some common bryophytes adds to the amateur's joy in the field. In fact, Grout's old book, Mosses with a Hand Lens, demonstrates that it can be done.

However, in all taxonomic studies but particularly in the smaller life forms more sophisticated equipment is needed for identification requiring high magnification and dissection or sectioning of leaves and stems. Also, now that greater emphasis is being made on subtle characteristics and biochemical information, it is becoming difficult for amateurs and other nonprofessional bryologists to do this work. Up to a point, it is mostly exhilarating fun, but suddenly one hits a stone wall. This happens even to professionals who lack not only the money for physical equipment but also the indispensible reference books, especially as the older ones are becoming unavailable or are being reprinted. These, as well as the new books, are only available at astronomic prices.

The basic equipment for a serious amateur includes two microscopes, one compound for high magnifications and prepared slides, and one stereoscopic for magnifications up to 100x for gross examinations. Dissecting needles, razor blades (double-edged "blue blades" from which slivers can be broken off to serve as knives), slides and cover-glasses, a propane burner for heating dry specimens in water, and a few chemical reagents, are all necessary. Depending on the student's affluence, a fiber-optic light source and a digital camera mounted on the compound scope will make life easier.

Colorado bryophytes have never been treated fully. The earliest paper attempting to list them is that of Leo Lesquereux (1874). A. J. Grout visited Colorado once and published a short paper (Grout 1916) on his collections near Tolland, in Gilpin County. Fred J. Hermann (1970, 1987) published a few papers, one on Rocky Mountain National park and the other on additions to the flora. Hermann's life and career is reviewed by Voss and Reznicek (1988). Craft and Craft (1952) taught at the Adams State College at Alamosa and published a three page paper listing by county 40 mosses they had collected in southern Colorado, but cited no specimens. They implied that the collections were in that institution. H. S. Conard, Frederick McAllister, and others have collected mosses sporadially but no publications are based on their collections.

The real founder of the bryophyte herbarium (COLO) at the University of Colorado has to be Geneva Sayre (Pfister 1993). She came from Menlo, Iowa, and went to Grinnell College, where she came under the wing of Henry S. Conard. When she graduated, during the Great Depression, she could not afford to go to graduate school, so Dr. Conard invited her to stay another year at Grinnell so that he could continue to be her mentor. In 1934 Conard arranged for her to spend the summer with Dr. Grout in Newfane, Vermont. With Grout she worked on the preparation of the text for the Splachnaceae, Timmiaceae, and Aulacomniaceae for the Moss Flora of North America. In 1935 she received the Master's Degree at the University of Wyoming, her thesis being the work just mentioned. In 1935 she went on to get a doctorate at the University of Colorado under Joseph Ewan.

"Jimmy" had no mentors in the region so she kept in touch with Conard and Grout. The difficulties of writing a moss flora for Colorado (her thesis subject) were immense. There was no moss herbarium here; the library was poor. Grout's flora was only two-thirds published. Nevertheless, she did write a thesis. She was never very proud of it, through no fault of hers; the information was just not ready for her, and field work must have been extremely difficult without a car. During this time she studied the vascular plants. It is said that she wrote a spring flora of the Laramie area and a mountain flora of Boulder County; these were probably never really published and evidently no longer exist. But she did leave a small collection of her bryophytes, some 2,000 specimens, which I discovered in a cardboard box soon after I arrived on campus. This was our beginning.

Geneva Sayre held an instructorship in the Biology Department and taught at the University Camp from 1938-1941 before taking an Assistant Professorship at Russell Sage College in Troy, New York. There she had to teach almost everything in the Biology Department (except bryophytes!) and spent her entire career there, becoming head of the department in 1946. She took early retirement in 1972 and took a part time position at Harvard in the Farlow Library and Cryptogamic Herbarium.

Because of the pressures of her various academic duties and the lack of an herbarium, she was never able to pursue a career in the taxonomy of bryophytes. Instead she embarked on several bibliographic projects that have produced extraordinarily important basic tools for research in botanical bibliography (see Sayre 1957-1975).

I got to know Geneva Sayre at national meetings and always looked forward to finding her there. She was a bright young lady and full of ideas and conversation topics in bryology. In her biography there is nothing said about her having anything to do with the formation of the bryological herbarium at Boulder, but I feel we owe her a great deal for what she achieved, and it is a great tribute to her mentors that she fulfilled her mission in life so abundantly.

Fred Hermann (see Voss & Reznicek 1988) came to Fort Collins in 1970, with the herbarium of the U. S. Forest Service, where he had served in Washington, D.C. and became the leading American specialist on the genus Carex. From that time on until his death in 1987 he devoted his spare time to collecting bryophytes, mostly in Rocky Mountain National Park, recruiting a now famous professor of philosophy, Holmes Rolston III, to the enterprise. Fred and I frequently went into the field together and we have at herbarium COLO duplicates of most of his Colorado collections. His principal collection went to the University of Michigan, where he started his botanical career as a freshman student in the 1920s. It was a great privilege to have had his companionship during those years.

Fred was fortunate to find a compatible colleague at the State University at Fort Collins -- Holmes Rolston III, a philosophy professor. We are fortunate that these two men spent many days collecting bryophytes in and near Rocky Mountain National Park. Holmes was just the companion Fred needed, for it is very difficult to be a bryologist and hike alone in safety in the wilderness during ones declining years. Holmes proved to have not only stamina, but an excellent eye, and collected a number of exciting specimens. However, when Fred died, Holmes was able to devote all of his time to his favorite study--the application of religious teachings and ethics to environmental conservation. He published many significant papers and traveled world-wide on the "lecture circuit", and in 2003, he earned the Templeton Prize for Progress toward Research about Spiritual Realities, joining Mother Theresa and Alexander Solzhenitsyn in the list of notable recipients. We bryologists can be very proud that such an individual has worked in our midst.

Hepatics are much less well known in Colorado. Alexander Evans published a list of Colorado Hepatics (Evans 1915), but never visited Colorado to see them in the field. T. C. Frye & Lois Clark published a work on the North American Hepatics (Frye & Clark 1937, 1946) and cited specimens from Colorado, but likewise did no visit the state. Their work at least permitted us to compile a check list from the literature. While Rudolph Schuster published a giant six-volume work on the Hepatics of part of Eastern North America (Schuster 1966-1992), he did not include observations on the western American flora. At the present time Won Shic Hong, College of Great Falls, is publishing papers on the western American leafy liverworts and has examined our collections and provided identifications (Hong 1986-2000) which have proved very helpful to us, but the American West needs more active workers in the field.

There is no single guide that covers all of the Colorado mosses. Seville Flowers, Mosses: Utah and the West lacks several of our genera and species, as does Crum, Steere, and Anderson, Mosses of Eastern North America; Sharp, Crum, and Eckel, The Mosses of Mexico, treats several Colorado species that otherwise are unknown in North America. Steere's Mosses of Arctic Alaska has most of the disjunct arctic species but no keys are provided. The most useful book covering our flora actually is Nyholm's Moss Flora of Fennoscandia, which has been most useful to us because so many species are common to northern Europe and the Rocky Mountains. And lastly, a few of our mosses have been added to our flora as the result of recent monographs such as Blom's revision of the Schistidium apocarpum group in Norway and Sweden, which has added over ten more species in this group than was known. A few mosses are widely disjunct from Middle Asia, one of them, Didymodon anserinocapitatus, having been described little more than a decade ago. Thus, the present manuscript should be of great value in providing information about species missing from the other available books.

The first edition of this Guide to the Mosses of Colorado was published in July, 1973, as Occasional Paper No. 6 of the Institute of Arctic and Alpine Research. In the 28 years following this publication, numerous additions and changes have been necessitated.

There were no bryologists in Colorado during the historic exploratory period of the nineteenth century (the United States Geological Surveys (see Rothrock, 1878) and the Hayden Surveys (see Foster, 1994). Not that the Lewis and Clark expedition would have collected many mosses had it marched through what is now Colorado. Only two specimens of bryophytes, Hypnum (= Kindbergia) oregana and Bazzania trilobata were known to be collected (Moulton 1999), both of them are exceedingly common northwestern species that merely could have been separated from some vascular plant's roots during the preparation of specimens. There were hardly any botanists either; most were medical men attached to expeditions. It is no secret that botanists are the last in line for positions on general exploring expeditions. They are usually held hostage by the demands of the more important members. Their collecting spots were usually accidental, such as rest stops along the way. When the geographers, surveyors, military, geologists, and zoologists wanted to move their camp, the botanists had to go along with them. Even those who have been identified as "botanists" collected only vascular plants. The list is small; most were male.

Fortunately, in the historic period there were two great bryologists in America, who may rightly be called jointly, the "fathers" of American bryology -- William Starling Sullivant and Leo Lesquereux (see Rodgers 1940). Thanks to the efforts of these men and their collaborators, Thomas P. James, and Coe Finch Austin, they put together the first "comprehensive" moss flora of North America (Lesquereux & James 1884) ).

Biddlecome, Miss Hannah J. Amateur bryologist from Springfield and Columbus, Ohio. Bryum biddlecomei, from Alma, Park Co. Andrews, in Grout (MFNA 2:224. 1935), wrote: "The original collector is not known with certainty. Miss Biddlecome [who never was in Colorado] had received it from Mrs. Haines, who may have received it from someone else."

Brandegee, Townshend Stith (1843-1925). Civil engineer with the Atchison, Topeka and Santa Fe Railroad, assigned in 1871 to Canyon City area, became the county surveyor of Fremont County, and is said to have "laid out' the town of Florence. He served as botanist on the Hayden Survey in 1875 and provided an early and very useful description of the flora of southwestern Colorado. His bryophytes came from "S. W. Colorado" or "within 100 miles of Cañon City." He was the only resident botanist that collected any mosses here. A few of the fragmentary specimens on which new species were based have not been located again. Brandegee later went on to have a distinguished career in California, where he was a botanist at the University of California, Berkeley.

Conard, Henry Shoemaker (1874-1971). Important American bryologist and teacher of bryologists, professor at Grinnell College, collected some mosses near Buena Vista in the summer of 1941. His herbarium is at Iowa State University.

Cooper, David. Contemporary plant ecologist, specializing in the floras of Colorado high altitude fens and responsible for important collections of Sphagnum and Amblystegiaceae.

Craft, James H. professor at Adams State College, Alamosa, published a list (1952) of mosses collected in several southern Colorado counties, mostly determined by Conard. These evidently have not been preserved.

Crandall, C. S. (1852-1929). Professor of Botany at Colorado Agricultural College, collected some mosses in what is now the Roosevelt National Forest in 1894.

Downie, Timothy Campbell, 1830-1875. Nicknamed "Major Downie", he collected some mosses in Colorado in 1868, at Twin Lakes, with the Hayden Survey (Ewan 1981). According to Hayden (1869), the expedition came into Colorado first from Cheyenne, visiting points in the outer Front Range to Colorado Sprigs and Raton Pass. It returned to Colorado from Santa Fé, up the Colorado River, through the San Luis Valley, over Poncha Pass, through South Park, and into the upper Arkansas River Valley. Presumably Twin Lakes was their last camp before returning to Denver. The collections were to be turned over to the Smithsonian Institution, but we have been unable to find them.

Flock, JoAnn W. (***) Student at Univ. of Colorado, collected and published on the inventory and ecology of alpine lichens and bryophytes on Niwot Ridge, Boulder County (Flock 1978). Also made inventories of the mosses and lichens of Cape Prince of Wales, Alaska.

Goodding, L. N. (1880-1967), Wyoming botanist, student of Aven Nelson, collected a few mosses in Rout County in the Park Range (Ptychostomum pallescens, No. 1781 [COLO]) in August, 1903. Goodding's main collections were at the Rocky Mountain Herbarium, whose bryophytes were given to the University of Tennessee, where they were later destroyed in a fire that destroyed the herbarium.

Grout, Abel Joel, (1867-1947). Grout made a short visit to Colorado in 1916, and collected in the vicinity of the University of Colorado's summer camp at Tolland, Gilpin County. See appendix, and Steere (1948).

Haines, Mrs. Mary Parry (1826-1884). A naturalist of eclectic tastes, custodian of the Paleontology Dept. of a museum in Richmond, Indiana, and an amateur horticulturist. Although she never visited Colorado she was responsible for funneling several odd specimens [cf. Orthotrichum hainesiae) collected by her friends, Mrs. E. J. Spence and T. S. Brandegee, to bryological specialists (Flowers 1942).

Hall, Elihu (1820-1882). A farmer and amateur botanist in Athens, Illinois. Collected with Parry in Colorado in 1862.Also collected plants in Texas and Oregon.

Hermann. Frederick J. (1906-1987), received his doctorate at the University of Michigan. His career was spent at the United States Forest Service Herbarium, where his specialization was the genus Carex and Vicia. He retired to Fort Collins, Colorado, where he collected a fine series of bryophytes, especially from the Front Range and Rocky Mountain National Park. His career collections are at the University of Michigan, with many duplicates at the University of Colorado. See Voss & Reznicek (1988).

Holmen, Kjeld (1921-1974), Danish bryologist who discovered Oreas martiana on Mount Evans. Specialist in bryophytes of Greenland. See Warncke (1975).

Holzinger, J. M. (1853-1929), German-born Minnesotan, collected bryophytes in Colorado in Arapaho and Pike National Forests in 1896, and his Colorado collections contributed to his published exsiccati, Musci Acrocarpi Boreali-Americani.

Holzinger, Marie. Collected in Boulder County in 1892.

Jamieson, David, contemporary bryologist, professor at Fort Lewis College, Durango, and specialist in the moss flora of the southwestern San Juan Mountains. A student of Wilfred Schofield, Univ. of British Columbia, and monographer of the American species of Hygrohypnum.

Kiener, Walter B. (1894-1959), Swiss-born Colorado mountaineer, collected on Long's Peak (1938), where he was a climber and guide. His bryophyte and lichen collections were acquired by the University of Nebraska in 1960. Kiener was involved in a winter climb of Long's Peak on Jan. 10-12, 1925, in which his companion, Agnes Vaille, died in a fall.

Lehr, Paula, contemporary amateur botanist, collected bryophytes in the Gunnison Basin and Elk Mountains, and stimulated interest in mosses among amateurs and Forest Service staff in the local area.

Nelson, Lawrence T. (1862-1932). Primarily a mycologist, collected for the U.S.D.A. as a forest agent occupied in locating poisonous plants in Colorado and elsewhere. He collected a few mosses from the Gunnison area. Nelson taught at an extraordinary number of colleges and universities in many states. His bryophyte collections (1902-1910) are at Duke University. See Gier (1953).

Nelson, Patricia (1940-), contemporary amateur bryologist, Librarian at the University of Colorado Medical School in Denver, studied the mosses of Clear Creek Canyon, and for her master's thesis prepared a very useful Random Access Key to the Genera of Colorado Mosses. An amateur bryologist, she is credited with the discovery of the only collection of Leptodon smithii in North America.

Parry, Charles Christopher (1823-1890). "During that year [1865] Rocky Mountain mosses were received from Dr. Parry, who a few years before had named Colorado's Rocky Mountain peaks for Dr. Gray and Dr. Torrey)" (Rodgers 1940). Only Bartramia stricta was specifically attributed to Parry by Lesquereux & James (1884). See Weber (1997).

Porter, Thomas Conrad (1822-1901). Botanist and clergyman, was attached to the Hayden Surveys from1869-1874 ((Foster 1994). His moss collections were few and incidental to vascular plants. Professor at Lafayette College, Lancaster, Pa. What collections of his remain are at the Philadelphia Academy of Natural Science.

Rolston, Holmes, III. (***). Contemporary amateur bryologist, Distinguished Professor of Philosophy at Colorado State University, "father" of the concept of environmental ethics as a modern academic discipline. Collected bryophytes with Fred Hermann for several years, and is an excellent field bryologist.

Rothrock, Josiah Trimble (1839-1922). A student of Asa Gray, and a botanist and surgeon on the Wheeler Expedition. He was the supervisor of John Wolf. Many vascular plants bear his name; whether he really collected bryophytes is questionable.

Sayre, Geneva (1911-1992), Collected mosses in the Front Range and in Mesa Verde National Park and wrote but did not publish a Moss Flora of Colorado. Did extensive bibliographic work on cryptogamic exsiccati. See Pfister (1993).

Spence, Mrs. E. Jane. Reported as having collected at Manitou and vicinity in July, 1897 (Ewan 1981). Possibly the collector of Neckera douglasii?

Stiverson, Clare L. Bryology student, produced a Masters Thesis for the University of Denver in 1951: A preliminary survey of the mosses in the Mount Evans area, 45 pages. Some of her collections are in herbarium COLO. accession numbers B-9263-9284.

Wikel, Patricia nee Thomas (***). Collected hepatics in the Front Range and San Juan Mountains in the 1970s. Collections are at COLO.

Wolf, John (1820-1877). Member of the 1873 Wheeler expedition. Assistant to Rothrock. Several vascular plants (cf. Ribes wolfii) bear his name. It is questionable whether he had any real interest in bryophytes.

In 19*** the University of Colorado hosted a series of field trips during the first ICSEB meeting in Boulder, A large number of bryologists attended, including William C. Steere, Lewis Anderson,

Wet meadows on the eastern plains these very poor in mosses, but usually support Drepanocladus aduncus.

Carrs are willow-dominated wetlands common on mountain floodplains. These are not very productive of bryophytes because of erosive scouring by waster, and density of Carex stands. However, at the bases of willow shrubs, a characteristic cluster of moss species includes Climacium dendroides, Helodium blandowii, and Tomenthypnum nitens.

Colorado has no true bogs. What have been called bogs in Colorado are really fens. McQueen (1990) described the differences. We paraphrase some of these statements to apply to our mountainous region. Bogs are wetlands that are very poor in nutrients because most of the minerals are obtained from precipitation rather than from ground water. They are also referred to as ombrotrophic peatlands, meaning simply that all nutrients come strictly from precipitation. They are usually characterized by a pH of 4 or less. Sphagnum is the dominant vegetation of a bog.

Fens are very similar to bogs, but they receive their nutrients from the surrounding ground water and are wetter. Many of the same plants that are found in bogs are also found in fens. Fens are often referred to as minerotrophic peatlands because if the greater influx of nutrients from ground water The term fen may be somewhat misleading, because the category includes some diverse habitats. For example, the word fen is used to describe some alkaline wetlands that are dominated by sedges and grasses and very few peat mosses.

Poor fens generally obtain more water from precipitation than from run-off from elevated places, and their pH is higher (5 to 6.5). High Creek Fen, in South Park, may be so characterized. Rich fens occur in level areas into which water drains from higher altitudes and flows through and out although at a rather slow rate. Chattanooga Fen, in the San Juan Mountains, is an example.

The extensive willow/sedge meadows along the subalpine streams do not hold water for long periods and may become quite dry seasonally. The relatively meagre number of bryophytes are commonly restricted to the slightly raised bases of willow clumps.

Sites that are dominated by running water are the most varied habitats in the mountains, and are not easily characterized. They vary from slowly flowing or intermittent streamlets to rushing cataracts. Some very common species, such as Brachythecium rivulare Hypnum cupressiforme run the gamut, while certain species require some subtle characteristics of the stream that are not easily pin-pointed.

***

A great deal has been written about the moss vegetation of walls in Europe, much less in America. In Colorado, at least, our buildings are not very old, .and we lack the stone walls separating agricultural fields. Our dry climate does not encourage very much wall vegetation.

Nevertheless, wall mosses can be found here, especially on the buildings of the University of Colorado, where native stone, particularly the Lyons sandstone, has been used for facings. Mosses occur particularly on mortar in the cracks between the flagstone layers on east-facing walls. There are also retaining walls on the campus that support mosses. When these occur in areas that are shaded the cumber of species increases.

Some of the species that are common on mortar in horizontal cracks are: Bryum argenteum, Barbula convoluta, Ceratodon purpureus, Coscinodon calyptratus, Didymodon rigidulus, Grimmia anodon, G. pulvinata, Orthotrichum hallii, Ptychostomum angustifolium, Schistidium confertum, Syntrichia virescens, and Tortula muralis. Shaded retaining walls, in addition to supporting these species, have some species of more mesic preferences, such as Brachythecium erythrorrhizon, Rosulabryum flaccidum, Tortula atrovirens, and T. mucronifolia..

Lawns that are over-irrigated or poorly drained support mostly Brachythecium erythrorrhizon and Ceratodon purpureus. Sidewalk cracks, as they become eroded, are commonly filled with Bryum argenteum. For many years the University of Colorado had a system of shallow concrete ditches collected to a major irrigation canal. These ditches were used to flood the campus mall and are now no longer used and dry, but receive enough seepage moisture that their vertical sides are often plastered with mats of Amblystegium varium! A drain-pipe on the corner of Denison Building watered the surface at its base enough to support a nice colony of Gemmabryum subapiculatum until the Facilities Management people decided to cleanse the building of asbestos and built a temporary shack over the site. Most of the moss disappeared but is slowly coming back despite the lack of water!

Alkaline flats are seasonally wet stands. On the western slope these are commonly filled with greasewood (Sarcobatus vermiculatus). In the shade of these shrubs a few species of mosses, such as Crossidium and Syntrichia are frequent.

Gypsum/salt domes retain water longer than the flat desert pavements. The most notable one in Colorado is in Paradox Valley. The sypsum retains rainwater longer than the adjacent sandy flats, and is densely covered by a veriety of cruatose lichens. The moss flora is poor but especially interesting. Didymodon nevadensis occupies bare faces between the lichens, while Bryum nanoargenteum and Syntrichia caninervis occur in the shade of low Atriplex shrubs.

There are two wonderful things about bryophyte distributions which have nothing much to do with the bryophytes themselves. It is a boon to scientists that the Lord did not distribute bryophytes, or plants, for that matter, other plants, at random. If He or She did, we scientists would have a terrible time, because we would have to scour every inch of the planet. Bryophytes are tied very closely to their microhabitats, which are discrete and can be recognized by an astute observer. Furthermore, bryological collectors move at a snail's pace compared to wild flower people. We might walk a few miles in a day -- longer distances if we close our eyes until we reach the destination where we know we will find a particular microhabitat. Usually we tend to find an interesting spot at the roadside or only a few hundred yards away, and spend the day right there!

A corollary to the first blessing, we should be very thankful that man has built roads and trails, despite the fact that we tend to resent these intrusions on pristine habitats. But for these we might never find some of the rarer species. When looking at a dot distribution map of a bryophyte we must remember that this is where collections have been made. A dot map will give us a general idea of other localities, but few areas are so well collected that one can make detailed dot maps covering the smaller geographical divisions such as counties, townships, ranges, and sections. Thus it is not very meaningful to declare a bryophyte rare because few collections have been made. Few bryologists collect the same common moss in more than a few counties in a state. At the same time, because we have found a rare microhabitat does not mean that we have exhausted the possibility that other such places may still be found farther from the road.

Bryophytes often have much larger or more widely disjunct distribution patterns than do flowering plants. The Southern Rocky Mountains draws its flora from the obvious migration pathway afforded by the immense north-south extent of the Western American Cordillera. Every drainage system that radiates from the core of the system also serves as a highway of migration for riparian and lowland or desert species. An apt analogy is that of a great wheel, whose hub, the Southern Rockies, preserves the most ancient survivors, and whose axle and spokes--the Cordillera and the drainage systems, provide the pathways along which migration has taken place through time under the pressures of climatic shifts and orogenic movements.

The Colorado bryophyte flora is predominantly boreal-montane, as testified to by the fact that all but a very small number of species are common to Colorado and Scandinavia. A bryologist trained in Scandinavia or Siberia will be very comfortable in Colorado for there are so many species in common.

A significant number are restricted to protected ravines in the eastern slope of the Front Range, where they persist as relicts of the Pleistocene. These species of eastern woodlands (some having extensions to the Pacific Northwest) formerly radiated to the foot of the Rocky Mountains. Following the drying out of the high plains, they were eliminated from the intervening territory. Among these are Amphidium mougeotii, Anomodon spp., Brachythecium acuminatum, Cnestrum schistii, Dicranum brevifolium, D. flagellare, D. fulvum, D. montanum, Didymodon tectorum, Entodon cladorrhizans, Grimmia anomala, G. pilifera, Leskea polycarpa, Mnium hornum, Neckera complanata, Plagiothecium cavifolium, P. laetum, Platygyrium repens, Rhytidiadelphus triquetrus.

A few desert and lowland species are common to desert and steppe areas here and in Asia (Anoectangium handelii, Didymodon anserinocapitatus, Jaffueliobryum, Crossidium, Aloina, Pterygoneurum). A very small number of world-wide disjuncts can be considered to be ancient relicts of Tertiary or older floras (Bryoxiphium norvegicum, Gemmabryum alpinum, Leptodon smithii, Oreas martiana, Voitia nivalis). A very few species follow a North American Cordillera-Andean path ranging from Alaska through Colorado and down to southern South America (Anacolia, Bartramia potosica, Leptopterigynandrum, Rhexophyllum subnigrum, Syntrichia bartramii).

Of the boreal-montane element, the majority of our species belong to taxa of relatively dry mountain areas with continental climate. Species requiring constantly high humidity (suboceanic species) such as those of the forests of the Pacific Northwest are either absent here or they occur in small remnants of what might have been during the Pleistocene larger areas suited to their growth (, Hylocomiastrum pyrenaicum, Hylocomium splendens, Pleurozium schreberi, Ptilium crista-castrensis, Rhytidiadelphus triquetrus).

***Endemic Pacific Northwest: Brachythecium leibergii, Roellia roellii ...

Bryophytes are obviously closely tied to particular substrates, not only the general types of rock, but the features of rocks. having to do with exposure, seepage areas, crevices; tree and shrub habitats: the kinds of bark, the height from the ground, the exposure; soil types and conditions: soil types and conditions, whether heavy, light, alkaline, compacted or loose; the surrounding forest canopy. Others may come to mind.

Koponen (1982b) presents an interesting discussion concerning how bryophyte ecology may give insight to their taxonomy. He says "It is much easier to find examples of species which are strictly limited to a special micro-habitat than of species which grow in a large variety of habitats, such as Ceratodon purpureus." Habitat specificity often is correlated with the evolution and hence the taxonomy of the species. This is why it is so important to include habitat information with collections. When I was identifying, "site unseen" Donald McVean's Australian alpine collections, I sometimes suggested a name. "It can't be that; the habitat is all wrong!" He was right a hundred percent of the time. Koponen says. "I am also pondering why we should not accept a specific status for a taxon which differs from its nearest relative in several characters and also has a different ecology?" I feel that this question is pertinent to our consideration of the taxonomic position of the high-alpine "variety" of Aulacomnium palustre. This moss never grows with the species proper, its leaf shape is unique; it produces no gemmae, it is disjunct in high mountains of the Eastern hemisphere; it does not fruit; it conceals what stem rhizoids it possesses within the axils of the leaves; I have not encountered "intermediates."

Because of their physical nature, their existence as predominantly haploid organisms, and their close attachment to the very thin layer of substrate on which they grow and must survive, bryophytes, more than any other plants, are recognized as being associated, if not dependent on, various physical substrates, often having to do with the availability of water as actual liquid or as mist or dew; if liquid, whether the water is running or not, whether it is augmented by chemical ions coming down from a hillside or by runoff from mine tailings, etc., hat is, what the pH of the substrate is, acid or basic or neutral. Other characteristics of the substrates on and in which they grow are important. Bryophytes, at least many of them, are categorized as being calciphiles or basiphiles (associated with calcium or other substances including potash, similar in chemical behavior); disturbance plants, nitrophiles (requiring or tolerating nitrogenous material); dung-"loving" mosses; minerotrophic mosses.

Not very many mosses are well enough known for them to be categorized with any of these terms, and moreover, it is not understood what the roles of substrate really plays in the occurrence of mosses. Mosses that are considered calciphiles on one part of the world may not behave as such in another. Moss floras in Places like Scandinavia are more thoroughly classified than others because of a long history of interest in them and because calciferous rock associated with abundant moisture, makes calciferous substrates more obvious selectors of plant species. In the Rocky Mountains, however, we not only have had little serious interest in mosses, and the so-called calciphiles are often not definitely connected to calciferous substrates. Very little note has been made in Colorado concerning the calcium content of the substrate.

Here is a partial list of Colorado species that are considered by Crum (1973a) as being calciphiles, or likely calciphiles. A few more have been added by the present writers. Those indicated as (!Crum) we consider surprising, since in Colorado they do not necessarily behave as such.

Disturbance: Ceratodon, Bryum argenteum, B. angustifolium, Gemmabryum subapiculatum

Nitrophile: Ceratodon purpureus

Basiphile: Ditrichum flexicaule

Minerotrophic: Coscinodon cribrosus, Mielichhoferia

Fire ash: Funaria hygrometrica, Leptobryum

Dung: Splachnaceae (Tayloria, Splachnum, Voitia)

We used to think that the first snow would bring an end to our work in the field, because, we thought, once the mosses become covered with snow, there would be no use looking for them until May or June. At Hall Ranch and on the outwash colluvium around Boulder, we found that this is not necessarily the case.

In 1005 we have been very successful in finding mosses on what ordinarily would be considered extremely dry habitats at low altitudes. However, in the winter, snow cover is spotty, and ground mosses such as Tortula acaulon, can be very abundant locally in grasslands, their sporophytes perfectly formed although green, ready to pop when spring comes. On the Hall Ranch, Syntrichia calcicola and Grimmia longirostris are active in the sheltered gulches under the shrubs. On the rim-rock, Grimmia anodon bears young capsules, Didymodon rigidulus and D. anserinocapitatus are active, and in the shallow crevices of the rim-rock, Pterygoneurum subsessile, Encalypta vulgaris, and Weissia ligulifolia also are developing capsules. In the extremely deep, narrow crevices where there is a littl moisture, one can usually find Anoectangium handelii, as well as Fissidens sublimbatus.

This situation surely obtains in the canyons and plateaus of the Western Slope, but unfortunately we have not had the opportunity to visit these areas in the winter. A resident bryologist needs to explore the gypsum flats for Crossidium and Aloina, and the rim-rock for Entosthodon sonorae, Didymodon convoluta, Funaria americana, and other minute ephemeral species. There is much work to be done also on the high plains, where there is always a possibility of discovering Aschisma kansanum on the underside of quartz pebbles in the easternmost counties. The weather is always a problem in the winter and spring, and the midges can prove daunting.

(In preparation)

Ref: Hedenäs papers

Pleurocarpous mosses in fens generally are suggestive of Amblystegiaceae, but belong to diverse families. For ecologists working in fens, it is very useful to be able to identify the plants in the field. The following field key will make the job easier. Note: Leaf features refer to the stem leaves, not the branch leaves.

1a. Leaves broadly ovate, ovate or rounded-triangular, in upper part suddenly rounded-narrowed or apiculate (note: the inflexed upper leaf margins may give the apex a more a mucronate appearance) (2)

1b. Leaves straight or falcate, from ovate or triangular basal portion gradually narrowed towards leaf apex (5)

2a. Costa short, double or single, not or hardly visible with a hand lens; shoots turgid, slightly and irregularly branched; alar cells not hyaline, in an indistinctly delimited group. Pseudocalliergon turgescens

2b. Costa long, reaching leaf middle or further, single or branched, usually easily seen with a hand lens (3)

3a. Leaves usually ovate or narrowly ovate; plants of alpine pools. (4)

3b. Leaves usually broadly ovate or broadly rounded-triangular; green or brownish-green species, sometimes with a pale pinkish hue. Calliergon

4a. Red colors common, otherwise green to dark green; usually strongly branched; stem leaf apex usually distinctly apiculate, at least in young leaves; leaf point often bent inwards; very rarely found with leaf-borne rhizoids. Warnstorfia sarmentosa

4b. Pale- or yellow-green, usually sparsely branched; leaf apex rounded or rounded-obtuse; leaf-borne rhizoids common near the leaf tips. Straminergon stramineum

5a. Costa short, usually double, not or hardly visible with a hand lens (6)

5b. Costa long, reaching middle of leaf or further, normally single, usually easily seen with a hand lens (8)

6a. Very large species; shoots often turgid, not flattened; green or often with brown, yellow-brown, red, or blackish colors. Scorpidium scorpioides

6b. Medium sized species; shoots often somewhat flattened; green or pale to yellow-green species (7)

7a. Leaves with large and distinct groups of alar cells; costa short and forked, or lacking. leaves strongly falcate Calliergonella (See Hypnaceae)

7b. Leaves with small and indistinct groups of alar cells; costa very short and inconspicuous; leaves weakly falcate. Breidleria pratensis (See Hypnaceae)

8a. Leaves straight, lanceolate, strongly plicate (9)

8b. Not as above (10)

9a. Underside of the straight stems densely clothed with brown tomentum. Tomentypnum

9b. Underside of stems not tomentose. Brachythecium turgidum

10a. Stem leaves usually more or less broadly triangular to very broadly cordate, rather quickly narrowed to the acumen; branch leaves much smaller, falcate; alar groups large, triangular and well-delimited, reaching from margin to costa; plants usually densely pinnate; leaves not plicate. Cratoneuron filicinum

10b. Leaves rounded-triangular, ovate or broadly ovate to linear, more gradually narrowed toward the apex; alar groups usually less distinct; branching rarely densely pinnate (tomentum may occur in Palustriella falcata, Tomentypnum, and Conardia). (11)

11a. Paraphyllia present (tear off a few leaves; the paraphyllia are usually distinctly visible with a hand lens); large species with falcate, distinctly plicate leaves; costa strong; tomentum frequently present. Palustriella falcata

11b. Paraphyllia absent; tomentum lacking except in Tomentypnum (12)

12a. Leaves conspicuously plicate; commonly fruiting, the capsules normally horizontal (not common in fens but very common in wet spruce forests). Sanionia uncinata

12b. Leaves not plicate or indistinctly so (13)

13a. Leaves from more or less straight and rather erect basal portion with rather strongly curved upper part; alar groups undifferentiated or very small and not visible in the field. (14)

13b. Leaves curved more or less along their entire length, or nearly straight; alar groups distinctly differentiated and usually visible on torn off leaves (indistinctly differentiated in Pseudocalliergon angustifolium) (15)

14a. Shoots relatively stiff and more regularly branched; green, yellow-green, brown, or brown-red (a unique color combination in contrast to the next); leaves green or yellow-green with brown-red costa (often also leaf base), somewhat dull (due to relatively short leaf cells with squarish ends). Scorpidium cossonii

14b. Shoots somewhat larger and less branched; red, blackish red (brown-red) or green; glossy (due to long cells with gradually narrowed ends. Scorpidium revolvens

15a. Costa long-excurrent; shoot and branch apices sometimes pencil-like; usually growing submerged in pools (16)

15b. Costa not excurrent -- in case of doubt (occurring in species with radially branched shoots) the shoot and branch apices are not pencil-like; submerged or not (17)

16a. Shoot and branch apices pencil-like; leaves straight or curved, often deep red-purple; young axillary hairs long and brown (visible with a hand lens if a few leaves are torn off); shoots radially branched. Warnstorfia trichophylla

16b. Shoot and branch apices never pencil-like; leaves usually curved, never becoming red; young axillary hairs small, hyaline (not visible with a hand lens); shoots distichously branched. Drepanocladus longifolius

17a. Yellow-brown, brownish-yellow, or green species, typically with a golden gloss when dry; alar cells indistinctly differentiated; plants of strongly calcareous habitats. Pseudocalliergon angustifolium

17b. Color varying, not with golden gloss when dry; alar groups usually large and more or less distinctly differentiated; in less strongly calcareous sites (18)

18a. Shoots distichously branched, never red; rhizoids never growing from the leaves; mostly in nutrient-rich habitats (19)

18b. Shoots more or less radially branched, most distinct when growing with stem in vertical position; sometimes with red coloration, and sometimes with rhizoids growing from the leaves; in less nutrient-rich situations. Warnstorfia

19a. Plant habit "Drepanocladus-like", that is, with leaves falcate-secund to sometimes straight and erect; leaf acumina in straight-leaved plants plane or at most slightly furrowed. Drepanocladus aduncus

19b. Plant habit "Campylium-like", that is, with leaves from straight and erect bases usually with leaf acumina more or less spreading or squarrose; leaf acumen furrowed. Drepanocladus polygamus

Aquatic pleurocarpous mosses that grow over seeping rocks or attached to rocks in small streams may belong to genera of several families. Brachythecium rivulare may be mistaken for Amblystegium, but the leaves are more broadly ovate and finely serrulate distally, and the alar group is conspicuously inflated.

la. Alar cells inflated, sharply differentiated; paraphyllia usually present; leaf cells more or less papillose (from protruding cell ends). Cratoneuron and Palustriella

lb. Alar cells various; paraphyllia absent; leaf cells smooth or nearly so (2))

2a. Leaves usually squarrose. Campylium and relatives (See Campyliaceae)

2b. Leaves never squarrose (3)

3a. Leaves with a single costa extending at least to mid-leaf (4)

3b. Leaves lacking a costa, or the costa short and double (8)

4a. Leaves cordate-ovate, oblong-ovate, or oblong, never long-acuminate; margins entire. Calliergon and its relatives

4b. Leaves always distinctly acuminate; margins either entire or serrulate (5)

5a. Leaves usually falcate (except in some submerged forms) or plicate, or both. Drepanocladus and its relatives

5b. Leaves neither distinctly falcate nor plicate (6)

6a. Costa strong, percurrent or excurrent; plants submerged in running water. Amblystegium

6b. Costa shorter; plants terrestrial or in wet places, rarely submerged (7)

7a. Leaves very small, 2 mm or less long; plants small and delicate; plants of forests (8)

7b. Leaves larger; plants rather coarse; plants of wetlands (9)

8a. Leaves and branches very small and slender, leaves up to 0.45 mm long, ovate-lanceolate, never squarrose, entire or often serrulate at the base; 2-3-celled gemmae present in the leaf axils. Platydictya (see Hypnaceae)

8b. Leaves larger, otherwise not as above. Amblystegium

9a. Leaves small or large, less than twice as long as wide or, if longer, then somewhat falcate; basal cells rarely with pitted walls; on wet rocks in or near streams. Hygrohypnum

9b. Leaves usually 2 mm long or longer, commonly twice as long as wide or more, deeply concave, never falcate; walls of basal cells often pitted; not attached to rocks, but either submerged in fen ponds or in loose gravel of snow-melt rills (10)

10a. Leaves more or less falcate-secund, at least those of the tips of the stems and branches; alar cells hyaline, thin-walled and inflated, in small, inconspicuous groups. Scorpidium

10b. Leaves loosely imbricate to spreading, not at all or at most very slightly secund; alar cells shortly oblong and subquadrate, neither hyaline, thin-walled, nor inflated. Pseudocalliergon

1a. Leaves less than 1 mm long, generally serrulate for at least part of their length; basal marginal cells and adjoining alar cells subquadrate or short-rectangular to transversely elongate, firm-walled. A. serpens

1b. Leavesw more than 1 mm long, entire; basal marinal cells short- to rather long-rectangular, the adjoining alar cells quadrate to rectangular and rather lax. A. riparium

A. riparium (Hedwig) Bruch & Schimper. Common in very wet places, particularly in areas where livestock or other pastoral activity is strongly evident. I have found it abundant in drinking troughs, on check-dam sluices and in relatively still water through the middle altitudes.

In our distribution of specimens, we made a horrendous mistake of thinking a large, robust, submerged population in the bed of an intermittent stream was Fontinalis hypnoides. Evidently we were not alone. Crum & Anderson, p. 994, write: "Amblystegium laxirete represents an extreme development of a Fontinalis-like habit. It sometimes occurs in long, streaming masses in swiftly flowing waters. . . .In this habitat, the plans are striking, but in less vigorously flowing waters they are less distinctive." A number of other named forms evidently are variants of this extraordinary modification. Moral: Don't assume that everything that looks like Fontinalis, is Fontinalis!

A. serpens (Hedwig) Bruch & Schimper var. juratzkanum (Schimper) Rau & Hervey. A small, nondescript species, usually fruiting. The species is generally distributed on wet boulders, soil, bases of saplings or shrubs, exposed roots, rotten logs, etc., from the foothills up to the subalpine. The leaves wide-spreading, about 0.6 mm long and gracefully acuminate, with costa to about mid-leaf. The median leaf cells are about 6:1 or 35m x 5-7m, smooth and moderately thick-walled. The leaf margin is slightly denticulate from projecting distal ends of the cells. The alar region is quite clearly differentiated, consisting of quadrate and broader cells forming a more or less triangular patch at the basal angles. The capsule is curved, the operculum conic, and the urn strongly constricted below the mouth. We are inclined to follow Nyholm in treating A. juratzkanum as a variety of A. serpens. The variety is distinguished by having the leaves more widely spreading and with the marginal basal cells rectangular instead of quadrate.

A. varium (Hedwig) Lindberg. An exceedingly variable species of running water in the lower altitudes, having had many forms recognized as separate although indistinct species, place either in Amblystegium or Hygroamblystegium. Crum and Anderson (1981) amply show the frustrations inherent upon separating them. Hygroamblystegium has been a terrible problem because of its variability. Recently. molecular biology has come to the rescue (see Vanderpoorten 2004), in which the author demonstrates that H. fluviatile, H. humile, H. noterophilum, and H. tenax are justifiably synonymized under Amblystegium varium.

Calliergon is generally not a plant of the edges of swiftly flowing brooks. C. richardsonii grows submerged in still pools. C. cordifolium occurs on saturated, swampy ground in forest clearings filled with tall willows, C. giganteum was found on sloping rock faces beside a quiet backwater on a level bench of a mountain stream. A few species formerly included in Calliergon are common in alpine pools. These include Warnstorfia sarmentosa and Straminergon stramineum.

1a. Costa usually ending well below the leaf apex, with short branches or forked at the apex; shoots with rather long, thick branches, more sparsely branched and with branch leaves more erect or imbricate than in C. giganteum (see below). C. richardsonii

1b. Costa ending almost in the leaf apex (appearing to reach the apex as seen with a hand lens) (2)

2a. Costa strong; alar groups of stem leaves large, triangular and sharply delimited from surrounding cells, extending from leaf margin to or almost to the costa; leaves broadly triangular (shoots when well-developed densely branched like a spruce tree, with more or less spreading branch leaves except near branch apices. C. giganteum

2b. Costa weaker; alar groups similar but diffusely limited from surrounding cells; rarely found in permanently submerged habitats). C. cordifolium

C. cordifolium (Hedwig) Kindberg. Pools and lakeside swamps, upper montane and subalpine. Our records are from the plateaus of western Colorado and fens in the Front Range.

C. giganteum (Schimper) Kindberg. We have two collections: Larimer Co.: Cirque Meadows trail, vic. Pingree Park, streamlet tributary of Fall Creek, 9,600 ft., Hermann & Rolston 80114. Same locality, Weber & Wittmann 112731.

C. richardsonii (Mitten) Kindberg. Submerged in fens, Boulder, Larimer, and Clear Creek counties. The Boulder County specimens are extremely large-leaved, resembling C. megalophyllum (so reported by Weber, 1973).

C. filicinum (Hedwig) Spruce. This species is abundant in wet sites in the subalpine forests and willow carrs and fens. The plants are small, pinnately-branched, and the stem leaves are conspicuously larger than the branch leaves. The branch leaves are usually falcate. A characteristic feature are the paraphyllia, of various shapes and sizes, minutely leaflike, which unfortunately vary in abundance and are sometimes almost lacking. The stem leaves are broadly ovate, witrh a very strong costa. The median laminal cells are narrowly rhomboid, an the alar cells are enlarged, in a conspicuous triangular group. The species is said to be strongly calciphilous, and is common in calcareous fens in the subalpine.

Crum and Anderson accept Drepanocladus and Scorpidium in a traditional sense, which for field botanists seems reasonable, and they reject the separation of several other species into the genera Limprichtia, Warnstorfia, and Scorpidium. However, we follow the recent works of Hedenäs.

Hedenäs suggests one way to distinguish Drepanocladus from Warnstorfia: "It is always good to look for rhizoidal initials near the leaf apices. If you study 10-15 leaves (or a shoot apex with several leaves left) in the microscope, there are almost always at least some leaves with such initials in Warnstorfia species, but never in Drepanocladus s. str."

1a. Leaves from more or less straight and erect bases, usually with the acumen more or less spreading or squarrose; leaf acumen furrowed; plants more golden-brown than green. D. polygamus

1b. Leaves strongly falcate-secund; leaf acumen in less falcate-leaved plants plane or almost slightly furrowed; plants green with no other tints (2)

2a. Costa of stem leaves ending well below the leaf apex; leaf margin entire, or only occasionally very finely denticulate. D. aduncus

2b. Costa of stem leaves excurrent, rarely ending a few cells below leaf apex; one or both leaf margins usually partly finely denticulate. D. longifolius

Drepanocladus aduncus (Hedwig) Warnstorf. An extremely common and variable species, very wide-ranging in altitude from the plains up to the alpine. The elongate, inflated alar cells in a row continuing almost to the costa, is diagnostic. The plant is weak, not standing erect, yellowish-green, never with reddish or brown colors, very little branched and then not at all pinnate; the costa is slender and elongate. However, some collectors have confused this with Calliergonella (was Hypnum) lindbergii. The latter stands stiffly erect in close order, the leaves are very broad at the base (triangular-ovate) and with large alar cells at the basal angles just where the leaf becomes decurrent. The costa is absent or short and double, and faint, and the leaf margins tend to curve inward near the apex.

Drepanocladus longifolius (Mitten) Paris. In subalpine pools. We have one excellent collection: Grand Co.: Rocky Mt. Nat. Park: Floating on pond along Ute Trail south of Lake Irene, 10,400 ft., Hermann 26506. Hedenäs (BFNA) accepts this for Colorado, saying: "D. longifolius [D. capillifolius of authors] differs from all other American Drepanocladus species in its excurrent leaf costa. Because of the latter it could hardly be confused with any earlier Drepanocladus s. l. in North America except Warnstorfia trichophylla. However, the leaves are always green, there are never any rhizoidal initials, and the shoots are distichously pinnately branched. In Warnstorfia trichophylla the shoots are radially branched, and the shoots and branch apices pencil-like, the leaf margins are more strongly denticulate than in D. longifolius; it frequently becomes red when emergent whereas D. longifolius never gets red, and the axillary hairs consist of 1-7 early brown upper cells (1-2 elongate hyaline cells in D. longifolius).

Drepanocladus polygamus (Bruch & Schimper) Hedenäs. One record, from a subalpine fen on Diamond Lake, in the Front Range, collected by Hermann (!Lawton). See Flowers, plate 119:9-11. (Campyliadelphus, Campylium, Drepanocladus). This has until recently been placed in Campylium and indeed has more the aspect of that genus. The plant is definitely stouter and the leaves less falcate than D. aduncus and has more brownish tints. D. polygamous is autoicous, while D. aduncus is dioicous but this is difficult to determine without capsules.

Ref.: Jamieson (1986a)

*** Add H. styriacum (Limpricht) Brotherus Check also (11). Have we left out something?

1a. Stem cross-section with an epidermis of enlarged, fragile, thin-walled cells, the inner cortical cells in 3-4 layers, thick-walled, the core cells large and thin-walled. H. ochraceum

1b. Stem cross-section with an epidermis of several layers of small, thick-walled cells, the outer layer not fragile nor thin-walled (2)

2a. Leaves all straight, but sometimes somewhat secund (3)

2b. Leaves falcate (or some leaves on different branches falcate and/or straight). (11)

3a. Leaves broadly ovate to orbicular (4)

3b. Leaves ovate to oblong-ovate or ovate-lanceolate (8)

4a. Median marginal leaf cells 60 m long or more. H. bestii

4b. Median marginal leaf cells rarely longer than 55 m (5)

5a. Alar cells clearly differentiated, either thin-walled or incrassate. H. duriusculum

5b. Alar cells undifferentiated or formed of a few quadrate or short-rectangular cells which are incrassate or thin-walled (6)

6a. Costa usually single, stout, extending to mid-leaf or slightly beyond, sometimes forked or short and double; plants very coarse and rigid. H. smithii

6b. Costa almost always short and double; if single, the costa slender and the plants soft and pliable (7)

7a. Leaves deeply concave to cochleariform, usually 0.8-1.2 mm long, the apex tapering to an obtuse or broadly rounded tip; inner perichaetial leaf margins entire and recurved; endostomal cilia 2 or 3. H. smithii

7b. Leaves concave but never cochleariform, usually 1,0-1,7 mm long, the apex tapering to an acute but blunt point; inner perichaetial leaf margins coarsely denticulate and plane; endostomal cilia rudimentary or absent. H. molle

8a. Alar cells clearly differentiated, either inflated and mostly thin-walled, or smaller, incrassate and quadrate to short-rectangular. H. luridum

8b. Alar cells undifferentiated or with but a few quadrate to short-rectangular cells which do not form a recognizable group (9)

9a. Costa predominantly single to mid-leaf or beyond, sometimes short and double (leaf apex acute, plants coarse). H. smithii

9b. Costa usually short and double, rarely single to mid-leaf (margin of leaf apex uneven to denticulate). H. molle

Hygrohypnum is found attached firmly to rocks which are inundated by water of swift-flowing streams or at least wet periodically by spray. They are often coppery in color. Species with falcate leaves might be mistaken for Drepanocladus but lack the prominent percurrent costa. Drepanocladus does not occur firmly attached to rocks. H. bestii and H. dilatatum might be mistaken for Scorpidium turgescens but the Scorpidium likewise is not firmly attached to rocks and grows in snow-melt rills (see key also). Unfortunately we do not understand the subtleties of the microhabitats for these species or we probably could separate them quite nicely on their preferences. They probably do not occur together in the field. Taxonomically they are very difficult if one considers relationships of the species over a broader range. Quite possibly there are more species in our area than we recognize at present.

H. bestii (Renauld & Bryhn) Holzinger ex Brotherus. "Of the broad-leaved species, H. bestii is distinguished by the very long marginal leaf cells, which range from 60 to 250m, the large leaves which reach 3 mm long x 2 mm wide, and the dioecious sexuality. In many instances, discoloration in the basal leaf cells imparts the appearance of a radiating 'sunburst' in the leaf base which is typical of the species" (Jamieson, unpublished thesis).

H. cochlearifolium (Venturi in DeNotaris) Brotherus. "A soft species; leaves very small, deeply concave, often with margins explanate or more or less recurved. Often gets saturated with mud; much smaller than H. smithii, not at all falcate, alar cells not inflated, poorly developed." (Jamieson, voce). Collections from San Juans (Spencer Basin) and La Plata Mts. (Owen Basin), Mount Evans (Vaarama and Weber).

H. duriusculum (De Notaris) Jamieson. "Under the microscope, the species can be recognized best by its usually oblong-elliptic to broadly ovate leaves and the well-defined group of thick-walled, usually discolored, quadrate, short-rectangular or slightly irregular alar cells." Jamieson, thesis (H. dilatatum).

H. luridum (Hedwig) Jennings. I find H. luridum and H. ochraceum difficult to distinguish without making stem sections, and of course this is impossible to do in the field. Both species have parallel green or reddish, straight or falcate-leaved, imbricate or spreading-leaved forms, and there seems to be a wide range in leaf size. There is also speculation that the outer cortical cells may be showing differences in response to environmental changes, in which case the species may not be separable. "An exceedingly variable species. . . .The species varies virtually continuously in nine or ten features, which have been used singly or in various combinations as criteria for the recognition for numerous subspecific taxa." (Jamieson, thesis). "The alar cells are enlarged, "blistered", more well-developed than in H. luridum, the stems are usually more or less julaceous an may be falcate. Fruiting material: the capsule of H. luridum has a persistent annulus; in H. styriacum it is deciduous." (Jamieson, voce). Good luck!

H. molle (Hedwig) Loeske. "H. molle may be distinguished from other species in the genus by the broadly ovate, concave leaves, which generally taper into an acute but blunt point and often denticulate apex, the undifferentiated alar cells, and the inner perichaetial leaves in which the cells on the abaxial surface of the leaf apex are prorulate" (Jamieson, thesis). For distinctions between this and H. luridum, see chart on thesis, p. 186. Jamieson suggests we ignore the discussion in Crum and Anderson.

H. ochraceum (Turner ex Wilson) Loeske. A very common moss on wet rocks in subalpine and alpine rivulets. ". . .a polymorphic, yet very distinctive species, which may be distinguished. . . by the dioicous sexuality, the outer layer of inflated cortical stem cells, the variable costa, and the nature of the alar cells." (Jamieson, thesis).

H. smithii (Swartz) Brotherus. "May be recognized by its coarse, rigid habit and the usually broadly ovate to orbicular, loosely imbricated to spreading leaves and the stout, generally single costa." (Jamieson, thesis). This is a rare species occurring in high, wet tundra (H. cochlearifolium).

H. styriacum (Limpricht) Loeske. We have one collection, from Clear Creek Co.: Summit of Loveland Pass, at inlet of small pond called "Summit Lake", 3,600 msm, Weber 111133-34 (verified by Jamieson, 2002). "H. syriacum looks like a 'weird' luridum; the leaf tip is flexed sideways, the leaf base is bowed out more than in other species, the leaves are triangular ovate, widest just above the base. there are few alar cells" (Jamieson, voce).

Ref.: Ochyra (1989)

Palustriella falcata (Bridel) Hedenäs. A moss of cold running water of streamlets in the alpine and subalpine (Cratoneuron falcatum). We had been calling this C. commutatum, but the species does not occur in America. Stems pinnately branched; leaves falcate-secund, plicate, broadly triangular-ovate, abruptly narrowed at the base, the cells linear; paraphyllia abundant (strip leaves from the stem), linear, 2-3-cells wide; rhizoids red, branched, with thick cell walls; alar cells not differentiated but a few rows of basal cells inflated or much enlarged.

Ref: Hedenäs (1990a)

P. angustifolium Hedenäs. Fairly common in subalpine wetlands. This is a Drepanocladus-like plant with essentially straight leaves, little branched and lax, the leaves gradually long-attenuate with very slender apex; costa ending below the apex. The alar cells are not very inflated but rectangular, with thickened walls which turn yellow; a band of short, almost quadrate cells run across the leaf base. In Scandinavia the plant is usually in calcium-rich wetlands. It is commonly confused with Drepanocladus aduncus, which is always clear green, never brownish, reddish, or yellow.

P. turgescens (T. Jensen) Loeske. A large, sparingly branched, flaccid, brown moss very slightly falcate and with broad usually obtuse, concave leaves. The terminal leaves of the year are yellow-green. Commonly found floating in rich willow-peat fens and forming short, dense mats in tundra snow-melt rills, South Park and vicinity. See Crum & Anderson (1981) vol.2, p. 994-997 for excellent plates and discussion. The species is sterile here. Hedenäs (1989, 1990) feels that this does not belong to either Scorpidium or Calliergon and suggests that its proper designation is Pseudocalliergon turgescens. In Scandinavia it is considered a calciphile. See Hedenäs (2002). He says: "The species is always sterile here. . . ."The easily detached apical shoot buds, a means of vegetative reproduction, probably partly make up for the low sporophyte production."

Ref.: Hedenäs (1989c)

Sanionia uncinata (Hedwig) Loeske. Very common, the only species found in relatively dry sites in the montane and subalpine forests, at the bases of trees and shrubs, and up to the tundra, but not in the wetter fen sites (Drepanocladus uncinatus). The clearly plicate leaves suggest Hypnum revolutum, and care must be exercised not to mistake the leaf folds of that species for a costa.

Ref.: Hedenäs (1989a)

1a. Large species (stem leaves 0.7-2.4 mm wide); stem leaves strongly concave, almost orbicular and obtuse to acuminate, or from broadly ovate to broadly ovate-lanceolate base narrowed to apiculate, acute, or acuminate point, falcate or (rarely) straight; costa usually double, more rarely single or lacking, rarely reaching above mid-leaf; hyalodermis of stem often incomplete. S. scorpioides

1b. Small species (stem leaves 0.45-1.09 mm wide); stem leaves concave, from ovate to ovate-lanceolate base narrowed to shortly or longly acuminate apex, falcate; costa single, ending in upper half of leaf; hyalodermis of stem complete. (2)

2a. Mid-leaf cells (of stem leaves) 14-95(-120) microns long, with squared to shortly fusiformly narrowed stem cells. Dioicous; outer peristome layer of exostome predominantly (>70%) dotted in lower part. S. cossonii

2b. Mid-leaf cells 61-140(-178) microns long, with shortly to long fusiform cell ends. Autoicous; outer peristome layer usually predominantly (40-50%) cross-striolate in lower part. S. revolvens

S. cossonii (Schimper) Hedenäs. This resembles somewhat S. revolvens, but is a smaller plant without the deep golden-brown color. One might mistake it at first for a husky Drepanocladus aduncus, but the color is variable on the same stem (in aduncus always green), and the stem is stouter, the branching rather stiffly spreading at right angles (never so in Drepanocladus) the leaves are very strongly circinate, and the color of the leaves varies from young to old from yellowish-green through pink to darker reddish. We have collections from many counties, in subalpine fens. With a little practice it can be distinguished easily in the field without a lens. One caution: when the leaves are stripped from the stem, often some of the stem epidermis comes along with them. The stem cells are inflated-rectangular, and often red-brown. They may be mistaken for alar cells except for the fact that they begin where the leaf base (as measured by the costa base) ends. The leaf base is also slightly decurrent alongside these stem cells.

A Drepanocladus-like pleurocarp with distinct pinnate branching, closely ranked falcate-circinate leaves forming prostrate mats. The color alone is enough to separate it from D. aduncus, a clear green moss for which it has been repeatedly mistaken. The Scorpidium is not green, but has a variety of brown and reddish tints. The stem is stout and brown here, but weak and slender in the Drepanocladus. In size the leaves are closer to the Drepanocladus than to Scorpidium revolvens, and the matte appearance according to Hedenäs may be due to the broader, less long-pointed ends of the cells of the glossy S. revolvens. The species is not confined to fens but evidently is common enough in stream-side situations. The same may be said of the Scorpidium. It seems amazing that this common plant has only recently been recognized in America.

S. revolvens (Swartz ex Anon.) Rubers. A beautiful dark blackish-copper-colored species of alpine fens and tundra pools, not abundant (Drepanocladus revolvens). The leaves commonly are quite circinate, much more so than in S. cossonii, with the narrow tip curved back again. The alar cells are very few and small. The terminal branches show the leaves completely curved into a circle. With experience, the shape and size of the median leaf cells is critical. In S. revolvens the leaf cells are so long and so narrow that they are difficult to measure, and the ends of the cells are acute. In S. cossonii the leaf cells are elongate but their borders are easily discerned, and the end walls are more blunt. S. cossonii is definitely a smaller plant and more pinnate. See Hedenäs (1989), for discussion of this and Scorpidium cossonii (dioicous); S. revolvens is said to be autoicous).

S. scorpioides (Hedwig) Limpricht. Floating in pools in calcareous subalpine fens, South Park and Guanella Pass. A gigantic water moss, with very soft and pliant, densely foliate stems with short pinnate branches. No other aquatic moss looks anything like this. The plant is black except for the very apices of the leaves exposed above the water level. The leaves are short, very convex, and falcate-secund, The stems reach up to more than a decimeter long.

Straminergon stramineum (Bridel) Hedenäs. Common in very wet ground along the edges of tundra pools and in shallow water, upper subalpine and alpine. When growing with Sarmentypnum, it characteristically is light green in color and has fairly narrow leaves with rounded tips (Calliergon stramineum). The leaf tips often contain one or more colorless cells that can give rise to brown rhizoids.

1a. Leaves ovate to narrowly ovate, not falcate, the upper part suddenly narrowed to a rounded-apiculate apex, often bent inwards over the leaf. W. sarmentosa

1b. Leaves gradually narrowed to the apex, usually falcate (2)

2a. Shoots distichously branched, never red; rhizoids never growing from the leaves; mostly in nutrient-rich habitats. See Drepanocladus

2b. Shoots more or less radially branched, most distinct when growing with stem in vertical position; sometimes with red coloration, and sometimes with rhizoids growing from the leaves; in less nutrient-rich situations. Warnstorfia (3)

3a. Costa strongly excurrent; shoot and branch apices sometimes pencil-like; leaves straight or curved. W. trichophylla

3b. Costa not excurrent; shoot and branch apices not pencil-like (4)

4a. Alar groups mostly large, triangular and well-delimited; costa rather strong; green or partly to entirely red to blackish-red plants. W. exannulata

4b. Alar groups either large, triangular and well-delimited, or indistinct and more or less ovate; costa weak or strong; plants usually green to brownish, hardly ever red, but sometimes reddish brown. W. fluitans

W. exannulata (Bruch & Schimper) Loeske. Common in still water of subalpine and tundra pools. The plants are usually brown, purplish or blackish. The curvature of the leaves is usually pronounced, but may vary. The leaves do not have an excurrent costa, and are distinctly serrulate especially near the apex. The alar cells are inflated, rectangular, and form a triangular patch that reaches the costa.

W. fluitans (Hedwig) Loeske. Very common in fens and still water of ponds, especially in the San Juan Mts. Nyholm stated: "When well-developed, this species is easily known and distinguished from W. exannulata. . . by its stem leaves mostly slightly decurrent and distinctly narrowed towards the base, a narrow plano-convex costa reaching about half-way up the leaf, slightly wider leaf cells, longer seta and larger spores."

W. sarmentosa (Wahlenberg) Hedenäs. Very common in tundra pools, where it forms floating or submerged purple-red masses along with Straminergon. Formerly Calliergon and Sarmentypnum.

W. trichophylla (Warnstorf) Tuomikoski & Koponen. This occurs in the fens of Guanella Pass. Most other reports have been considered to be misidentifications. The leaves have a long, excurrent costa, the alar cells are large (sausage-shaped), and the leaf margins denticulate. It is a beautiful red-purple moss, and the leaves are extraordinarily long and tapering to a needle-point. Our only collection is Clear Creek Co.: Guanella Pass, 3550 msm, just below the pass along trail to Mt. Bierstadt; in shallow water on margin of the largest pool at the base of the slope beside the boardwalk, 18 June 2000, Weber & Wittmann 110855.

January 8, 2001

Ref: B. Murray (1987).

This family is characterized by the peculiar dehiscence of the capsule, which splits into four valves with the valves remaining attached at the top and bottom, elastically constricting the capsule in the manner of a paper-lantern. The sporophyte parts are not strictly homologous to those in other mosses, so that this family forms a separate class of mosses. Andreaea is a very slender-stemmed moss growing in dense tufts on granite rocks, usually above timberline. The stems are more slender than most species of Grimmia, and usually there are at least a few of the characteristic capsules present. Didymodon subandreaeoides Kindberg, a very similar moss that forms very dense, wide "turfs" on limestone terraces and is never fertile can be easily mistaken for Andreaea.

The genus Andreaea and the family Andreaeaceae are named for Johann Gerhard Reinhard Andreae (1724-1793). He was born in Hannover, the son of a pharmacist. After training in a Frankfurt pharmacy, he studied in Leiden and England. Returning to Hannover, he took over his father's pharmacy. He did field work in Switzerland and became interested in chemistry and mineralogy, describing 300 types of soils. Besides natural history, he read great literature in various languages, especially loved the English poets, and was a fine pianist. Friedrich Ehrhart, with whom he worked, named this genus after him (Frahm 2001).

1a. Costa lacking; leaves lanceolate. A. rupestris

1b. Costa present; leaves subulate. A. heinemannii

A. heinemannii Hampe & Müller Hal. Very rare, on granite boulders on the highest peaks, our records from Mount Evans and Quandary Peak. Easily distinguished in the field by the loosely spreading, acuminate leaves. The costa is weak, flattened above, 4-cell-layered and often absent in the base of the leaf. Murray states: "Andreaea heinemannii is easily identified by its small size, untidy look due to divergent leaf tips, obtuse leaf apices, the more or less flattened subula and the costa often very weak or lacking basally."

A. rupestris Hedwig. More frequent than the last, but always a welcome discovery. Although most collections are from alpine tundra, one locality in Grand County was from boulders along a stream in subalpine spruce forest.

1a. Leaves ending in a short or long, hyaline hair point; leaf margins revolute. A. rostratus

1b. Leaves not ending in a hair-point; leaves acute or merely apiculate; leaf margins plane. A. attenuatus

A. attenuatus (Hedwig) Hübener. In addition to the merely acute or just apiculate leaf apices, this differs from the next by having coarser, conical papillae We have a very few collections, all from the outer foothills, where the plant forms extensive carpets over the downslope edges of granite boulders just above the high-water marks of Boulder and South St. Vrain creeks at about 6,500 feet altitude.

A. rostratus (Hedwig) Schimper. Our few collections are from sandstone cliffs in the southern and western plateau and canyon country at low altitudes: Montrose Co.: foothills of La Sal Mts., near Buckeye Reservoir, west end of Paradox Valley, 30 May 1960, Weber B-5547 (5-6,000 ft.).; Las Anmas Co.: Along Purgatoire River, T30S R59W, at spring on cool, N-facing slope of the canyon, with Populus tremuloides, 5 Sept. 1983, Cooper B-85560. The stems when dry have the leaves appressed, more or less catenulate; when wet, they are widely spreading, and the stems are very densely foliate. The leaves are multipapillose with very fine, sharp papillae, easily seen along the recurved margins of the leaves. The costa is very conspicuous, and appears sunken; the cells are elongate and lack papillae.

July 18, 2004

1a. Leaves small, mostly less than 1.5 mm long; basal leaf cells green, not distinctly different from the upper ones, unistratose; stems characteristically with a terminal pseudopodium bearing a spherical cluster of few-celled, fusiform gemmae; habitat on rotting, often charred, wood. A. androgynum

1b. Leaves larger, 2-4 mm long; basal leaf cells often enlarged, bistratose; pseudopodia, when present, naked or bearing relatively few green, leaf-like, many-celled gemmae in an erect flabelliform cluster; terrestrial (2)

2a. Leaves often yellow-green, variously spreading, contorted when dry. Stems matted together by conspicuous masses of red-brown rhizoids. Gemmae usually present. A. palustre

2b. Leaves usually green, erect and imbricate, not contorted when dry. Stems separating easily, the rhizoids largely hidden by the leaves. Gemmae absent. A. imbricatum

A. androgynum (Hedwig) Schwägrichen. An infrequent species of a very specific habitat: rotten, often charred, wood, on the ground on slopes from the foothills canyons, under Pseudotsuga to the subalpine forests.

A. imbricatum***. Restricted to the highest wet alpine areas. This is a plant that strongly suggests the Arctic A. turgidum (but with smaller leaves) or A. imbricatum (but with rounded leaf apices). Collections: Boulder, Clear Creek, Larimer, Summit Counties. This has been considered to be merely a high altitude modification of no taxonomic significance, a response to the peculiar climatic conditions of the high alpine zone. However, this form never produces gemmae, does not expose its rhizoids, hence the stems do not cling to each other; the tufts do not grow intermixed with other mosses and tend to be darker green rather than yellowish. Vitt (in litt.) has grown this in the greenhouse and reports that under those conditions it reverts to type, but we have not seen vouchers of this. We believe that no other mosses in our region exhibit such a pronounced ecological modification. A. imbricatum is extremely well developed and abundant on the saddle between Mount Evans and Mount Epaulet, at 13,500 ft. alt. Collections: Boulder, Clear Creek, Larimer, Summit Counties.

The most recent, and really the only description of this plant is given in C. C. Pedrotti, Flora dei muschi d'Italia: Sphagnopsida, Andreaeopsida, Bryopsida (I parte). 2001, p. 686. I translate:

"Cushions dense. Stems scarcely tomentose; leaves erect-imbricate and acute, linear-lanceolate, apex obtuse, margin entire. Vegetative reproduction absent. Ecology: Environment [stesso ambiente] similar to that of the species, subalpine and alpine flats. Because of the erect-imbricate leaves in the dry state it may be confused with A. turgidum, a species not yet reported for Italy which differs in having ovate leaves with obtuse or rounded apices [spesso] cucullate-concave, the cells not or scarcely papillose and with a distinctly sinuose costa."

Usually with modifications there are intermediate forms along an ecological gradient. This is such a strikingly different plant in the field from the abundant lower altitude A. palustre that we feel justified in giving it specific status, especially in view of the fact that it occurs in isolated tundra sites in Austria (the type locality), Norway, Italy, and Russia (Lake Baical).

A. palustre (Hedwig) Schwägrichen. One of the most abundant mosses of wet areas, occurring in willow swamps and fens in the upper montane and subalpine. The plants typically have narrow, contorted or twisted yellow-green leaves and abundant tomentum on the stems.

Ref: Flowers (1952), Griffin (1989, 1998)

la. Leaves from a more or less clasping or sheathing, differentiated and greatly enlarged basal portion, the lamina subulate. Bartramia

lb. Leaves from a non-clasping, non-sheathing, usually ovate base, the lamina subulate or broader (2)

2a. Lamina subulate or linear; tufted or matted plants of cliff-sides, not associated with seeps, springs, or running water (3)

2b. Lamina lanceolate or ovate; with terminal branches frequently in whorls, with the spreading, short branches subtending antheridial buds; plants of springs, seeps, streamsides and fens. Philonotis

3a. Leaves broadly linear; stem 3-angled in cross-section; plants with erect stems, forming rounded tufts, usually fruiting. Plagiopus

3b. Leaves with subulate lamina; sterile in our area. Anacolia

According to Griffin (in litt.), sterile material of Anacolia can be distinguished from Bartramia by the cross-section of the stem. In Anacolia the cortical cells are highly mammillose, and there is no hyalodermis. In Bartramia a hyalodermis is present and thus the cortical layer is smooth. The axillary hairs (difficult to make out) are of two cells, the basal one more or less quadrate and brown, the terminal one globose and hyaline. In Bartramia they are of 3 or more cells, the terminal cell elongate, thick- or thin-walled, the intercalary cells hyaline or with pigmented cross-walls, the basal cell(s) sometimes brown (Griffin 1998).

1a. Distal cells of leaf papillose at the ends on both surfaces; inner basal cells linear; upper lamina 2-3-stratose. A. laevisphaera

1b. Distal cells smooth or only a few with low papillae at the ends on the abaxial surface; inner basal cells quadrate or short-rectangular; upper lamina 1-2-stratose. A. menziesii

A. laevisphaera (Taylor) Flowers. One record (verified by Flowers and by Griffin) from eroding soil over sloping granite outcrop on west side of Boulder Falls. The site has been destroyed by foot traffic. This otherwise barely gets into the United States in New Mexico and southern Arizona. We have one collection from northern New Mexico.

A. menziesii (Turner) Paris. A very beautiful moss, forming sprawling mats. Frequent on north-facing granite cliffs in the Front Range near Boulder. A. menziesii roughly follows the distribution of the redwood forests of California, with outliers in northeastern New Mexico and northwestern Wyoming.

1a. Leaves rigidly erect-appressed, the shoulders of the sheath with very thin-walled, elongate cells that often appear collapsed; dioicous. Bartramia potosica

1b. Leaves laxly appressed to flexuose; synoicous (2)

2a. Seta short, 3-4 mm long; endostome rudimentary; exclusively high alpine. B. subulata

2b. Seta long, 5-7 mm long; plants of middle altitudes up to subalpine. B. ithyphylla

B. ithyphylla Bridel. A fairly common species of the subalpine and alpine, the only Bartramia that will be generally met with. It is commonly fruiting.

B. potosica Montagne. A rare species, this occurs on rock ledges in the foothills canyons, on north-facing cliffs. It appears to be distributed from Larimer to El Paso County, but is nowhere abundant. It is always sterile here. The remarkable sheath, with its margin of thin-walled cells, at first suggests a member of the tropical family Calymperaceae! It is distributed along the Front Range of Colorado and New Mexico, throughout Mexico and Central America, and in the Andes from Peru to Argentina. Its occurrence in the Front Range reinforces the pattern of very ancient Tertiary disjunct species.

B. subulata Bruch & Schimper. A very rare species usually occurring on wet tundra at high altitudes, known in Colorado from Summit Lake on Mt. Evans, and Blue Lake in Summit Co. However, our best material comes from the montane-subalpine ecotone along Lost Creek, in Park County, where it occurs on wet rocks along a small rivulet in the wet meadows of the valley floor. The very short and stout setae, erect, symmetrical capsule which becomes black and loses its peristome at maturity, and short, stiffly appressed leaves distinguish it from B. ithyphylla. In America this is restricted to the Rocky Mountains from Colorado to Alaska. Its distribution in the Old World is not clear.

Sven Fransén has provided a table for the separation of our species:

1a. Leaf cells with a single papilla projecting from the lower (proximal) end; perigonia (male buds) terminal on the stems, when fresh open wide, when dry appearing nut-like; some marginal leaf cells with a projection at each end, suggesting a double tooth. P. fontana

1b. Leaf cells with a papilla projecting from the upper end or from both ends; perigonia similar; marginal leaf cells with a single projection or tooth. P. marchica

A note on the papillae: When a fresh leaf is placed under the microscope, little round objects with distinct edges can be seen over the cell lumen. They are not papillae. The papillae are not distinctly margined, but can be seen only when one moves the objective slowly up and down. The papillae are visible as faint mounds. In P. fontana these are at the bases of the cells. In P. marchica they are at the distal ends. Flowers suggests that boiling the leaf in a drop of lactophenol will make the papillae more obvious.

The plants are dioecious. The perigonial (male) shoots have strongly appressed leaves; below these, the stems of the previous season have falcate leaves. The perichaetial (female) stems commonly have radiating whorls of subterminal branches. Entire tufts thus will either be male or female and very different in appearance.

P. fontana (Hedwig) Bridel. Extremely common in wet places throughout, especially abundant on seepage areas along highways, where its yellow-green color makes it easily recognizable from a moving car. The subspecies pumila (P. tomentella) is characterized by its very dense, compact growth form, the copious production of tomentum, the non-plicate leaves that are slenderly acuminate and with a long-excurrent costa. This expression of P. fontana tends to be most frequent at very high latitudes in the Northern Hemisphere and at high elevations in the temperate mountain ranges.

P. marchica (Hedwig) Bridel. Evidently relatively uncommon. Our records are mostly or all from the San Juan Mountains. It possibly is restricted to areas of high mineralization.

Further notes: The present treatment is very much simplified from that of most texts, where the species and varieties are legion. Dana Griffin III writes: "In his dissertation on Philonotis for North America north of Mexico, William Zales. a student of Wilfred Schofield, attacked the P. fontana complex in a way different from that of most other bryologists. He brought several specimens into culture and grew them in light boxes and in the greenhouses at Univ. of British Columbia. What he found out from this approach is that many of the characters used to define species allied to P. fontana are (to use his term ecophenic characters). Changes in the local environment can influence the elaboration of these characters. In other words, such characters form a very poor basis on which to justify a particular taxon. Zales found three characters that, taken together, help identify P. fontana: Discoid perigonia, adaxial cells of the lower lamina with postical projections and some marginal cells showing paired teeth.

"The paired teeth character needs to be clarified. For Zales, it means a tendency in some contiguous marginal leaf cells to develop apposing and recurved projections. When well developed, this produces a 'double tooth' that might recall the bifid teeth in the leaves of some Hookeriaceae. However, here we are not dealing with a bifid tooth (which is produced by a single mother cell that bifurcates but rather two side-by-side projections that meet and then curve away from each other.

"If one accepts this as the basis for identifying P. fontana, it then becomes largely a matter of taste (sounds better than bias) how many subspecific entities one wishes to recognize. P. tomentella, also known as P. fontana var. pumila is characterized at the varietal level by its very dense, compact growth form, the copious production of tomentum, the non-plicate leaves that are slenderly acuminate and with a long-excurrent costa. This expression of the species tends to be most frequent at very high latitudes in the Northern Hemisphere and at high elevations in temperate mountain ranges."

P. oederianus (Swartz) Crum & Anderson. This species forms dense and extensive sods over seeping rocks in the subalpine and lower alpine zones. Mrs. Nyholm writes (Moss Flora of Fennoscandia), "In moist, deeply shaded habitats the tufts become shorter and denser and the leaves shorter and more crowded. On dry alpine rocks forma alpina, with smaller capsules, grows in small tufts." This form is very abundant in the upper valley of Monte Cristo Creek in Summit County below Hoosier Pass.

March 19, 2005

Note: In order to understand some of the problems involved with the taxonomy of this family, read Robinson, H. 1962. Generic revisions of North American Brachytheciaceae. Bryologist 65:73-146. 1962. This is an important paper and deserves more attention.

la. Leaves concave, cucullate, rounded at the apex but then abruptly extended into a long filiform point; stems often julaceous. Cirriphyllum

1b. Leaves without the above combination of characters (2)

2a. Leaves blunt-pointed or rounded at the apex, erect-spreading when moist, serrulate from base to apex; costa ending below the leaf apex, the terminal cell protruding from the dorsal side of the lamina as a small spine. Rhynchostegium

2b. Leaves definitely acute or acuminate; variously smooth or denticulate along the margins; costa not protruding as a dorsal spine (3)

3a. Leaves linear-lanceolate, very strongly plicate, appressed, glossy; basal cells not differentiated from the median ones (4)

3b. Leaves broader, lanceolate or ovate, plane or more or less plicate; basal leaf cells shorter and broader than the median cells (5).

4a. Stems with masses of brown rhizoids on the older parts; shoots essentially straight; plants of fens. Tomentypnum (see Campyliaceae)

4b. Stems without masses of rhizoids; stems with the branches up-curved when dry, of a shining golden color; plants of vertical cliff faces in the foothills. Homalothecium

5a. Stems complanate, the leaves more or less distant, never plicate; alar cells not differentiated. Rhynchostegium

5b. Stems not complanate, the leaves usually not distant, often imbricate, plane or plicate; alar cells usually differentiated (6)

6a. Small plants in deep compact cushions on moist calcareous sandstone cliffs or ledges; leaves never plicate; margin serrate, the teeth at the leaf base often double and recurved; short-cylindric gemmae commonly produced along the costa near the leaf apex. Conardia

6b. Medium-sized to large plants on the forest floor, soil of canyonsides, fens, or tundra; leaves plane or plicate; teeth of leaf base not as above; gemmae absent. Brachythecium

March 8, 2004

1a. Leaves strongly falcate-secund, almost circinate (resembling Hypnum or Drepanocladus), biplicate; branches strongly pinnate and also curved; plants in extensive, tight mats (as in B. collinum). B. leibergii

1b. Leaves either straight or sometimes falcate but never circinate, and larger (2)

2a. Leaf margins entire or nearly so, sometimes slightly serrulate at the apex only; seta smooth (or nearly so) (3)

2b. Leaf margins serrulate in the upper half or sometimes to the base; seta rough or smooth (5)

3a. Plants robust, in loose, somewhat shiny, green or golden-brown tufts, the stems subjulaceous, erect-ascending, sparsely branched, the leaves appressed; strictly alpine; autoicous. B. turgidum

3b. Plants of medium size, mostly creeping, much branched; dioicous (4)

4a. Leaves narrowly lance-ovate, gradually narrowed to long acumen, appressed to the stem (as if combed); the branches short, curved at the tips, flattened; plants growing in dry duff on forest floors, light yellow-green. B. erythrorrhizon

4b. Leaves broadly ovate, more abruptly narrowed, concave, some somewhat falcate, the bases rounded to the the stem, very slightly decurrent and producing a shaggy appearance; plants of more moist sites, green. B. salebrosum

5a. Plants small; leafy stems less than or 2 mm wide; leaves 1 mm long, not plicate; capsules short (6)

5b. Plants medium-sized to somewhat robust; leafy stems more than 1 mm wide; stem leaves 1.5 mm long or more, entire, often plicate; capsules often elongate (9)

6a. Plants true green; stems not julaceous, leaves spreading (7).

6b. Plants bright, yellow-green, mats not soft or silky; stems julaceous, leaves appressed (8)

7a. Plants in soft, dark green, silky mats; leafy stems less than 1 mm wide; branch leaves serrulate throughout; leaves somewhat falcate, long-tapering to the slender point; alar cells in a short group, not nearly reaching the costa. B. velutinum

7b. Plants with bright green foliage, not in silky mats; leafy stems broader; leaves concave, abruptly narrowed to a slender tip, very faintly serrulate near the apex; alar cells more numerous, reaching close to the costa. B. plumosum

8a. Capsules straight and erect. B. fendleri

8b. Capsules inclined to horizontal. B. collinum

9a. Leaves 1.5 mm long, broadly ovate, acuminate, decurrent, 1.5 mm long; seta rough (use microscope). B. oedipodium

9b. Leaves over 1.5 mm long; seta (smooth or rough) (10)

10a. Stem leaves triangular-ovate, slenderly long-acuminate; alar cells inflated, forming auricles; seta rough; weak plants of subalpine willow fens. B. nelsonii

10b. Stem ovate, but not triangular; leaves more or less abruptly acute or shortly acuminate. . .

(11)

11a. Leaves entire or minutely denticulate; alar cells inflated, numerous across the leaf base; wiry plants tending to have the branches erect or arching on seeping forest slopes and edges of swiftly flowing streams. B. rivulare

11b. Leaves distinctly serrate, especially near the apex; alar cells small, not inflated nor conspicuous ; plants soft in texture; on rocks in ravines, but not in really wet sites. B. acuminatum

B. acuminatum (Hedwig) Austin. A mid-western species, recently discovered in Boulder County, in the Boulder Mountain Parks, in the saddle between Green Mountain and Flagstaff Mountain, Weber & Wittmann 112049, and in El Paso Co.: Williams Canyon, Weber & Wittmann 113146. We do not believe it to be distributed widely over Colorado, for it evidently belongs to a group of plants that are relictual on the east side of the mountains. The leaves are appressed when dry, the branch tips not curved, leaves spreading at a narrow angle when moist; leaves biplicate, shortly acuminate, recurved in the lower half, the margins serrulate distally; costa vanishing below the acumen; alar cells numerous, quadrate, the cells across the base of the leaf similar.

B. collinum (Müller Hal.) Bruch & Schimper. Our most abundant species on slopes in the forested outer foothills where it inhabits sites that are dry except in springtime. It also occurs in optimum moss tundra and probably throughout the middle altitudes. Along with B. fendleri, it is our smallest species. The plants are yellow-green, with short and commonly julaceous branches; the leaves are closely spaced along the stem, non-plicate, convex, appressed, and are sharply serrate from apex to near the base. The alar cells are numerous and quadrate. Fortunately, it fruits in the lower foothills, where it occurs with B. fendleri and is distinguished by the inclined rather than erect capsules. B. collinum, B. fendleri, B. curtum, B. leibergii, B. oedipodium, and B. velutinum form a group of species that have much smaller leaves (1-1.5 mm long) than the group that includes B. erythrorrhizon, B. nelsonii, and B. rivulare.

B. erythrorrhizon Bruch & Schimper. Our most abundant species in the forested regions. Our records are scattered through the middle altitudes and upper subalpine, where it occurs on the forest floor and over rotting wood. In suburban areas it commonly occurs mixed with Ceratodon purpureus and Hypnum cupressiforme in poorly drained areas of lawns. The plant forms loose, spreading patches, with light green or straw-colored foliage, long-acuminate, slightly falcate, plicate leaves, so that the branches are somewhat curved. At intervals along the stem, there are clusters of long, reddish brown rhizoids, hence the specific epithet erythrorrhizon.

Lawton identified a specimen (Hermann 23257a) from Box Canyon, Ouray, as B. asperrimum. The branches are distinctly falcate, the leaf margin is faintly, not strongly serrulate, and there are no capsules. In B. asperrimum the seta is very strongly papillose. In the absence of more and better material I would hesitate to add B. asperrimum to the list.

B. fendleri (Sullivant) Jäger & Sauerbeck. Frequent in rocky canyonsides in the outer foothills, especially in the southern counties. The erect capsule serves to distinguish this from the very closely related B. collinum. Evidently B. fendleri is a plant of the southern counties or in low elevations elsewhere. There should be some vegetative distinction, but thus far we have not found any significant one.

B. leibergii Grout. A species of the Pacific Northwest, known in Colorado only by a few collections from humus and rotting logs in dry spruce forests. Routt Co.: Park Range, near beaver pond 1-2 mi above Slavonia, E of Clark, Weber & Nelson B-49372; Boulder Co.; N Fork Middle Bopulder Creek 8 mi N of Nederland, Flowers 9819 (COLO 54933). With its small, strongly falcate-secund leaves, this is like no other Brachythecium, and except for the costa, it might be mistaken for a tiny Hypnum. It is a very much smaller plant than Sanionia uncinata, which also combines a costa with falcate leaves, and its habitat separates it from Drepanocladus. The seta is rough.

B. nelsonii Grout. A beautiful species of wet ground in willow fens and wet tundra. It does not occur along flowing streamlets and is a plant of high altitudes. It usually does not form dense clumps. The branching is pinnate, with the lateral branches more or less at right angles with the stem. The stem leaves are very translucent, rather remotely spaced, and are broadly triangular ovate, with distinctly auriculate alar cell groups. The branch leaves are distinctly smaller (B. latifolium Kindberg. Based on Hypnum latifolium Lindberg, 1879, not Taylor 1848).

B. oedipodium (Mitten) Jäger. This relatively small species is fairly common in dry forests in the montane and subalpine. The leaves are small, and serrulate or serrate, so one might think it to be a very loosely branched form of B. collinum. However, the branches are not julaceous, the leaves are weakly serrate, the quadrate alar cells are larger (about 20 m) and there are some oblong cells along the basal margin. The seta is rough. Crum and Anderson state that this is the plant that has been called B. starkei and B. curtum in American floras.

B. plumosum (Hedwig) Bruch & Schimper. A very uncommon species, of which we have only a few collections:Larimer Co.: Moist face of granite boulder along trail near summit of Greyrock Mt., 7,400 ft., 10.5 mi WNW of LaPorte, Hermann 23652 (B-38552); Rocky Mt. Nat. Park, trail to Mills lake, Hermann 26957 (COLO 111799). Some specimens of Rhynchostegium pulchellum have been called this by Lawton, but in that species the leaves are strongly serrate and with a blunt or even rounded apex. Flowers does not appear to have understood this species either. His Colorado collections belong to B. oedipodium. B. plumosum does not appear to have very slender points except under the microscope. We find the rather short, not filiform, and abruptly narrowed apex, combined with the concave leaves, good for recognition. The leaves are concave, somewhat spreading and often turned to one side. Very few treatments do a good job of finding distinctiveness, but it is one of a few species that have small bright green leaves.

B. rivulare Bruch & Schimper. An abundant species wherever there are springs and moss-lined forest seeps and brooks. This is really an easy species to recognize. Most other forest species of Brachythecium are soft, forming flat mats on the ground. Unlike them, B. rivulare is stiff and somewhat wiry, with somewhat arching stems with erect, not overlapping, branches (some call these somewhat dendroid!). The stem leaves are strongly decurrent, and are difficult to remove with a tweezers. The stem leaves are larger than the branch leaves. The leaves are broadly ovate, the apex acute but not drawn out, with the costa not entering the acumen. Very large patches of inflated alar cells continue down the decurrent margin of the leaf. The leaves are entire or very faintly denticulate in the uppermost part.

B. salebrosum (Weber & Mohr) Bruch & Schimper. Evidently very infrequent. Often B. erythrorrhizon has been mistaken for this. But the leaves in B. salebrosum are not appressed to the stem, nor do they have a combed appearance, and they are deep green. The bases of the leaves bow out so that the stem is subjulaceous. The leaf arrangement is somewhat loose and has a shaggy appearance. The stem tips are usually somewhat falcate. The alar cells are quadrate and generally not very clear in contents. We have collected a fine set of specimens from the vicinity of Caribou mountain peat "bog," at 10,000 feet, in peaty soil on a spring bank under willows, Weber & Wittmann B-11742. However, it is very infrequent in our experience.

B. turgidum (Hartman) Kindberg. A robust species, generally growing in almost erect clumps, and is the only large Brachythecium occurring in moist tundra. The stems are more or less erect, very sparsely branched, and subjulaceous. The leaves are densely packed, plicate, 2.5-3.0 mm long, abruptly narrowed to an acuminate tip, and strongly appressed to the stem. It is said to be a calciphile, but it does not seem to be thus in Colorado. We have specimens from BL, CC, EA, GA, LK, PA, ST.

B. velutinum (Hedwig) Bruch & Schimper. This species has been found in the outer foothills in the Pseudotsuga zone near Boulder. It is not common, and occurs in extensive mats on vertical faces of shaded boulders near small streams. The leaves are small, as in B. collinum, but they are widely spreading, falcate, and not at all julaceous, and the foliage is dark green, not straw-colored as in B. oedipodium. It is usually fruiting and the seta is rough throughout.

C. cirrosum (Schwägrichen) Grout. This occurs in the upper subalpine and alpine zones on wet tussocks in tundra. Often it occurs as scattered solitary stems mixed with other mosses, but occasionally pure stands occur as loose mats. C. brandegei (Austin) Grout does not appear to be more than an ecological modification, said to differ by a more caespitose habit, looser leaf cell areolation, and an entire rather than denticulate acumen.

C. compacta (Müller Hal.) H. Robinson This very much resembles Amblystegium serpens, but the leaves are distinctly serrulate from base to apex, the median cells are longer and narrower (sometimes up to 10:1), the costa going well into the apex, often expanding there, and stouter than in A. serpens. A very good separating characteristic is the conspicuous serrations, often recurved, along the basal margin. Also, the long , narrow median leaf cells separates it from Amblystegium serpens, and the well-developed costa from Isopterygiopsis. Fertile specimens may be distinguished from Amblystegium because the operculum has a narrowed beak in C. compacta but is conical in Amblystegium. When the characteristic gemmae are present there is no problem. C. compacta appeared to be restricted to calcareous sandstone cliffs and ledges at low altitudes, but we find it especially abundant at High Creek Fen! We also have collections from Boulder, Gunnison, and Montezuma counties. The plant is light green, often rather densely matted and mixed with calcareous deposits, which the moss evidently is instrumental in accumulating into a travertine deposit (Rhynchostegiella compacta).

Ref: Hofmann (1998)

H. aureum (Lagasca) H. Robinson. One of the most beautiful mosses of our area. When moist, its large size, more or less regularly pinnate branching and burnished golden color is very attractive on cliff-sides. When dry the species has a totally different aspect, with the short lateral branches having their leaves strongly appressed and the axes curved upwards, of a distinctly golden color. It is always sterile here. Our collections are mostly from the outer foothills of the Front Range, but we have one collection from Moffat County: Yampa River Canyon, Lehr 2020 (B-113301). The species is probably restricted to relatively low altitudes.

In his classic paper on the Brachytheciaceae, Robinson (1962) I believe justifiably synonymizes H. nevadense and a number of other northwestern species under H. aureum (Lagasca) H. Robinson, based on Hypnum aureum Lagasca, 1802.

Hofmann (1998) recognizes two subspecies under H. nevadense: subsp. nevadense and subsp. aeneum (Mitten) Hofmann, and cited specimens of each subspecies from Boulder Canyon, which seems unlikely. In subsp. aeneum the alar cells are 2-7 along the margin of detached leaves, more clearly visible, and mainly quadrate, and in subsp. nevadense the alar cells are 1-4 along the leaf margin, less clearly visible, and mainly irregular in shape. The capsule is curved to rarely nearly straight in subsp. aeneum, and straight or rarely somewhat curved in subsp. nevadense. Our plants have never been found in fruit. Furthermore, there seems to be no clear geographic separation.

1a. Leaves complanate, ovate, weakly serrulate, narrowed to a fine, twisted point. R. serrulatum

1b. Leaves not complanate, strongly serrulate, the apex broadly acute or obtuse. R. pulchellum

R. pulchellum (Hedwig) H. Robinson. This is very characteristic of mesic Pseudotsuga stands on moderate to steep slopes on north-facing slopes in the foothill canyons. Once recognized, it is easy to know. The leaves are short, broad, triangular-ovate, and serrulate from base to apex. Even under a hand-lens the neatly rounded, serrate apices on the glossy leaves spreading at a 45 degree angle have a very characteristic appearance shared by no other species. Macroscopically it might be mistaken for a species of Brachythecium.

Most of the Colorado collections from the relatively dry forested slopes belong to var. pulchellum forma praecox (Hedwig) Crum, Steere, & Anderson, in which the main stems are very short, the branchlets short and very numerous, and the leaves generally appressed to the stem when dry (suggesting Brachythecium collinum in size). Rarely, a very large form occurs in more mesic forests, resembling a large Brachythecium, with elongate main stems and strongly pinnate branching, with the individual leaves standing out from the stem. This is var. barnesii (Renauld & Cardot) Crum, Steere & Anderson. Superficially they are so different that its is difficult to reconcile the two types under one species, but there is nothing qualitative we can find to separate them taxonomically.

R. serrulatum (Hedwig) Jäger & Sauerbeck. We have only one collection: Montezuma Co.:Mesa Verde National Park: Wetherill mesa, under ledge of west pour-off of Bobcat Canyon, 6900 ft. alt., 8 Aug,. 1959, Erdman (COLO B-3688), where it was growing under the lip of a cliff over which water seeps intermittently (a fragmentary specimen!).

Isothecium : Reported by Grout, MFNA, from a collection by Shockley. Likely An error in transcription of the label. (Cal, for California read as Col, for Colorado). A plant of the Pacific coast. Shockley did not collect in Colorado. Such transcription errors have occurred frequently in the vascular plant collections.

Scleropodium obtusifolium (Mitten) Kindberg. This was once reported from a fish stomach but has not been collected since. It might be expected in the north-central tier of counties.

March 19, 2005

The Bryaceae are roughly characterized as being acrocarpous mosses, a complete peristome with both an exostome and endostome, and rhombic distal laminal cells without ornamentation. Most commonly the sporophyte, when present, is pendent, with a dome-like operculum. Sterile plants have been a real problem, causing even specialists to throw up their hands and give up on them, assigning them to such trash-heaps as "Bryum caespitosum." The principal genus in this family has been a bugaboo, at least to American bryologists, for over a century. Keys to the species of Bryum have proved to be frustrating, since they have relied so much on having mature sporophytes.

Unfortunately, in Colorado most species of "Bryum" in the broad sense are almost always sterile. A great number of the collections have been misidentified or left without names. However, during the past few years John Spence has examined the group and discovered that vegetative characters of the leaves, stems, and rhizoids are equally useful in classification. He has divided this monster genus up into a number of smaller ones based largely on their vegetative features. This has been a blessing for Colorado, for now with his insight we can feel confident in our identifications and, with a little practice, we can recognize most of them in the field.

Here are some characters that need to be recognized. Rather than use the generic names, a

specific epithet should be enough for an example.

The leaf itself.

a. The leaf shape is usually oblong-ovate or obovate. It may be narrowest at the base (angustifolium) or contracted to a narrower base (laevifilum). Many species, especially of Ptychostomum, have ovate-lanceolate leaves. The only linear-leaved species is Leptobryum pyriforme, which probably does not belong to this family.

b. The leaf margin may have a border of narrower cells; this may be inconspicuous (hardly differentiated in length and width from the laminal cells), conspicuous, or even bistratose. This border is called a limbidium. It might be compared to the frame of a tennis racket.

c. The basal margin of the leaf sometimes is decurrent (pseudotriquetrum).

d. The leaf will have a distinctive form when dry. It may be essentially flat, appressed to the stem (pallescens), or convex and curving toward the stem distally (calobryoides). Or it may be variously contorted, sometimes distinctly spirally arranged around the stem. The leaf form when wet or dry must somehow be governed by the presence and thickness of the limbidium.

e. Some species, especially those occurring in fens, such as P. pseudotriquetrum, may resist wetting. This is a common adaptation occurring in wetland mosses. Dry-land mosses commonly imbibe water immediately.

f. The costa may be red (angustifolium, pseudotriquetrum) at the base or not. It may not reach the leaf apex or may be excurrent to form a slender or stout awn. Very few species have the excurrent costa so slender and colorless to be called a hair-point. An excurrent costa is usually stout and best referred to as an arista.

g. The leaf color may be bleached, as in B. argenteum and B. lanatum that have almost no chlorophyll distally. The color may be deep pink or red in B. pallens and B. cryophilum. With practice, different constant shades of green, turquoise, or brown may help in recognition of species.

The stem and its leaf arrangements.

a. The stem may be red or green. It may be simple or branched; the stem and branches are erect and tend to make for a dense tuft. Stem color is often an important feature.

b. The leaves may be distributed uniformly the length of the stem (most species). They may also be clustered at the end of the stem, forming a bud-like tip or a rosette of leaves.

c. Stem length may be characteristic of the species. Stems may be very short, not more than a few millimeters . Medium length stems range from 5 mm to 30 mm. Long stems will run more than 30 mm.

Rhizoids.

The stem may or may not be clothed with brown rhizoids. Rhizoids have no chlorophyll, and the end walls of the cells are slanting. Rhizoids may be completely smooth or densely papillose. This feature is doubtless of taxonomic importance, but descriptions commonly omit mention of this character. A thorough survey is yet to be undertaken.

Cell detail (areolation types).

Spence (2004) describes four areolation types. These are basic to his reclassification.

1. Bryum type: Cells heterogeneous, the distal and median lamina cells elongate, mostly 4:1 or greater, long-rhomboidal or linear, the proximal cells wider and shorter than the median cells, quadrate or short-rectangular, the transition often abrupt, giving the areolation a heterogeneous appearance.

2. Plagiobryum type: homogeneous, all cells except in the alar region elongate, mostly 4-8:1. no strong distinction between the upper 2/3 and lower 1/3 of leaf.

3. Rhodobryum type: Heterogeneous, the distal and median lamina cells short rhomboidal to hexagonal, 2-4:1; proximal cells the same width or narrower, and the same length or longer, more regularly rectangular with squared-off ends, the transition rather gradual but still giving the areolation a heterogeneous appearance.

4. Pohlia type: Homogeneous, all lamina cells except those at the leaf base long and narrow, mostly linear to hexagonal, often thick-walled.

Asexual reproductive structures.

Mosses commonly reproduce by simple fragmentation when they are dry and brittle. But several species have specialized reproductive structures that are usually present.

a. Bulbils are short, plump structures that have a narrow opaque base and usually a few green, short, leaf primordia appressed to the sides. They fall out of the leaf cluster when manipulated.

b. Filiform axillary gemmae. These are found in the axils of the leaves, sometimes very abundant, looking rather like fat rhizoids that are filled with chlorophyll and with perpendicular cross-walls rather than the diagonal ones of rhizoids. They also fall free when the leaves are dissected out.

c. Rhizoidal gemmae. These are spherical multicellular bodies ranging from 120 microns to 250 microns. They are loosely attached to the slender rhizoids, and in our species are orange or red -brown. These are inconspicuous, weak and short-lived, weedy mosses. In order to detect rhizoidal gemmae, the specimen should be soaked in water, then gently teased away from the surrounding sand grains (sometimes these are very tenacious). The remaining rhizoids should be carefully scanned on a slide.

The seta and capsule.

a. The seta may be long and slender or short and thick. While the nature of the seta may sometimes be diagnostic, for the most part it is not.

b. The operculum. Some species have an operculum that equals the mouth in circumference; some have small opercula. We have not explored this in the Colorado species.

c. The capsule may be cylindric and slightly wide at the mouth. It may or may not have a distinguishable neck that becomes shriveled. The mouth of the capsule may have a prominent ring of red cells. The cells of the capsule wall may by thick or thin, long or short. We have not explored this.

d. Most capsules are pendent. A few are "leaners."

e. The outer peristome. The exostome is usually stout, yellow or brownish. In only one species (Ptychostomum pendulum) does the exostome exhibit any especially remarkable feature. Instead of the ladder-like sequence of cells from top to bottom, the adaxial side of these teeth have an additional layer that resembles circular or variously shaped chambers (this is the endostome that is adherent to the exostome). I believe that fertile specimens are needed, and they are usually present. I do not know how to recognize sterile specimens.

f. The inner peristome (endostome) has a hyaline basal half that consists of united segments, which separate upward to form teeth. Between two teeth there is usually a one-cell wide filament called a cilium, which may have one or more short lateral perpendicular branches, which makes the cilium "appendiculate." A peristome may be examined by softening the capsule by boiling momentarily, and cutting the upper part of the capsule free with a piece of a razor-blade and separating the resulting ring with fine needles.

1a. Capsule horizontal, the seta stout, the urn curved, bloated distally with the operculum pushed to the side. Plagiobryum demissum (this tundra species is not easily recognizable except with capsules, but these are almost always present.

1b. Capsules not as above, or plants without capsules (2)

2a. Leaves narrowly linear-setaceous; plants usually fruiting abundantly, the capsule pendent or horizontal, pyriform with a narrow neck; common on burned ground, burned peat etc., in middle and higher altitudes. Leptobryum

2b. Leaves broader than linear; capsules present or lacking, of various forms (3)

3a. Leaves silvery-white (at least most of them), giving this color to the tuft (4)

3b. Leaves some shade of green, yellowish-brown, golden or reddish (5)

4a. Forming dense, extensive tufts, tightly packed; leaves minute (0.3 mm), broadly ovate, acuminate, the costa not conspicuous, often disappearing in mid-leaf. Bryum, sens. lat. (9)

4b. Occurring as scattered stems in tufts of other alpine mosses; leaves 0.5 mm (the older stem leaves up to 2 mm), broadly ovate, suddenly recurved-apiculate; costa conspicuous to the apex; all cells of leaf pale and often the basal more or less collapsed. Plagiobryum zieri

5a. Inflorescences appearing lateral; leaves soft, with costa not reaching apex or percurrent, rarely weakly excurrent; laminal areolation Plagiobryum-like; alar cells somewhat differentiated from juxtacostal cells, quadrate; capsule shortly pyriform, lacking a peristome. Haplodontium

5b. Not as above. (The likelihood of finding Haplodontium is extremely slight. It is only found on limestone cliffs in the San Juans.) (6)

6a. Leaves very large and translucent, 5-6 x 2-3 mm, in a rosette at the stem apex, not much changed or scarcely contorted in drying, papery in texture; leaf cells thin-walled and very large (150 x 50 m). Roellia

6b. Leaves smaller, usually somewhat altered in drying; leaf cells never so large (7)

7a. Plants very short, forming extensive tight mats, strikingly turquoise-colored, very rarely fruiting; mostly restricted to mine tailings in mineralized areas of the San Juans, and on limestone cliffs in the Elk Mountains. Mielichhoferia

7b. Not as above (8)

8a. Median leaf cells 5:1 or longer; leaves rarely bordered, often serrulate distally; costa ending below the apex or percurrent, never excurrent. Pohlia

8b. Median leaf cells 4:1 or often shorter and broader, entire; if longer, then with at least one of the following characters: leaves broadly ovate, oblong, or oblong-ovate, usually not serrulate; apex obtuse to rounded; costa excurrent; leaf margins bordered (sometimes weakly) by longer cells. Bryum sensu lato (9)

9a. Stems short, julaceous; leaves minute, green, yellow green to silvery; if stems not julaceous then plants silver and basal cells quadrate; lamina areolation Bryum-type. Bryum

9b. Plants and leaves of various sizes; fertile stems not julaceous although innovations sometimes so; if julaceous, then leaves >2 mm long; lamina areolation various (10)

10a. Lamina areolation open, transparent (as stained glass windows), the cells usually broad; gemmae often present, either as tubers on the rhizoids or axillary bulbils; limbidium often very narrow or absent. Gemmabryum

10b. Lamina areolation various, limbidium distinct; vegetative propagula lacking or filamentose. . (11)

11a. Leaves ovate to ovate-lanceolate or lanceolate, widest at or below the middle, not distinctly rosulate; lamina margins smooth to finely serrulate but not strongly serrate; limbidium uni- or bistratose (most of the common densely tufted mosses belong here). Ptychostomum

11b. Leaves usually obovate or oblong to spatulate, widest above the middle usually rosulate; margins serrulate to serrate, unistratose; filiform gemmae commonly produced in the leaf axils. Rosulabryum

1a. Leaves, at least the upper parts, white and lacking chlorophyll (if on tundra, see Plagiobryum zieri); plants forming low, dense patches, mostly at low to medium altitudes (2)

1b. Leaves green, brown, or variable in color (4)

2a.. Plants hoary white; stems evenly foliate, slender, costa strong, excurrent in a stout hair-point, leaf apices recurved when dry; plants of undisturbed sites, having a shaggy appearance because of the long white leaf tips. Native plants of western and southern Colorado. B. lanatum

2b. Plants silver to silver-green; stems short, julaceous to gemmiform, costa not reaching apex; apiculus usually present, irregularly curved when dry (3)

3a. Plants tiny; stems 1-3 mm long; leaves broadly ovate, larger leaves broader than long; distal laminal cells short-rhomboidal. 2-3:1 Plants of desert gypsum flats.. B. nanoargenteum

3b. Plants larger; stems 2-10 mm long; leaves ovate, longer than wide; distal laminal cells narrower, 3-5:1. B. argenteum

4a. Stems very slender, ropy, tightly julaceous, the leaves appressed, narrow; leaf cells narrower than in most species; often growing solitary or a few stems mixed with other mosses. B. julaceum

4b. Not as above (5)

5a. Plants forming dense tufts, the stems sometimes elongate, the terminal leaves crowded into cabbage-like heads, very broad and short, circular, obtuse or rounded, cucullate and with a short apiculus, the leaves not shrunken when dry; common from foothills to alpine, never fruiting here. B. calobryoides

5b. Plants not forming tufts, the stems very short, gemmiform, in a terminal cluster; sporophytes always present, with very short seta and broad ovoid purple capsule; very rare subalpine species. B. blindii

B. argenteum Hedwig. One of the easiest of mosses to learn to recognize, and found in all of the continents including Antarctica. It is a weed in sidewalk cracks, lawns and gardens, but also occurs widely in naturally disturbed areas. It rarely fruits in our region. The capsules are on relatively short setae, and ovoid, not cylindrical, and are bright red. A fruiting colony was found around the fountain area in the garden of the Hamill House, in Georgetown, but the colony was destroyed by the remodeling of the fountain. An old collection from Pingree Park, collected by Pammel in 1896, is the only other fruiting collection we have seen. B. argenteum is the type species of the genus Bryum.

B. blindii Bruch & Schimper. B. blindii is a unique species, easily recognized with only a hand lens. The stems are minute, only a few millimeters long, julaceous, with pale green or straw-colored leaves. It has very small, rounded leaves and a very short and broad ovoid purple capsule on a short red seta. We have two collections, from Grand Co.: Fraser River Valley at north base of Berthoud Pass, Arapahoe National Forest, T2S R75W S34, 3,100 msm. On steep gravelly slope just above roadside on east side of lowest hairpin turn, 14 Sept. 1986, Weber & Dahnke B-91852. The moss was discovered by a group of us who sat down there to eat our lunch! The site has recently been scoured by bulldozers and is now barren, but probably more populations might be found above this area, which was obviously disturbed. The second collection is from Summit Co.: Blue Lake Dam area, Monte Cristo Creek Valley between Mount Quandary and North Star Mt., 3,000 msm. Limestone substrates. Rare, on gravels at the base of the dam., 26 Aug. 2,000, Weber, Wittmann, & Spribille B-111126. Evidently these are the only records in the contiguous United States.

Evidently Bryum blindii favors disturbed gravel habitats at high altitudes. The Blue Lake Dam site should be more carefully studied for other rarities. We already have found Aloina rigida and Isopterygiopsis alpicola there. The gravels were deposited below the dam during its construction, and the gravelly flat may have improved the habitat for these mosses.

B. calobryoides Spence. This is a unique species, not to be confused with any other, although it has been misidentified as B. gemmiparum! In Colorado, it is possibly restricted to the Front Range, where it occurs from 6,500 ft. to 9,000 ft. on cliffs, boulders, and in gravelly trail-sides. The populations form very compact, low, bright green tufts, with cabbage-like terminal stems, which may sprout slender microphyllous stems. The rhizoids are densely papillose. The terminal leaves are only up to 1 mm long and broadly obovate or circular, cucullate, often with a small slender recurved apiculus. The costa is strong, brown or reddish. The median leaf cells are hexagonal, almost isodiametric, nearly uniform on the apical leaves. The lower leaves are narrower and up to 1.5 mm long, less cucullate and lacking an apiculus. In the lower leaves the median cells are similar, but more of the laminal cells are rectangular, especially near the margin. A leaf border is indistinct in the short leaves and well-developed on the longer leaves. Our collections are from Boulder, Fremont, Grand, and Larimer Counties.

B. julaceum Schrader ex Gärtner, Meyer, & Scherbius. Very easily distinguished by the very slender ropy, julaceous stems (Anomobryum filiforme, A. julaceum, or Pohlia filiformis). Our early collections were made at Summit Lake, Mount Evans, but recently we have found it at low altitudes (6,000 ft.) in lower Boulder Canyon, on stream boulders, mixed with Bryum pseudotriquetrum. Jamieson has collected it often in the San Juan Mts. It probably is not a rare moss, but it occurs in very small patches mixed with other mosses of seasonally wet places. The stems are bright green or reddish, and according to Crum & Anderson, they often produce red, ovoid gemmae in the leaf axils. Shaw (1982) discusses the reasons which Ochi (1981) used to place this in Bryum rather than Pohlia. "It is the long [leaf] cells that might indicate a relationship to Pohlia, but this trait is to be found rather frequently elsewhere in Bryum. In addition, the costa is shortly excurrent, and the leaf margins are entire (or vaguely serrulate), features uncommon in Pohlia." In our collections, the median leaf cells are about 5 x 30-40 m, not vermiform. Spence (in litt.) suggests that we might possibly have B. concinnatum (Spruce) Lindb. in Colorado, which differs from B. julaceum in having shorter leaf cells

B. lanatum P. Beauvois. A "white" moss, resembling B. argenteum but with long slender leaf tips. It is a plant of arid sites and definitely a native species, while B. argenteum appears to be a weed, probably mostly introduced and commonly found in cracks in sidewalks. It is very abundant on the Western Slope and also known from the eastern foothills.

B. nanoargenteum Spence, sp. nov. ined. We have only one collection: Montrose Co.: Paradox Valley, 4 mi E of Bedrock; on red soils, under Sarcobatus.

Gemmabryum has several well-defined sections which could as well be considered separate genera. As a genus, it consists of mostly tropical groups of the southern Hemisphere. in Australia it is represented by many often handsome species, while our most diverse and common genus, Ptychostomum, is poorly represented.

1a. Medium to large plants; leaves imbricate, not twisted when dry; rhizoidal tubers or bulbils sometimes present (2)

1a. Small plants; stems evenly foliate; leaves strongly imbricate when dry, slightly twisted when dry; stem very slender, with bulbils in the upper leaf-axils or rhizoidal tubers at the stem base or on long rhizoids in the substrate; weedy. Small, often bud-like or weedy plants occurring on disturbed soil in towns, in the foothills, and arid flats in the western counties; (4)

2a. Leaves stiff, rigid, strongly imbricate, dark green or golden-brown; distal laminal cells long, incrassate and vermicular, >6:1. Very common on irrigated granite sloping rocks in the outer Front Range. G. alpinum

2b. Leaves less stiff, loosely imbricate, light green; distal laminal cells shorter, not vermicular, mostly 3-6:1.

3a. Leaves distinctly concave, somewhat distant when wet, the apex broadly acute to obtuse; costa not reaching leaf apex or only percurrent; arista lacking; common on floodplains of the canyon country. G. gemmiparum

3b. Leaves flat or weakly concave; crowded, the apex acute to acuminate; costa short but distinctly excurrent; plants uncommon, montane to alpine. G. mildeanum

4a. Short-stemmed stoutish, often cabbage-like tiny plants, usually with large axillary bulbils that fall away easily when dissected. G. bicolor

4b. Very slender plants with red or brown rhizoidal tubers; weeds of cultivated or disturbed sites. (5)

5a. Rhizoidal tubers 50-100 m diam. G. klinggraeffii

5b. Rhizoidal tubers over 120 m long. G. subapiculatum

G. alpinum (Withering) Spence, ined. A beautiful species, forming large soft deep turfs trending downslope on smooth, sloping surfaces of irrigated granite outcrops, from the base of the mountains to the alpine. Only one collection was made on sandstone: Boulder Co.: On sandy thin soil, Laramie Formation, White Rocks, 8 mi NE of Boulder, 5,500 ft. alt., 18 May 1967, Weber B-27930.The plants are hardly anchored to the rock but are held together by eroded gravel and sand. The plants are bright green when fresh, but when dry they become golden-brown, with a metallic sheen. According to Crundwell & Nyholm (1964) the rhizoids always have small round gemmae attached, but this is not always easy to demonstrate, since the rhizoids are so mixed with detritus. The anatomy of the leaf does not suggest a Bryum because the cells are long and narrow, as in Pohlia! Although this appears to be a plant of the foothills, we have one alpine specimen, unusual in having bright red foliage, and leaves smaller (0.5-0.7 mm long). The material is very scant. Summit Co.: Blue Lake, on seeping rock terraces, 2550 m, Spribille 9912 (B-111475) !Spence.

G. bicolor (Dickson) Spence, ined. This is a small species with leaves no more than 1 mm long, broadly ovate, with the costa not reaching the leaf apex or only percurrent, somewhat resembling B. argenteum but lacking the hyaline leaf apex, or G. calobryoides because of its somewhat julaceous stems. However, leaves are dull, the stems are plump because a leaf axil encloses a single large green gemma with red base; the green leaf primordia are well-developed and arise from near the base and upward, curving over the apex. Collections: Boulder Co.: Eldorado Mountain, 29 April 2001, Weber & Wittmann B-111734; Larimer Co.: Soil at edge of sandstone slab, open Pinus ponderosa slope, 5,400 ft., Spring Canyon, 5.5 mi SW of Fort Collins, 25 Dec. 1971, Hermann 24576; San Juan Co.: Deadwood Gulch, 9,500 ft., 2 mi SSW of Silverton, Hermann 24397, !J. Shaw.

G. klinggraeffii (Schimper) Spence & Ramsay, ined. According to Nyholm (1993), this species is characterized by having multicellular globose gemmae on the rhizoids. She writes: "This species is easily recognized by its pale orange rhizoids and usually abundant small bright crimson gemmae with protuberant cells." We have two collections. The first, a specimen verified by Crundwell, from Logan Co.: South Platte River below Crook Check Station, 2 mi S of Crook, 3700 ft,. alt., on moist river bottom, with willows and cottonwoods, 30 May 1981, Rolston 81025. The specimen is a sliver of clay with a few scattered stems of Barbula unguiculata, and one or two stems of a very tiny Bryum, sterile and unpretentious. To analyze it for gemmiparous rhizoids would destroy the specimen. The second collection was separated from a packet containing Pterygoneurum subsessile, from Weld Co: Pawnee Buttes National Grassland; on friable sandstone erosion fan at base of West Pawnee Butte, NE of Keota, 5,000 ft. alt. Pale green sterile plants; brown spherical gemmae 20, 60 m diam produced on the most slender hyaline branches of rhizomes. 20 May 1971, Weber & Hermann B-37484b. The plant should be common at this site.

G. barnesii (Wood) Spence, ined. is shown as being in Colorado by Vanderpoorten et al (2002), as Bryum, but no specimen was cited. It is a species with several gemmae per leaf axil. However, the dot is at the same spot on the map as that for G. bicolor, which may have more than one gemma. Spence (in litt.) agrees that this is not likely to be found in Colorado.

G. gemmiparum (De Notaris) Spence, ined.. This is not a gemmiparous moss, nor is it usually sterile. The "gemmae" for which is gets its name are clusters of short shoots at the stem apex (which are not necessarily present!) It is a species of medium size, and is distinguished probably best at first sight by having oblong leaves that are not at all acuminate or attenuate into hair-points. They do not collapse on drying. The costa is strong and reddish. The border is well-developed but fades at the leaf apex. The leaf cells are elongate-hexagonal and with somewhat thickened walls. Capsules are usually well-developed, pyriform, without a long neck. The outer peristome is yellow, not divided along the mid-line, smooth except for the papillose apex; the endostome basal membrane is very high, and the cilia swollen at the nodes but not appendiculate.

This is a species of silty ground along flood plains, and particularly common in the canyon country of western Colorado. We have very few collections. Mesa Co.: Koel Canyon, Colorado National Monument, Weber et al B-112047; Moffat Co.: Yampa River near Deer Lodge; moist crevices and angles of sandstone in bottom of ravine, drainage way, and seepage, Flowers 8790 (B-55483).

G. subapiculatum (Hampe) Spence & Ramsay, ined. A weed moss, growing in urban settings, known so far from Boulder, but undoubtedly present elsewhere. We have excellent collections from Boulder Co.: U. of Colorado campus: alongside Denison Building. surrounding a manhole cover. Rhizoids brown, beautifully papillose (sculptured); Gemmae red, spherical, 150-160 m diam. See also B. klinggraeffii. This site, under a drain-pipe, was destroyed by construction workers in 2004.

G. mildeanum (Juratzka) Spence. Somewhat similar to G. alpinum, but the leaves loosely imbricate (2-2.3 mm long), pale to golden green, lanceolate, with little curvature to the margins, while in G. alpinum the leaves are 1.5 mm long, more ovate, the margins are curved. It also forms much more slender stems in looser tufts, and varies in altitude from montane to alpine; the common G. alpinum is a plant of the outer foothills. Collections: Clear Creek Co.: Summit Lake, Mount Evans, 14 July 1953, George Neville Jones 19957; Larimer Co.: near summit, Greyrock Mt., 7,400 ft., 10.5 mi W of LaPorte, Hermann 23649; Mesa Co.: Grand Mesa; 2 mi E of Mesa Lakes, 10,300 ft., Hermann 16521. Moffat Co.: Douglas Mountain, Dutch Oven Spring, Weber 101111, !Spence.

H. macrocarpum (Hooker ex Drummond) Spence. Shaw (1992) reports this from the San Juans (Red Mountain Mine area; this was called Mielichhoferia macrocarpa. Bryum nelsonii Grout is synonymous with M. elongata). Jamieson has it from Cascade Creek Trail at Engine Creek Falls, on the damp, deeply shaded face of an eroded limestone grotto behind the plunge basin of the foot of the falls (Jamieson 13703). Anent M. macrocarpa (Hooker) Bruch & Schimper, Grout (MFNA, Vol. 2:261) claimed that Bryum nelsonii Kindberg, type from Gunnison, collected by N.L.T. Nelson, is M. macrocarpa, but the Gunnison County populations all belong to M. mielichhoferi and M. elongata. So do our collections from Summit County. Shaw et al (1992) mention all three species occurring together at Red Mountain. Utah specimens have come from limestone cliffs on Mount Timpanogos. This is an exceedingly rare and disjunct species, described first from Disko Island, and known also from Greenland and a few localities in western Canada.

L. pyriforme (Hedwig) Wilson. Frequent in disturbed soils, often on burned ground and peaty banks which have been quarried, definitely a moss of weedy tendencies, mostly found in the middle and higher altitudes. The combination of filiform leaves and nodding, pyriform capsules, is definitive. However, when not bearing capsules, the moss may be overlooked or be a puzzle for beginners. Here are some pointers: The leaves are very slender, but not secund, just somewhat spreading in all directions. The costa is quite wide and the leaf cells are slender but blunt, not long tapering at the ends. The margin may be slightly denticulate. Farther back from the apex of the shoot, the leaves are rather suddenly very small and broad, and the stem is black! This plant when sterile certainly does not suggest a Bryum! In fact, Spence proposes moving this genus to the Meesiaceae!

1a. Leaves brilliant turquoise blue when growing, less than 1 mm long, ovate, the cells wide and transparent; costa slender; stems more or less julaceous, the leaves appearing blunt; rhizoids smooth. M. elongata

1b. Leaves yellowish-brown, more than 1 mm long, narrowly triangular-lanceolate, tapered to a point, the cells narrow and densely packed; costa strong; leaves not tightly appressed, and commonly somewhat secund; rhizoids densely papillose. M. mielichhoferi

M. elongata (Hooker) Nees & Hornschuch. This is extremely abundant around the mines of Red Mountain Pass, in San Juan County. Carpets of it may seen from the highway! Although the individual stems of this moss are very tiny, the effect of them in the mass is overwhelming. When green, the great masses of Mielichhoferia that cover the ground of the mine tailings can be seen from the main highway as a turquoise-colored carpet! The leaves are ovate, green, minute (<1mm), appressed to the slender (up to 0.3 mm wide) stems; stems without rhizoidal mat; rarely fruiting, the capsule elongate-pyriform, the operculum conical; inner peristome well-developed, papillose; plants found only on substrates containing copper. A critical study of the two species (Shaw & Rooks 1994) was concerned with the possibility of intergradation between them. A second site is in Gunnison County, where Hermann collected both species, of which Hermann 26235 (B-57071) contains parts of two polsters, the largest one being M. elongata.

M. mielichhoferi (Funck) Loeske. Our major locality is in the vicinity of Red Mountain in the San Juan Range of southwestern Colorado where both species grow together. We have other collections from limestone cliffs: Gunnison Co.: 3,200 msm, just below Schofield Pass, July, 2001, Lehr 111886, Aug. 2, 1976. Gradstein B-56659, 21 July 1976, Hermann B-9494. One collection is particularly exciting because Hermann 27235, same place and date, consists of two pieces, one of which is M. elongata! A general account of the copper mosses is given by Persson (1956) and Shacklette (1967); the ecology of this species in Colorado was discussed by Hartman (1969). The spelling "mielichhoferiana" cannot be used because it was published a few months later than "mielichhoferi."The papillosity of the rhizoids was first noted by Nyholm (1993).

Spence, ined. proposes transferring this genus to the Mniaceae or "Pohliaceae".

November 9, 2004

Plagiobryum is an exclusively alpine genus, not too well separated from Bryum sens lat. However, sterile plants of P. zierii are easily recognized by the pale pinkish leaves; when present, the sporophytes are elongate and horizontally oriented, with a very long neck. Long-capsuled species of Pohlia, which have narrow elongate cells in the distal portion of the leaf and the leaves are deep green. P. demissum also is unmistakable if it has capsules, for the seta is stout and curved, and the urn is widest at the top (bloated, and the operculum is not continuous with the axis, looking somewhat like a species of Funaria or Meesia. Sterile plants tend to have a plump rosette of broad leaves that usually are quite reddish or brown.

la. Seta up to 2 cm long; capsule 5-7 mm long, curved and more or less horizontal, with the shriveled neck about as long as the oblong urn, more or less horizontal; sterile branches elongate, julaceous, with broad, concave, pink to silvery leaves. P. zieri

1b. Seta less than 5 mm long; capsule 3 mm long, horizontal from the bend in the short seta, narrowly cylindric, only slightly curved, the neck not much differentiated; sterile branches short, with ovate-lanceolate, more spreading leaves; new leaves green, the older ones brown. P. demissum

P. demissum (Hoppe & Hornschuch) Lindberg. Superficially similar to Ptychostomum pendulum, which grows in the same places, but the capsule is asymmetrical, as discussed above, not ovoid, and not pendent. Unlike P. zierii, this species forms very dense sods, often on rather hard turf. Rare or infrequent, in crevices of tundra rocks, on hummocks of solifluction lobes, in soil pockets of screes, etc., in the alpine, 12,000-13,500 ft. alt. Two collections: Clear Creek Co.: Summit Lake, Mt. Evans, sterile, Weber & Wittmann B-114282; McClellan Ridge above Santiago Mine, 12,500 ft. alt. fruiting, Weber, Koponen, & Nelson B-41407.

Several specimens collected as this species proved to be Pohlia greenii Bridel. Upon examination, their sporophytes are not asymmetrical, the operculum is terminal, the urn narrowly cylindric, not bloated, the leaves are lanceolate, distally denticulate, lack differentiated marginal rows of cells, and the distal leaf cells are narrow and vermiform. This minute Pohlia is a quite common component of the tundra flora.

P. zieri (Hedwig) Lindberg. Extremely rare species of subalpine seeping cliffs and moist tundra. The stems are usually tightly clumped, with broadly ovate, julaceous, delicate leaves with slightly recurved points. Their color is pale green, becoming pinkish in older parts. The cells are easily seen with a strong lens, and are swollen, reminiscent of a bowl of cut glass! We have specimens from Clear Creek Co., Summit Lake, Mount Evans, at 3900 meters, Hermann B-40082, Boulder Co.: Cirque above Rainbow Lakes, Weber & LaFarge B-43743, San Juan Co.: Lower Coal Creek along Lime Creek road, Jamieson & Schofield 10824, and Summit Co.: Blue Lake, Weber & Wittmann B-111138. The sporophytes are rare (our collections have a single abortive one!).

Ref: Shaw (1982). Spence, ined. *** suggests this genus be moved into a new family, the Pohliaceae.

How is Pohlia distinguished from Bryum? 1. Lanceolate, often serrate, leaves, usually without borders. 2. Costa ending before or in the apex. 3. Long, generally narrow and uniform leaf cells. 4. Peristome is double and the endostome is well-developed although cilia, when present, generally lack appendages. Because most of our species are usually without capsules, this is a rather iffy set of distinctions! But we seem to be stuck with them. Spence points out that the genus has a rather definite "Gestalt"; the plants and leaves have a stiff, rigid "hygrophobic" aspect shared with Mnium.

1a. Plants bearing axillary gemmae (2)

1b. Plants without axillary gemmae (8)

2a. Gemmae arising singly in the leaf axils, cylindrical to oblong, reddish when moist; leaf primordia green and conspicuous, scattered on the body and at the apex. P. drummondii

2b. Gemmae arising in clusters (3)

3a. Gemmae spherical to short-oblong, hardly longer than wide (4)

3b. Gemmae oblong, obconic or more or less elongate or vermicular (5)

4a. Leaf primordia of gemmae triangular-laminate (leaf-like), forming a dome over the apex. P. bulbifera

4b. Leaf primordia of some or all gemmae peg-like (just 1-2 cells broad at the base), erect or incurved over the apex. P. camptotrachela

5a. Gemmae small (about 70-120 m) and orange-red, with the primordia peglike and incurved. P. camptotrachela

5b. Gemmae larger, oblong, obconic, or linear-vermicular (6)

6a. Gemmae very large (200-500 m long), oblong to linear; leaf primordia laminate, at least 3-4 cells broad at the base (7)

6b. Gemmae small (up to 100 m long), linear-vermicular; leaf primordia peg-like, 1-2 cells broad at the base. P. proligera

7a. Gemmae oblong or obconic. P. andalusica

7n. Gemmae linear-cylindrical. P. tundrae

8a. Leaf cells broad (13 m or more wide) (9)

8b. Cells narrow, 6-10 (-12) m wide (10)

9a. Plants dull whitish, in calcareous seepy or wet areas. P. wahlenbergii

9b. Plants yellow-green, reddish or blackish at least below, in calcareous or acidic, usually wet, sites. P. obtusifolia

10a. Cells with thin to firm walls (11)

10b. Cells thick-walled; leaves ± serrate at the apex (14)

11a. Plants whitish, with the leaves often erect and seriate (12)

11b. Leaves shiny or dull green, the leaves sometimes erect but not whitish (13)

12a. Leaves erect and often seriate (the shoot appears pentangular). P. bolanderi var. seriata

12b. Leaves more or less spreading, not seriate, broadly lanceolate to elliptic. P. cruda

13a. Leaves long-lanceolate, erect, coarsely serrate near the apex. P. longicolla

13b. Leaves broadly lanceolate, spreading to erect-spreading, serrate or serrulate near the apex. P. cruda

14a.. Rare plants of wet roadside ditches and swampy ground, subalpine; rarely fertile; new growth bright yellow-green, the older growth black; cushions often deep. Leaf cells very broad for a Pohlia. P. obtusifolia

14b. Very common plants of various habitats, usually in forests, usually fertile; leaves dark green. P. nutans

1a. Endostome segments narrowly or not at all split along the keel; cilia rudimentary or absent, rarely long (2)

1b. Endostome segments broadly split along the keel; cilia long (5)

2a. Plants dull; leaf cells thick-walled (3)

2b. Plants ± glossy; cells thin-walled (4)

3a. Capsule 4-7 mm long; basal membrane ½-1/2 the exostome length; segments keeled; cilia present. Alpine plants. P. greenii

3b. Capsule 2-4 mm long; basal membrane to 1.3 the exostome length; segments scarcely keeled, often irregular; cilia lacking; forest plants. P. elongata (not yet found here but expected)

4a. Plants dioicous; leaves serrulate to serrate at the apex. P. bolanderi

4b. Plants paroicous; leaves coarsely serrate at the apex. P. longicolla

5a. Basal membrane low; segments narrow, scarcely keeled though broadly perforate; cells lax and thin-walled, 10-20 m wide. P. obtusifolia

5b. Basal membrane ½ the exostome length; segments broad and keeled; cells linear, 6-10 m wide (6)

6a. Leaves dull; cells with thickened walls. P. nutans

6b. Leaves slightly to strongly glossy; cells with thin walls. P. cruda

P. andalusica (Höhnel) Brotherus. A gemmiparous species occurring on disturbed sites in the subalpine. It has a slender, glossy appearance and very large green gemmae (200 m or more) arising in clusters from its upper leaf axils, tapering to the base, with 3-6 laminate primordia. We have collections from Clear Creek Co.: 7 mi SW of Georgetown, 13,000 ft., Dry, rocky tundra below Argentine Pass, Hermann 29065 (B-80209) (distributed as P. annotina), and Gilpin Co.: Weber & Wittmann 110942.

P. bolanderi (Sullivant) Brotherus var. seriata Shaw. An alpine species found in snow-melt areas. The type locality is on the road to Corona (Rollins) Pass. It is a dull, whitish species in dense tufts. The leaves are ± distinctly arranged in 4-5 vertical rows. P. bolanderi has recently been discovered relictual in Europe for the first time, in southern Spain at 2-3,000 meters, on Mulhacén,, the highest peak in the Iberian Peninsula (Shaw et al. 2004).

P. bulbifera (Warnstorf) Warnstorf. A gemmiparous species of disturbed places where the soil is compacted and hard. The gemmae in the leaf axils are spherical, with the laminate leaf primordia closing over the top as a dome. It is not necessarily an alpine species but found along slumping forested slopes. However, we have a fine collection from Summit Co.: McCullough Gulch, 12,450 ft.; below a saddle, on a rocky, steep slope near a snow bank, 16 July 2002, Redner B-112718.

P. camptotrachela (Renauld & Cardot) Brotherus. A gemmiparous species characterized by its dull rather than glossy appearance, and gemmae that are roughly spheroidal and less than 150 m diameter with 1-4 peg-like leaf primordia at the apex that are incurved over the apex. It occurs in the forested areas along unstable soils.

P. cruda (Hedwig) Lindberg. Not as common as P. nutans and found in more moist sites. The pale green leaves with an iridescent sheen are easily recognizable. It has long-cylindric capsules and well-developed endostome with broadly perforate segments and long cilia.

P. drummondii (C. Müller Hal.) Andrews. Evidently a frequent species on tundra, but rather difficult to recognize unless the large red gemmae with 3-6 laminate green leaf primordia are present, scattered mostly at the apex.

P. greenii Bridel. A very small species of peat hummocks at the edges of subalpine fens. The capsules are very small and not strongly inclined. Probably this is common in subalpine and alpine fens. We have it abundantly at Guanella Pass (Weber & Wittmann 112510). Shaw considers this to be a variety of P. elongata. However, from his discussion (below), we feel that it is probably justifiable to recognize var. greenii (Bridel) Shaw at the species level until more is known about the variability of both. Shaw wrote: "Rock crevices in tundra areas, less frequently on exposed soil, Alaska, Washington, California, Colorado, Utah, Newfoundland; Greenland. Reported from northern Europe. . . . Although no single character is diagnostic, several morphological attributes distinguish the var. greenii from so-called typical expressions: smaller overall size, shorter, broader leaves, more strongly inclined, broader, shorter capsules, low basal membrane, irregular segments, and complete lack of cilia. . . . Sporophytic and gametophytic differences between the varieties, coupled with geographic differences, warrant separation, at least on the basis of North American plants. Unlike P. spectabilis, the var. greenii differs from the var. elongata in more than just capsule shape, and notably so in endostome morphology. P. carneoides Dixon & Badw.*** of the Himalayas represents the var. greenii at its extreme. The plants are minute, with capsules hardly longer than wide." Shaw's figure 5, p. 136, shows these features very clearly.

P. longicolla (Hedwig) Lindberg. A robust, shiny green plant occurring on soil of road banks and streamsides at high altitudes. It fruits frequently. The capsules are elongate-cylindrical, often with a narrower neck. We have found it in peatlands in Boulder County.

P. nutans (Hedwig) Lindberg. Copper Wire Moss. A very common moss in shaded areas throughout the mountains, except for alpine tundra. It usually fruits abundantly, with hanging, oblong capsules. The thick-walled cells give the plant a dull appearance. When sterile, it is difficult to distinguish from P. elongata, which, fortunately, is rare or very infrequent and confined to the foothills

P. obtusifolia (Bridel) L. Koch. When sterile (the usual case), this is often a very large plant up to 4 cm tall. The leaves are somewhat cucullate, with broad cells (12-13 m or more wide, suggesting a Bryum). In these tall forms the plants are black except for the short apex of fresh, pale green leaves. We have found it along ditches along roads in the subalpine where it receives snow-melt water in the early spring. In an iron fen in upper Geneva Park this moss forms extremely tight, dense, black mats on the edges of limonite sheets.

P. proligera (Lindberg ex Breidler) Lindberg ex H. Arnell. A gemmiparous species of roadsides and paths in the forested regions. The gemmae are found in dense clusters in the upper leaf axils. They are linear-vermicular to narrowly oblong, with 1-2 peg-like leaf primordia often bent at an angle.

P. tundrae Shaw. A gemmiparous species of the alpine tundra that has narrowly cylindrical gemmae with laminate leaf primordia. Collections verified by Shaw are from Boulder, Chaffee, Clear Creek, Grand, Gilpin, Pitkin, and San Juan counties. They were originally identified as P. bulbifera, P. proligera, and P. annotina var. loeskei. A Gilpin County collection (Weber 3471) contains P. andalusica, P. cruda, and P. tundrae mixed in the same tuft!

P. wahlenbergii (Weber & Mohr) Andrews. Characterized by whitish or dull green erect-spreading leaves on long, slender stems. It resembles Bryum weigelii but is never reddish nor with decurrent leaves. It is a beautiful moss when fresh and grows in dense, tall tufts in seeps at high altitudes. It has also gone under the name of Pohlia albicans.

Ptychostomum is the most diverse genus, with a number of fairly-well-defined subgenera; these are indicated in parentheses)

1a. Stems mostly evenly foliate, not comose; leaf base same color as rest of leaf, usually green; limbidium indistinct, bistratose; distal laminal cells lax, wide and short, 2-4:1, rhomboidal; proximal laminal cells longer and narrower, rectangular (2)

1b. Stems evenly foliate to comose with upper leaves distinctly enlarged; leaf base mostly reddish; limbidium distinct to indistinct, unistratose; distal laminal cells not lax, 3-5:1, rhomboid to hexagonal; proximal laminal cells similar in width and length but rectangular, occasionally quadrate (7)

2a. Plants medium to large, dioicous; peristome mostly perfect; spores small (<22 microns) (Pallentia group) (3)

2b. Plants small to medium, monoicous; capsule elongate, with a long neck, curved, peristome reduced; cilia short to absent; spores generally large (>25 microns) (Amblyophyllum group). P. cernuum

3a. Leaves strongly and broadly decurrent; costa percurrent; leaves generally pink in living plants, but this color does not persist in herbarium, specimens; never with capsules. P. weigelii.

3b. Leaves not or weakly decurrent, obtuse, acute, or acuminate; costa not reaching apex to excurrent (4)

4a. Leaves broadly ovate to suborbicular; leaf apex rounded to obtuse; costa not reaching apex to percurrent (5)

4b. Leaves ovate to ovate-lanceolate; leaf apex acute; costa typically percurrent to short-excurrent (6)

5a. Plants often red; leaves strongly concave, not strongly contorted when dry; costa often percurrent; very rare alpine. P. cryophilum

5b. Plant dull green, lacking red tints; leaves weakly concave or flat, very strongly contorted when dry; costa not reaching apex; rare, mostly in subalpine peat fens. P. cyclophyllum

6a. Capsules present, short-pyriform, constricted under the mouth; alpine. P. turbinatum

6b. Capsules not present in Colorado material, longer; subalpine; young leaves pink- or red-tinged; leaf margins recurved. B. pallens

7a. Plants small, densely caespitose; leaves ovate-lanceolate, limbidium indistinct or absent; proximal laminal cells shorter than median and distal cells; dioicous. (Caespitibryum group) (8)

7b. Plants medium to large, caespitose; leaves broadly ovate, ovate-lanceolate to lanceolate, flat or concave; proximal laminal cells same length as distal ones or somewhat longer; sexuality various (Pseudotriquetra group) (9)

8a. Plants very small, <1.5 mm; distal lamina in older leaves becoming hyaline, the proximal laminal cells short-rectangular to quadrate; costa variable, from not reaching apex to excurrent in a slender hyaline hair-point. P. kunzei

8b. Plant medium, most leaves 1.0-2.5 mm long; distal lamina green; proximal laminal cells long rectangular; costa strong, excurrent in brown or yellow hair-point. P. angustifolium

9a. Plants comose to caespitose, monoicous; peristome reduced, the cilia short or absent; basal membrane low, mostly ½ the height of the exostome; spores large, 18-50 microns (Penduliformia group) (10)

9b. Plants evenly foliate to loosely comose; sexuality various; peristome perfect; basal membrane > half the height of the exostome; spores small (10-22 microns) (12)

10a. Endostome strongly adherent to exostome, giving exostome teeth a chambered look; leaves acuminate, costa excurrent in a short arista; capsule short-pyriform only 1 mm long, not constricted at the mouth; operculum sharply conic; spores large (22-35 microns); alpine. P. pendulum

10b. Endostome not fused to the exostome; leaves acute to acuminate; costa not reaching apex or long excurrent; capsule various; spores 25-50 microns (12)

11a. Leaves lanceolate, not concave or keeled; arista long excurrent, denticulate; mouth of mature capsule red; spores small, 18-28 microns; endostome membrane about half the height of the exostome; cilia rudimentary. P. amblyodon

11b. Leaves ovate, strongly concave, keeled, acute; costa percurrent to very short excurrent; mouth of mature capsule yellow or pale orange; endostome membrane low, less than half the height of the exostome; spores 25-50 microns; cilia absent. P. knowltonii

12a. Stem elongate; leaves distinctly decurrent; limbidium strong; costa prominent but not with a long arista (13)

12b. Stem short, or if elongated, the stems tightly packed; leaves not decurrent; limbidium weak, arista well-developed (14)

13a. Leaves long-decurrent and usually remotely spaced on the stem; dioicous; long-stemmed plants of fens. P. pseudotriquetrum

13b. Leaves inconspicuously decurrent and closely spaced, overlapping on the stem; synoicous; relatively short-stemmed plants of wet streamsides. P. bimum

14a. Stems elongate, branched (the branches ascending with the main stem) forming deep sods bound by rhizoids, with the annual growth often very clear by the difference of color of the old leaves of succeeding years; leaves of the season turquoise green, the arista weak and not darkened; spores about 20-22 microns. P. pallescens

14b. Stems short, unbranched, not forming sods; leaves of the year darker green, often brownish, the arista strong, brown; spores 12-15 microns (15)

15a. Limbidium narrow, yellowish, 2-3 cells wide; plant synoicous. P. creberrimum

15b. Limbidium wide, 3 or more cells wide, the same color as the lamina; plants polyoicous. P. lonchocaulon

P. amblyodon (Müller-Hal.) Spence. This species occurs commonly in the subalpine wetlands and alpine tundra, where it may replace B. pallescens on drier sites. This has been known as Bryum stenotrichum. Although the species is characterized technically by having no cilia and a low endostome membrane. I believe it can be recognized on sight. The leaves are very long and narrow, loosely spreading and twisted, with a long pale excurrent costa, which on close examination is denticulate. The innovations are weakly julaceous and their leaves imbricate (one synonym is Bryum imbricatum). The limbidium is prominent and revolute from leaf base to apex. The capsule is straight and elongate.

P. angustifolium (Bridel) Spence & Ramsay. A weedy moss, usually sterile, and common in sandy soil on the eastern plains and outer foothills. It probably does not occur in the high mountains. It is a plant with very low stems, only a few millimeters tall, forming dense turfs. The yellowish leaves are broadly ovate, with intensely red broadly rectangular basal cells (often with conspicuous oil droplets).The costa is long excurrent into a hair-point. The species is dioecious, and the male plants occur mixed in with the female sporophyte-bearing ones. The (smaller) male plants terminate with a bulbous cluster of leaves with flaring points. It has long been called, incorrectly, Bryum caespiticium.

P. bimum (Schreber) Spence. This has usually been considered a variety of P. pseudotriquetrum, but the leaves are very shortly and faintly decurrent, and not very widely spaced along the stem The habitat is quite different from P. pseudotriquetrum. Instead of being a fen-plant, it forms clumps on stones along running streams and is never a tall plant. It is not a rare plant and ranges from the alpine tundra down into the lowest foothills. Spence has verified one collection from Colorado: Boulder Co.: 2 mi NW of Lyon; Spring Gulch, 5,600 ft. alt., N402458 W1051742; on rock along granite-lined stream-bed; plant bright green when moist, 7 April 2002, Weber & Wittmann B-112397. Actually, most of our Colorado collections seem to fit this species or variety, while the elongate-stemmed P. pseudotriquetrum appears to be found only in fens.

P. cernuum Hornschuch. We have a few collections, from Clear Creek Co.: Clear Creek canyon, Weber & LaFarge B-43499, verified by Jon Shaw; Gunnison Co.: Cement Creek canyon, wet cliffs in the Narrows, 9,400 ft., N385229,W1064750, 14 Aug. 2000, Weber, Wittmann, & Lehr B-111062; Montezuma Co.: Pine Canyon, S end of Moccasin Mesa, Mesa Verde, Welsh & Moore 2129. This species grows in places subject of light flooding or dripping rocks. It may be associated with Distichium capillaceum. It is very easily recognized by its very narrow elongate capsule. The leaves are very unusual, being well-separated on the stems, erect-spreading when wet and shriveled and incurved when dry. The cells are quite short and broad, and the margin is strongly bordered and narrowly revolute. There is a superficial resemblance of the capsule to that of Pohlia elongata and our specimens have been misidentified as such (Bryum uliginosum).

P. creberrimum (Taylor) Spence & Ramsay. This is evidently a very common species, going under various names in the herbarium (B. cuspidatum, B. lisae). It is most closely related to P. pallescens, but can be distinguished by the special key. Well-developed plants have very short, unbranched stems, and the leaves are less appressed, with stouter excurrent costae, and the leaves are more brown, whereas in P. pallescens the leaves of the season are almost turquoise.

P. cryophilum (Mårtensson) Spence. A very distinct alpine species, disjunct from the Arctic. The plants are delicate, often strongly reddish, the leaves are rounded-oval, concave, not strongly contorted in drying. Collections: Boulder, Clear Creek, Larimer Co.: On edges of pools and springs, Niwot Ridge, Summit Lake, Trail Ridge.

P. cyclophyllum (Schwägrichen) Spence. A very rare species occurring in peatlands. We have one verified collection: Boulder Co.: Peat bog source of Left Hand Creek, 10,600 ft., 3 mi W of Ward, 9 Sept. 1961, Weber & Dahl B-6983, !Spence, 2002. The site has since been destroyed by peat mining. The very slender stems, dull, bluish-green, oval leaves, rounded-obtuse at the apex, with the costa not quite reaching the leaf apex, remotely scattered on the stem and greatly contorted when dry, and the occasional presence of cylindrical, filamentous-branched, gemmae in the upper leaf axils, characterize this species.

P. knowltonii (Barnes) Spence. An infrequent species of the moist tundra. Collections: Jackson Co.: A little west of the Roxy Ann Lake turnoff from the Rainbow Lake Trail approaching the Continental Divide, 1 mi N of Mt. Ethel, 3400 msm., N404000 W1064035; on boggy flats, alpine tundra, 8 Aug. 1981, Rolston 81106 (B-80243) !Crundwell, Lawton; Clear Creek Co.: Seepage in granite bluff beside St. Mary's Glacier, 11,000 ft., 6 mi NNE of Empire, 31 July 1977, Hermann 28028. An especially fine collection, sterile but consisting of deep sods, is Summit Co.: Blue Lake Dam area, Monte Cristo Creek Valley between Mt. Quandary and North Star Mt., 3,000 msm, Ten Mile Range; on limestone terraces irrigated with snow melt, 16 Sept,. 2000, Weber & Wittmann B-111211.

This is a very distinctive moss because of the extremely low basal membrane and the extremely slender endostome segments and totally lacking cilia, but it is easily recognized even when sterile. The stem and leaves are not minute and resemble those of B. pallens but the leaves are concave and cucullate, bowed at top and bottom, standing close to the red stem. The leaves are broadly oval, rounded or obtuse at the apex, not decurrent, with a very stout, percurrent costa, very indistinctly bordered and strongly recurved in the lower half. The median cells are 3-5:1. The rhizoids are densely papillose.

P. kunzei (Hornschuch) Spence. This species is a miniature version of P. angustifolium. The plants have very slender stems and form dense, often deep sods, and the leaves are very small, only 0.5-1.5 mm, smaller than the closest relative, P. angustifolium. They may or may not have an excurrent costa. It appears to be restricted to the canyon lands of the western tier of counties. In Utah is is common on the flats around Great Salt Lake. In Colorado its habitat is on cliffs and rocks, whereas P. angustifolium is a very common weed moss on soil. Collections: Mesa Co.: Colorado National Monument: Kodel Canyon, just west of Fruita entrance; on sandstone rocks along lower part, up to the exposed Pre-Cambrian cliffs, 12 May 1981, Weber, Rector, & Siplivinsky B-81042. Montezuma Co.: just west of bridge crossing the San Juan River along Hwy north of Four Corners Monument; on dry soil and soil over rock on south side of river, about 30 meters west of bridge, 26 Dec. 2003, Spence 5629.

P. lonchocaulon (Müller Hal.) Spence. This is so similar to P. creberrimum that it can easily be overlooked. It should be sought at low elevations in the western counties. We have one collection: Montezuma Co.: Mesa Verde National Park, Chapin Mesa, seep draining water tank, 9 June 1963, Welsh & Moore 2113 (COLO B-55123), verified by Spence, 2005 (previously determined by Flowers as Bryum angustirete Kindberg)

P. pallens (Swartz) Spence. A species of wet ground in subalpine willow carrs and pond shores. In this the leaves are broad up to the apex, and either obtuse or with a slight one-celled apiculus. Immature sterile plants may have very small pink, concave leaves, 0.5-1.0 mm, appressed to the stem, unchanged in drying, while fertile plants have longer leaves (up to 2 mm). The areolation is very transparent, the leaf margin narrow. The leaves are not at all decurrent. The capsule is about 2 mm long, plump pyriform. The peristome is pale yellow, the endostome segments narrowly perforate, cilia are short, not appendiculate. Sterile plants are not hard to recognize. The distantly spaced leaves that are not much changed on drying, the abruptly short-acuminate ovate leaves that are not at all decurrent, the short and broad, clear leaf cells an the red stem are good characters.

P. pallescens (Schleicher ex Schwägrichen) Spence. One of the most common Brya of wet ground, especially at lower altitudes. The tufts are often very deep, and the stems tightly packed so as to obscure the branching, which is erect. The current year's shoots have pale green leaves that have long excurrent tips but they are not stout or reddish. The colonies often form distinct tiers of annual growth, and the lower stems densely clothed with brown rhizoids. Commonly the leaves of previous years are bleached, and the seta of previous years' sporophytes persist. The median leaf cells are short, about 3:1, and the limbidium is prominent. The costa is strong and excurrent, and red at the base (but the leaf base itself is not red). Sporophytes are usually abundant. The capsule is cylindric, with a distinct narrow and shrunken neck about as long as the urn. The exostome is yellow, and the endostome segments have very large and broad gaps in the middle; cilia are very well developed and appendiculate. This species is similar to P. creberrimum but has larger spores (18-22 microns).

P. pendulum Hornschuch. Locally abundant on moister parts of tundra. It is usually abundantly fertile and, except for this, might be overlooked because of its small size. The leaves are only 1 mm long, in a densely crowded rosette, broadly ovate, abruptly aristate, red-based, widest at the middle, the median cells about 40 x 15 microns. The capsules are short, fusiform-ovoid, (if opercula are present the capsules are pointed at each end). The seta is curved just below the urn, and the peristome teeth are unique. The endostome is adherent to the exostome. The lamellae (divisions separated by horizontal walls) have several cross-walls that are irregularly widened and not transverse (showing diagonal and circular lines); this is most easily seen from the ventral side. In age the capsules are white. It can hardly be confused with any other alpine species, being recognizable on sight without having to examine the peristome.

The habitus depends a good deal on the substrate and growing conditions. On gravelly polygonal ground the plants are extremely short with the stems crowded at the summit of the extremely short stem (B-16824 from Summit Lake, Mount Evans), On well-developed moss tundra the plants form a relatively deep mat with several growth periods apparent as in P. pallescens (B-16827 from Niwot Ridge).

P. pseudotriquetrum (Hedwig) Spence & Ramsay. A common species of wet places from the foothills to the alpine, not usually occurring in large pure stands but consisting of few scattered stems in amongst other wetland mosses. It is handsome, with quite large leaves distinctly decurrent on the stem. It does not seem to fruit regularly here. See discussion under P. bimum.

P. turbinatum (Hedwig) Spence. An uncommon species thus far found only in the high alpine area. This species is distinguished by the short-pyriform (turbinate) capsule with a small diameter mouth, the urn constricted under the mouth, somewhat resembling that of P. pendulum, but the peristome is not unusual for the genus. Collections: Boulder Co.: Green Lakes Valley, 10,500-11,500ft., on rocks in snow melt drainage, 10-12 Sept. 1972, Weber B-41730, 42005.

P. weigelii (Sprengel) Spence. A very unusual species with very distant leaves, growing in springs and half-submerged in willow fens of the subalpine. It is always sterile, and may be mistaken for Pohlia wahlenbergii, but the leaves of B. weigelii are greatly shrunken and contorted when dry, and much more strongly decurrent.

Ptychostomum sp. indet., from seeping granite cliffs on Flagstaff Mountain, Boulder, Weber & Wittmann B-113917, which we had tentatively suggested might be "B. nano-argenteum", sp. nov. ined. Spence comments (13 Nov. 2004) as follows: "What you have is a mystery, although I have several other collections of it from various places. It is somewhat reminiscent of P. kunzei, but the hair point is shorter and not so obviously hyaline. The leaves are more like G. bicolor. Most specimens have medium-sized red rhizoidal tubers, ca. 100-200 m across. This puts it in among the "Microerythrocarpum" group, near but not G. subapiculatum. I think that it is undescribed. I have material from CA, NV, AZ, and UT as well as your collection. One characteristic of all collections: they occur in areas that are at least seasonally wet or seepy, on soil or rock."

Ref.: Vitt (1993). Note: Spence (in litt.) feels that this is a monotypic family, the Roelliaceae.

R. roellii (Brotherus ex Roell) Andrews ex Crum. This handsome moss occurs on the forest floor under spruces in the subalpine of Rocky Mountain Park and in Middle Park. It can be mistaken for nothing else but the large species of Rhizomnium. However, those species grow in wet places in the forest and have strongly bordered leaves without marginal teeth, contorted when dry, and the stems are clothed with rhizoids. The old report of Rhodobryum roseum (Porter, 1876), which is now referred to R. ontariense (Kindberg) Kindberg, probably represents a misidentification of Roellia. Rhodobryum has erect stems arising from horizontal underground stems, while Roellia lacks rhizomes.

Ref.: Spence(1996)

1a. Innovations rosulate, most leaves clustered at the stem apex; leaves obovate, flat when moist (2)

1b. Innovations evenly foliate; leaves ovate, concave when moist. R. flaccidum

2a. Leaves in a tight spiral, leaves appearing very narrow an dark green when dry, with a strog excurrent costa; gemmae lacking; stems lacking brown micronemata; fertile, capsule with a distinct neck R. capillare

2b. Leaves in a loose, irregular spiral, leaves not appearing very narrow when dry; filamentous red-brown gemmae in the leaf axils, sometimes very copious; stems with numerous micronemata. R. laevifilum

R. capillare (Hedwig) Spence. The tightly massed strongly curved and spirally arranged (as if brushed) rosulate leaves with stout aristae are very clearly different from those of R. laevifolum. Our specimens are fruiting, while R. laevifilum is usually sterile. The rhizoids are papillose, and the small rhizoidal gemmae are concolorous. Thus far we have only three collection: Boulder Co.: 8 mi NW of Nederland, North Fork of Middle Fork of Boulder Creek, 10,300 ft., Flowers 9833 (B-55251); Douglas Co.: Castlewood Canyon State Park, 6,400 ft. alt., Lederer B-102658; Larimer Co.: Big South Trail, under Alnus, bank of Cache La Poudre River, 8,700 ft., 2 mi. ENE of Chambers Lake, 19 Sept. 1977, Hermann 28159. This probably has been overlooked but likely restricted to the Eastern Slope foothills.

R. flaccidum Bridel. This appears to be a rather tiny relative of the next. It was first discovered in Colorado by Carl Darigo, who found it "in the developed area of a hotel in Boulder" where he assumed it to have been an introduction. We now know it is an indigenous species: Boulder County: Marshall Mesa Open Space, on slopes of northwest-facing friable sandstone outcrops of Laramie Formation, in partial shade of Pinus ponderosa, 6,000 ft. alt., 28 Feb. 2004, Weber & Wittmann B-113853. It is also abundant on granite outcrops in the outer foothills. Also in Larimer and Jefferson Counties.

R. laevifilum Syed. A common moss in the outer foothills, characterized by the leaves with excurrent costae, distinctly twisted spirally around the stem when dry, the capsule that leans but does not usually become really pendent, and the characteristic green or brown filamentous and papillose gemmae that are usually found in clusters in the axils of the upper leaves. This species seems to prefer growing on thin soil over rock rather than on deep soils.

Bryum gemmilucens. There is a dot on the distribution map in Vanderpoorten & Zartman (2002) on the B. bicolor group, but we do not know the source of the report. A specimen from Weld Co.: Pawnee Buttes, 20 May 1971, Weber, Hermann & Arp B-37484b, belongs definitely to this group, but we have been unable to find the characteristic axillary gemmae.

Pohlia elongata. Our specimens have been re-examined and we find that they are Ptychostomum cernuum. However, there is no reason to believe that the species will not be found here, so we have left it in the key. The species differs from P. nutans in having a very narrow, elongate, not nodding capsule; the endostome segments are narrowly, rather than widely perforate.

May 8, 2003

A monotypic family consisting of the single genus, Bryoxiphium, with one or possibly two, species, depending on the taxonomic interpretation. Probably one of the rarest and most sought-after mosses in the world, each new discovery justifies a publication.

The moss is unique in its morphology. Arising from a small bulb-like base, the single stem has two rows of leaves. "Usually they are closely imbricated, erect, and rather folded, but at the same time they are so small that in a superficial observation they might be taken for appressed hairs. The lower leaves... are usually characterized by a blunt apex, and the longest leaves are clearly apiculate and tapering" (Löve & Löve 1953). The sporophyte is extremely rare, undoubtedly because male and female plants do not occur anywhere near each other. The shiny, light green or golden little stems with their unusual leaves are impossible to mistake for any other moss.

Bryoxiphium is undoubtedly a Tertiary relict, occurring as it does in highly disjunct areas of the world, including Iceland, Greenland, Japan, Kamtchatka, the American midwest, Mount Rainier in Washington, and Mexico, and now, most recently, in Colorado.

B. norvegicum (Bridel) Mitten. Sword Moss. Until recently, only one collection was known: Montezuma Co.: on overhang of cliff, ca. 6 mi NNW of Dolores along Beaver Creek, ca. 7,500 ft. alt., 0.5 mi NW of junction of Dolores River and Beaver Creek, 12 June 1958, Pursell 3246a (COLO B-65336). This evidently has been destroyed by high water level in the McPhee Reservoir. Jamieson has found a second colony in Conejos Co.: upper tributary of Navajo River, 11,500 ft. alt., Jamieson s.n. (COLO B-113241). Botanists desiring to find this species might well be guided by the Löve's observation: "Bryoxiphium seems to prefer a substratum porous enough to hold water and at the same time might seem to avoid calcareous soils. Sandstone or volcanic material might seem to be preferred in most areas." The leaves are so closely appressed that the stem looks like an undivided flat ribbon!

May 12, 2004

la. Capsule dark chestnut-brown, glossy, broadly ovoid, the face broad and flat, nearly perpendicular to the seta. B. aphylla

lb. Capsule green or becoming pale brown, not highly glossy, narrow, subterete and elongate, erect or merely oblique to the seta. B. piperi

B. aphylla Hedwig. Both of our species are very rare, and are found only in virgin spruce forests with a moist to wet and boggy floor. Both species have been found in the Gothic Natural Area, in a part of Colorado with a very high annual precipitation. The forest floor is laced with many seasonal run-off streamlets and with a tangle of down timber. B. aphylla occurs on mounds of clay soil probably raised by the falling of a tree and stabilized by algae and moss protonema, a very precise habitat and very locally developed. Buxbaumia species are usually associated with small liverworts such as Lophozia and are most likely to be found by one with eyes focused on them..

B. piperi Best. This species occurs on soggy, rotting wood and seems to be somewhat more plentiful than B. aphylla. Grand Co.: Middle Park; West St. Louis Creek, 6 mi W of Fraser, Weber & Vaarama B-11086; Gunnison Co.:Gothic Natural Area, 10,000 ft. alt., on rotting logs, Weber B-17070, Khanna 15328; San Juan Co.: San Juan Mts., Weminuche Wilderness, mesic spruce forest near confluence of Vallecito and Trinity Creeks, T39N R6W NE1/4 Sec, 4, 10,600 ft., Jamieson 11441, 11490 (herb. Jamieson). One specimen was associated with a few stems of Pohlia cruda, the other with minute plants of Leiocolea badensis.

1a. Leaves straight, strongly plicate; stem densely tomentose. Tomentypnum

1b. Leaves squarrose or slightly falcate, not plicate; stems not tomentose (2)

2a. Costa lacking or very short and double (3)

2b. Costa single, ending at or somewhat above the leaf middle (6)

3a. Leaves squarrose-recurved; margins revolute below; yellow-green; leaf bases closely overlapping. Campylophyllum halleri

3b. Leaves wide-spreading to more or less squarrose; margins not revolute (4)

4a. Plants slender, green, with creeping stems; leaves smooth, minute (0.5 mm), finely serrulate all around especially at the base; alar cells small and quadrate. Campylophyllum sommerfeltii

4b. Plants robust, with erect-ascending stems; leaves larger (2 mm), striolate when dry, entire; alar cells somewhat inflated, subrectangular (5)

5a. Plants usually erect; stem leaves 1.7-2.8 mm long, the acumen constituting 40-65% of leaf length. Campylium stellatum

5b. Plants usually creeping; stem leaves 1.0-2.3 mm long, the acumen constituting 55-77% of leaf length. Campylium protensum

6a. Leaves erect-spreading, gradually acuminate; alar cells enlarged and inflated, becoming thick-walled in age. Drepanocladus (Campyliadelphus) polygamus

6b. Leaves 1.0-1.2 mm long, wide-spreading to squarrose, abruptly acuminate; alar cells not or only slightly inflated. Campyliadelphus chrysophyllus

Campyliadelphus chrysophyllus (Bridel) Kanda. Forming dense golden-green patches on rocks along streams in the foothills canyons, but also in the upper subalpine, nowhere very common (Campylium chrysophyllum). Dioicous, never fruiting here. Archuleta Co.: T36N R1E S5, Lyon B-114201.Boulder Co.: Boulder Canyon, 7,000 ft., Weber B-10559; Eldorado Springs, Weber, Wittmann & Mazurek B-111741; Gregory Canyon, Weber & Wittmann B-112134.

Hedenäs (1990) wrote: "The new taxonomic treatment of Campylium s.l. species is suggested by features of the vegetative leaf costa, alar group structure, laminal cells (prorate or not), pseudoparaphyllia, exostome and endostome structure, as well as by the overall size of the plants and the habitat preferences of the species. The spreading, channeled leaf acumina cannot outweigh this set of characters and must thus have evolved several times among these taxa."

C. protensum (Bridel) Kindberg. Similar to C. stellatum. Hedenäs has seen two collections of this: Flowers 9820 and 9777, from Boulder Co.: 8 mi NW of Nederland, North Fork of Middle Fork of Boulder Creek, 10,260 ft., submerged in small brooklet, and on wet soil among herbage, 21 Aug. 1964. Grand Co.: Head of Willow Creek, trail to Lost lake, 8,500 ft., Weber, Wittmann, & Tidball B-112466.

This is ecologically quite distinct from C. stellatum. It grows on moist ground on the forest edge where water percolates from snow patches under the trees, and it also grows as this did, on a log, intermittently sprayed. The leaves are very small (less than 2 mm long) compared to the wetland C. stellatum and the leaf base more suddenly narrows to the very long acumen (B-112440).

C. stellatum (Hedwig) C. Jensen. Very common in wet ground of subalpine and alpine willow and peat fens. Easily recognized by the large (2 mm), stiffly-spreading leaves with a rich russet-gold tint. See Flowers, Plate 119:1-7 (According to Hedenäs, this is the type species of Campylium and cannot be called Campyliadelphus). He distinguishes this from the next as follows: When seen from above in the field, shoots of C. stellatum often look like small stars. It is closely related to C. protensum, from which it differs in its more erect growth, its slightly larger size, and its relatively shorter leaf acumen. Paraphyllia have never been seen in C. stellatum, whereas scattered plants of C. protensum have a few paraphyllia.

C. halleri (Hedwig) Kanda. A subalpine species growing on rocks. It has very minute leaves of a bronze color, is autoicous and usually fertile, and differs from C. sommerfeltii by having the leaf bases overlapping and not exposing the stem, and the very short or absent costa. Our only record of this is from San Juan County: Deadwood Gulch, Hwy 550, 2.2 mi SSW of Silverton, 2850 msm., Weber & Wittmann B-111922. However, the species is evidently very common in the an Juan Mountains (Jamieson collections).

C. sommerfeltii (Myrin) Hedenäs. An infrequent or easily overlooked species on rocks, rotting wood, or duff. near stream sides in the Front Range from the outermost base of the foothills to the montane (6,000-8,900 ft.) (Campylium hispidulum of Colorado reports).The plants are deep green to yellow-green, with minute strongly squarrose leaves, inserted with their bases not overlapping, The plants are autoicous and are usually richly fruiting. Our specimens are mostly from Boulder County: Bluebell Canyon, on trail to Royal Arch, Roehrick B-6111; bottom of Skunk Canyon, Weber, Nelson & Litvak B-35450; 2 mi S of Ward, 1 mi E of Peak-to Peak Hwy close to Four-mile Creek, 8,900 ft., Shultz B-46217. We have one collection from Montezuma Co. (Colyer B-113281).

Ref.: Hedenäs 1987), Faubert (2004 in BFNA tentative treatment)

T. nitens (Hedwig) Loeske. A handsome moss with very evenly pinnate branching in one plane. The branches are slender, with smaller leaves about a fourth as long as those on the single main stem. The ovate leaves are loosely appressed to the stem, lack a costa, and the elongate cells have high papillae arising from their side or end walls. Paraphyllia of a few filaments of elongate clear cells, either simple or once-branched, arise from the leaf bases rather than from the stem. Masses of branched brown rhizoids clothe the stems.

One of the most common and constant components of subalpine peatlands and willow fens, commonly occurring with Sphagnum and Helodium blandowii. In unusually cool, moist sites, the species may occur in ravines along streams in the foothills, down to 7,000 feet, but this is a rare situation. Hedenäs (1987) proposed putting this in the Amblystegiaceae.

This is a monotypic family, with only one genus and species.

C. nigritum (Hedwig) Bridel. This is a turf-forming plant of the boreal regions of North America and Eurasia occurring in Park Co.: South Park: High Creek Fen (a Nature Conservancy reserve), 10 mi S of Fairplay, 9,700 ft. alt., 29 June, 2000, Weber & Wittmann 112549. Clear Creek Co.: Mount Evans, Summit Lake, 12800 ft alt., 7 August, 2004, Weber, Wittmann, Lehr, & Tidball B-114276. In the West it is also known from Montana. The plant forms dense turf on the vertical sides of shallow channels in the center of the fen where the obvious associates are small Picea trees, Betula glandulosa, Salix candida, S. myrtillifolia, and S. serissima dotting the sedge carpet. The leafy shoots are undistinguished, with leaves resembling those of Ceratodon, but the sporophyte is unique in the mosses. The capsule is minute, orbicular, and bent to the side, resembling a miniature golf club.

C. dendroides (Hedwig) Weber & Mohr. An easily recognized species, being our only moss with a dendroid habit. The primary stems are creeping and embedded in the soil. Erect secondary stems are dark and clothed with appressed, often brownish or pale leaves, and branch at the top, the radiating branches suggesting the fronds of miniature palm trees. Climacium is a common moss of wet streamsides from the lower foothills on up to the subalpine. The plants are sterile in our area, since they are dioicous and only the female plants occur. The branch leaves are ovate, strongly irregularly plicate, with a prominent costa disappearing just below the tip. The margin is coarsely serrate at the apex, the median cells are narrowly rhomboid to linear, and the basal cells nearest the costa are thick-walled, pitted, and orange. Toward the basal margin the cells become thinner-walled, hyaline, and form a distinct area in the cordate-auriculate leaf base.

1a. Plants extremely small, less than 0.5 mm, difficult to see without a lens! Capsules erect, goblet-shaped. On sandstone cliffs, often on shaded under-hangs. Seligeria (Seligeriaceae)

1b. Plants larger (2)

2a. Leaves distichous, lying in one plane when wet, with a broad, sheathing base suddenly narrowed into a divergent, filiform lamina consisting mostly of the costa. Distichium (Ditrichaceae.)

2b. Leaves spirally arranged around the stem, not lying in one plane (3)

3a. Cells of the distal half of the leaf papillose or mammillose (4)

3b. Cells of the distal half of the leaf smooth or only slightly mammillose (8)

4a. Papillae or mammillae low and rounded, never projecting saliently as teeth; plants forming dense mats on shaded cliffs; capsules with very short seta, the urn strongly ridged and constricted below the mouth (suggesting Orthotrichum). Amphidium (Orthotrichaceae)

4b. Papillae or mammillae high, sharp-pointed, directed forward and projecting from the leaf margins and back (5)

5a. Leaf margin bistratose (6)

5b. Leaf margin unistratose (7)

6a. Distal leaf cells coarsely and broadly conical-papillose; gametangia in a lateral perigonium, a bud on a short stalk; extremely rare, on granite outcrops, outer foothills. Cnestrum schisti (Dicranaceae)

6b. Distal leaf cells inconspicuously low-papillose; gametangia enclosed in a sessile perigonium, not a stalked bud; on rocks in subalpine forests. Cynodontium (Dicranaceae)

7a. Leaves broadly lanceolate; on wet rocks on cliffs and along snow-melt streamlets; capsule not striate. Dichodontium (Dicranaceae)

7b. Leaves narrowly lanceolate; capsule distinctly striate-ribbed; on vertical faces of granite rocks, subalpine. Cynodontium (Dicranaceae)

8a. Alar and basal cells sharply delimited, thick-walled, orange (9)

8b. Alar and basal cells not sharply delimited or, if so, not thick-walled or orange (10)

9a. Leaves coppery-brown or dark green, leaf tip blunt, rounded (microscopically); costa broad, occupying most of the distal half of the leaf; plants of overflowed rock slopes. Blindia (Seligeriaceae)

9b. Leaves lively green; leaf tip sharp; costa narrow; plants of dry rock surfaces or on peaty soil, at least not directly on flow lines. Kiaeria (Dicranaceae)

10a. Alar cells not or only slightly differentiated; leaf margin recurved, plane, or incurved (11)

10b. Alar cells differentiated (enlarged-quadrate, thin-walled); leaf margins not recurved (17)

11a. Leaves pale bluish-green, always partly inter-tangled with a weft of fine white flexuous organic crystals (resembling "mold" or "cotton candy"); rare plants of the subalpine and alpine slopes. Saelania (Ditrichaceae)

11b. Leaves never intertwined with white wefts (12)

12a. Cells of the distal part of the leaf mostly longer than broad, (13)

12b. Cells of the distal part of the leaf mostly quadrate (14)

13a. Capsule erect, strongly ribbed, not strumose; peristome teeth 16, not forked; seta yellow. Rhabdoweisia

13b. Capsule curved, smooth or ribbed, commonly strumose; peristome teeth forked; seta red. Dicranella

l4a. Capsules short, strongly ribbed, on a short, curved seta and usually hidden among the leaves, mostly not fruiting at all; plants forming very compact, deep golden-brown sods on alpine slopes. Oreas (Dicranaceae)

l4b. Capsule cylindric, on an elongate seta, well exserted above the leaves, commonly fruiting; plants of various habitats (15)

15a. Leaves narrowly lanceolate or linear; capsule erect, cylindrical, not strumose. Ditrichum (Ditrichaceae)

15b. Leaves broadly lanceolate; capsule strumose (16)

16a. Leaves lacking a sheathing base; capsule straight but more or less horizontal, purple-brown, with a few prominent longitudinal grooves; operculum conic; plants of dry, disturbed sites (probably the most abundant weed moss everywhere in our area). Ceratodon (Ditrichaceae)

16b. Leaves with a prominent sheathing base, the lamina abruptly spreading above it; capsule curved and inclined, furrowed when dry and empty; operculum prominently beaked; plants of fens, dripping ledges, or wet streamsides. Oncophorus (Dicranaceae)

17a. Plants very short-stemmed, forming loose tufts on decaying wood and occasionally on rocks in rather dry forested sites; leaves strongly crisped when dry; capsule symmetrical, erect, not strumose. Dicranoweisia (Dicranaceae)

17b. Plants often long-stemmed, forming compact sods on the ground (18)

18a. Costa narrow, thin or thick, in cross-section convex, without thin-walled hyaline cells on the ventral surface; laminal cells homogeneous from the costa to the margin. Dicranum (Dicranaceae)

18b. Costa broad and flat, often occupying the greater part of the leaf width; hyaline cells on one or both surfaces of the costa; laminal cells decreasing in size from the costa out to the margins (19)

19a. Plants small, with leaves less than 3 mm long; hyaline cells present only on the ventral side of the costa; branch tips deciduous, commonly seen lying on the tuft. Campylopus (Dicranaceae)

19b. Plants robust, with leaves commonly over 5 mm long; hyaline cells on both sides of the costa, enclosing central chlorophyllose cells. Paraleucobryum (Dicranaceae)

C. schimperi Milde. A common but easily overlooked species of the subalpine and alpine snow melt areas, occurring on saturated soil around lakes and solifluction terraces (C. subulatus). It is an inconspicuous and nondescript sod-former without any obvious field characters except that, with its very broad costa, it looks like a very small, non-falcate Paraleucobryum enerve. However, since the tips of the shoots break away from the stem to form vegetative propagula, it is very easy to recognize the species in the field because the tufts have yellowish shoot-apices scattered over the surface.

C. schisti (Wahlenberg) I. Hagen. An extremely rare species, known in Colorado from a single collection in Boulder County: west slope of Steamboat Mt., foothills, 6,000 ft. alt., ca. 2 mi NW of Lyons; in crevices of shaded granite ledges, up the main draw west of the old city dump, 21 Feb. 1963, M. Bowers B-10567 (! G. Mogensen). It resembles a very small Cynodontium, but the antheridia are born in a bud on a very short stalk, not sessile. The leaves are falcate when dry, and the distal ends are very coarsely papillose. We have not seen capsules on the Colorado specimen, but they are ribbed. Some authors retain this in Cynodontium. The leaves have papillae rather like those of Dichodontium, but in that genus the margins are unistratose, and the habitat is wet cliffs.

1a. Capsule straight, erect, deeply furrowed, indistinctly or not strumose, the red, erect peristome teeth very evident, the seta twisted; leaves strongly contorted. C. alpestre

1b. Capsule curved, furrowed but not as strongly so, distinctly strumose at the base; leaves only slightly and erect. C. strumiferum

C. alpestre (Wahlenberg) Milde. This is a fairly common species on moist cliffs in the middle altitudes (Rocky Mountain Park westward. The leaves are crisped when dry, the capsule is erect and strongly furrowed, not strumose, on a twisted seta. The red peristome teeth are erect and split down the middle part way. The leaves are very coarsely papillose-mammillose, with sharp points directed forward. This was called variously C. tenellum, C. polycarpum, or C. gracilescens. See Crum and Anderson, p. 183 for a discussion of the complicated nomenclatural history. Collections from Eagle, Gilpin, Grand, Larimer counties.

C. strumiferum (Hedwig) Lindberg. Here the capsule is curved, asymmetric, furrowed, and strongly strumose at the base. In both species the operculum is beaked and diverges from the capsule urn at an angle. Habitat is similar to the last but reported also from sandy river banks, perhaps on drier sites. This was reported earlier as C. polycarpon, which evidently does not occur in America. Collections from Grand and Larimer counties.

D. pellucidum (Hedwig) Schimper. Infrequent. occurring on wet stones in snow-melt streams and cliffs from the foothills to the upper subalpine or alpine. The strongly papillose or mammillose cells and often toothed margins of the leaves of this, Cnestrum, and Cynodontium may lead students astray, since the leaf morphology strongly suggests the Pottiaceae.

The species of Dicranella are very small, forming little tufts with very short stems and narrow leaves. They are usually fruiting. The seta is red in our species, and the capsules very small, oblong, either straight or curved, and the operculum is long-rostrate. Unlike the fairly similar Dicranoweisia, they grow on soil along trails rather than on rock.

1a. Perichaetial leaves or both stem and perichaetial leaves with obovate, sheathing bases and spreading to squarrose leaves (2)

1b. Perichaetial leaves scarcely differentiated from stem leaves; leaves wide-spreading to secund, not squarrose. D. varia

2a. Leaf cells rectangular, 7-15 m wide, 2-3:1; annulus lacking. D. schreberiana

2b. Leaf cells elongate-rectangular, 4-5 m wide, 6-8:1; annulus present (3)

3a. Capsule erect and symmetric; leaves squarrose distally, spreading-flexuous proximally (sterile plants can be distinguished by the attitude of the leaves). D. crispa

3b. Capsule inclined or nodding, the peristome appearing overly large for the urn; leaves erect or falcate-secund. D. subulata

D. crispa (Hedwig) Schimper. When fertile, this species is easily identified by its straight capsule that is slightly furrowed. The differentiated perichaetial leaves are not so easily recognized as they are in D. schreberiana because they are very small, and form a tight "knot" that needs to be dissected out. At first glance they seem to be no more differentiated than those of D. varia. However, upon dissection they are suddenly narrowed from an ovate base. The leaf cells are very definitely very long and slender; this, in combination with the erect and straight, furrowed capsule, make the identification sure. One collection, from Grand Co.: Rocky Mt. National Park, Adam's Falls, Vitt 15318.

D. schreberiana (Hedwig) Crum & Anderson. The very obviously broadened leaf bases and squarrose leaves form a "flower-like" perichaetium. Uncommon; known from a few collections: Larimer Co.: Jinks Creek, Chambers Lake, 8,400 ft., Hermann 28064; Rocky Mt. Nat. Park, on gravelly, peaty creek banks in spruce-lodgepole forests (Hermann 1976, 26596); San Juan Co.: Deadwood Gulch, 2.2 mi S of Silverton, Weber & Wittmann B-111930.

D. subulata (Hedwig) Schimper. Evidently this is our most common species. It is alpine and subalpine. Most of our specimens are fertile; the red seta, curved capsule, and very conspicuous and surprisingly oversized peristome with incurved teeth with gaping spaces between them, are diagnostic. The perichaetial leaves are small and inconspicuous but distinctly broad at the base on dissection, as in D. crispa. The leaves are merely somewhat curved or secund, not spreading in all directions, and the cells are long and narrow. It evidently occurs on soil in open spruce-fir forests. Larimer Co.: Rocky Mt. Nat. Park, trail to Lake of Glass, 10-11,000 ft., Weber & Pontecorvo B-18193.

D. varia (Hedwig) Schimper. This seems to be a trail-side species, not especially restricted to any specialized habitat. The plants are small, but usually heavily fruiting. The leaves are yellow-green, narrow, and merely somewhat wavy rather than straight but not at all curled. The capsule is asymmetrical and not strictly erect, and the operculum is rostrate. It is fairly common mostly in the northern counties (Boulder, Jackson, Grand, Larimer), Archuleta and La Plata, but is not often collected, probably because the plants are small and not conspicuous: Boulder County, from a Salix-Betula thicket beside a peat fen at 10,000 ft. altitude, Vaarama B-11082, !Crundwell; Grand Co: West St. Louis Creek trail, Weber & Dahnke B-91858, and another from La Plata County, along the Animas River (Jamieson 10639).

Although we have records of only one species, it is possible that in the moist forests of northern Colorado there is a second, D. cirrata. The key provided by Flowers (1956) which distinguished them is given below.

1a. Margins strongly and widely recurved in many leaves; lamina smooth, unistratose except for the margins; inner perichaetial leaves with a long, more or less clasping base but scarcely convolute, the upper portion acuminate, shorter than the base. D. cirrata

1b. Margins of the leaves plane, erect to incurved, sometimes narrowly recurved; lamina usually longitudinally striate with narrow cuticular ridges, these appearing in cross-section as minute papillae; lamina unistratose to bistratose in the distal half. D. crispula

D. crispula (Hedwig) Lindberg ex Milde. This is undoubtedly one of the most common mosses in the forested areas, occurring on decaying logs and on boulders. The old capsules persist on the tufts for a long time but lose their peristomes and become quite bleached in color. It is a relatively nondescript moss and for some reason has been often mistaken for Weissia controversa.

1a. Leaves small, usually less than 5 mm long and 0.5 mm wide (2)

1b. Leaves larger, usually 5-10 mm long and 0.5-1.0 mm wide; otherwise not as above (3)

2a. Leaf cells papillose from apex to middle; leaves acute and strongly curled when dry; plants usually on wood or rock. D. montanum

2b. Leaves not at all papillose; leaves spreading widely from a densely tomentose stem, and becoming crisped; slender, stiff branchlets with appressed, short leaves (propagula). In the terminal leaf clusters. On rock. D. flagellare

3a. Plants with most leaf apices broken off; leaves straight when dry; growing on wood. D. tauricum

3b. Plants with most leaf apices present, rarely with a few broken; leaves curled when dry or sometimes straight (4)

4a. Leaves slightly undulate or rugose, especially near the apex; setae aggregate. D. polysetum

4b. Leaves not undulate or rugose; setae solitary (5)

5a. Distal leaf cells usually elongate, sinuose, pitted (6)

5b. Distal leaf cells usually short (quadrate, rectangular, or irregularly angular), neither sinuose nor pitted (or with few pits) (9

6a. Leaves keeled distally, the margins strongly serrate in distal half; costa with 2-4 well-developed dentate ridges on abaxial surface in distal part of leaf. D. scoparium

6b. Leaves tubulose to somewhat keeled distally, margins entire to serrate in distal half; costa without or with poorly developed dentate ridges distally on abaxial surface (7

7a. Alar cells bistratose; leaves with a long, narrow subula, the apex acute to somewhat obtuse; capsule slightly to strongly arcuate. D. spadiceum

7b. Alar cells usually unistratose, rarely bistratose in part; capsule slightly arcuate to straight and erect (8)

8a. Leaves spreading to slightly falcate-secund, the margins serrate near apex; capsule 2-4 mm long. D. rhabdocarpum

8b. Leaves erect-spreading to erect-appressed, the margins entire; capsule 1.5-2 mm long. D. groenlandicum

9a. Leaves narrowly lanceolate, straight and appressed or very slightly falcate; plants of subalpine fens (10)

9b. Leaves broadly lanceolate, convex, widely spreading. Common dry forest plants on soil

(11)

10a. Proximal leaves with acute apices; proximal leaf cells usually less than 40 m long, the median cells pitted mainly proximal to mid-leaf. D. elongatum

10b. Proximal leaves often with blunt apices; proximal leaf cells usually more than 40 m long; cells pitted well distal to the middle of leaf. D. groenlandicum

11a. Leaves relatively unchanged when dry, only lightly crisped, appressed, not standing out at right angles, the stem slightly tomentose but not forming a dense covering, only lightly crisped when dry; common plants of dry forest floors. D. muehlenbeckii

11b. Leaves strongly crisped and narrowly tubulose when dry, the middle stem leaves standing out at right angles to the stem; stem tomentum white in young growth, becoming red-brown below; rare plants of very wet forests, especially streamsides. D. brevifolium

D. brevifolium (Lindberg) Lindberg. Uncommon or rare in the eastern foothills. Collections: Clear Creek Co.: E of Silver Dollar Lake, 11 mi SSW of Georgetown, 29 July 1977 (det. by G. Trucco, 1978, as D. acutifolium); Larimer Co.: spruce-fir woods, 9,800 ft., trail to Comanche Lake, Mummy Range, 10 mi SW of Rustic, 9 June 1977, Hermann 27808 (B-60367, ! Ireland); Twin Cabin Gulch, Wittmann & Weber 112844.

This is very close to D. muehlenbeckii; in fact it was formerly considered to be a variety of that species. In D. muehlenbeckii the leaves are very tightly tubulose all the way to the apex and the costa is not prominent abaxially in the lower half of the leaf. D. brevifolium has leaves that are not as tightly tubulose near the apex because in cross-section they are tong-shaped, and the costa is prominently bulging and shiny on he abaxial surface.

D. elongatum Schleicher ex Schwägrichen. Restricted to alpine and subalpine fens or wet moss tundra. Very easily recognized by its straight leaves and densely packed stems. See also discussion of D. groenlandicum.

D. flagellare Hedwig. A species unlike any other, characterized by its production of microphyllose stems in clusters at the stem apex. It is very small (about 1-2 centimeters), with densely tomentose stems and very narrow, tubulose leaves that spread at right angles to the stem becoming loosely crisped. The cells of the lamina are quadrate and thick-walled. The cells of the broader somewhat sheathing base are rectangular, and the basal and alar cells are large and thin-walled. We have only one station: Larimer C.: Buckhorn Canyon, northwest of Masonville, at mouth of Twin Cabin Gulch, about 100 yards in, 2134 msm (7,000 ft.), N403450 W1052310, 25 May 1981, Rolston 81018 (B-93702), Weber & Wittmann 114328. This is an eastern species, evidently relictual in the Front Range.

D. groenlandicum Bridel. We have one collection: Jackson Co.: Upper Slide Lake, Rainbow Lake Trail, 0.5 mi NE of Mt. Ethel, 10,720 ft., Rolston 82197 (COLO B-93870). Nyholm (p. 69) writes: "The tufts of subsp. groenlandicum are somewhat more robust than [those] of D. elongatum. The leaves are broader below, the nerve narrower. The best character for distinction, however, is afforded by the elongate, strongly incrassate and porose cells in the upper part of the leaf." Hegewald (1972) has beautiful illustrations of the cell detail of this and its close relative D. elongatum. In D. groenlandicum, not only are the stems coarser but the pitting of all the leaf cells is very clearly defined. In D. elongatum the pitting is hard to see but in D. groenlandicum it is very obvious because the cell walls are exceptionally thick.

D. montanum Hedwig. Inconspicuous and infrequent, thus far only found in the eastern Front Range foothills. Only a few collections are known, from a cliff under-hang, growing with Neckera pennata in South St. Vrain Canyon, and in the Boulder Mountain Park, Boulder County. The plants are very small rarely forming intact, dense tufts but usually belonging to the variety pulvinatum Pfeffer. Crum and Anderson describe this modification as follows: "It. . .produces weak branchlets with minute, strongly crisped, spreading, roughish leaves. Such branches are, in fact, scarcely different from the ordinary plants except in size. Whole colonies appear to consist of such leafy branchlets of plants reduced in size and resembling them." Our specimens are little balls intermixed in branches of Neckera.

D. muehlenbeckii Bruch & Schimper. The most abundant Dicranum throughout the forested regions of Colorado. It ranges widely in forests of the middle altitudes. Some specimens have been determined by Lawton as D. fuscescens Turner. The latter species is very common in Eastern United States and in the Pacific Northwest but its presence in Colorado is very doubtful. Ireland (BFNA ms.) calls B. muehlenbeckii a "somewhat rare species throughout North America." See discussion under D. brevifolium.

D. polysetum Swartz. We have only one collection, from depressions in a Sphagnum fuscum-Carex aquatilis fen in the Silver Lake Valley, 3000-3500 m, in the Boulder watershed, Weber B-44158, determined by Wilbur Peterson. Unfortunately the plants are without sporophytes but the vegetative characters are convincing. The leaves are rather stiffly spreading, have a golden sheen, and are strongly wavy-margined.

D. rhabdocarpum Sullivant. An abundant species in the foothill canyons and especially on shaded north slopes. In fruit it is easily known by its erect straight capsules, and sterile by its dense tufts with non-falcate leaves held together by a rich development of reddish tomentum. In winter these tufts fill up solidly with ice!

D. scoparium Hedwig. Common in forests of the foothills, easily recognized when typical by the strongly falcate-secund leaves as if combed in one direction. The Rocky Mountain plants are smaller and less strongly falcate than plants of the East and Far West. Our specimens were verified by Peterson (1979).

D. spadiceum Zetterstedt. We have two collections: Boulder County; Diamond Lake, 11,000 ft. Alt., 11 Sept,. 1974, Hermann 25915, !Ireland; Gilpin Co.: Tolland, July 1914, Grout (DUKE, !Ireland). It is very close to D. muehlenbeckii, but inhabits wet habitats (fens, wet meadows, willow carrs). According to Ireland (in litt.), the very best criteria are the cells of the costa cross-section. In D. muehlenbeckii the cells on the adaxial surface are enlarged, compared to the smaller stereid cells below. Rarely there are some such cells in D. spadiceum but not all the way across the costa as in D. muehlenbeckii.

D. tauricum Sapehin. Infrequent in the outer foothills (Pseudotsuga forests), usually on decaying wood. The straight, yellow-green leaves, many of them broken at the tip, and its habitat, make this an easy plant to recognize. (Orthodicranum strictum).

This is not a difficult genus to recognize in the field, although the technical features that characterize it are primarily microscopic. Its aspect is that of a very small Dicranum, about a centimeter tall. The leaves are about 3 mm long, suddenly narrowed from a broader base, tubular-involute, strongly falcate (curving into a full circle), with the costa excurrent. Instead of having a relatively long shoot with leaves evenly spaced along it, as in Dicranum, there are rather discrete clusters of leaves. The technical features of the leaves include rectangular-quadrate basal cells, of which the lowest tiers are orange. The cross-section of the leaf shows a narrow costa with little differentiation, only a median row of large guide cells, on either side of which are thick-walled cells not organized in stereid bands (unlike Dicranum, which always has stereids). These features require skill in sectioning the leaves by hand or with a microtome.

The capsule is curved, weakly furrowed, with a rostrate operculum. The peristome teeth are wide-spreading when dry, and divided about half way to the base. Crum & Anderson rightly point out that the best way to be sure of this plant is to see that there is a short cluster of leaves (bud) that contains the male antheridia, just below the perichaetium bearing the capsule.

1a. Leaves strongly and uniformly falcate-secund, almost circinate, with extremely long slender apex (the leaves strongly convolute, tubulose, lying on their sides on a microscope slide); distal marginal cells rectangular, smooth. K. starkei

1b. Leaves flexuose but not distinctly nor uniformly falcate-secund, (not tubulose and lying flat on the slide); distal marginal cells irregularly quadrate, mammillose, commonly slightly prorulate. K. blyttii

K. blyttii (Schimper) Brotherus. This species resembles a very small Dicranum muehlenbeckii, but lacks the stereid layer in the costa, and the slightly mammillose-prorulate distal marginal cell are distinctive. We have a single collection, lacking sporophytes, from Jackson Co.: Park Range, along trail to Rainbow Lakes, 9,900 ft. alt., on soil encroaching on surface of boulder at ground level, Weber et al. B-113503.

K. starkei (Weber & Mohr) I. Hagen. Most of our collections were made by Fred Hermann and Holmes Rolston, in peaty soil along subalpine trails and on the borders of small lakes. It is now known from Boulder, Eagle, Jackson, Larimer, and Routt counties. The regularly circinate-falcate leaves are distinctive. It is commonly found with sporophytes.

1a. Leaf limb not sharply differentiated from the sheathing base and not diverging sharply away from it when dry; upper portion of leaf not narrowly subtubulose, not strongly crisped; alar cells somewhat enlarged. O. virens

1b. Leaf base clearly sheathing, strongly clasping the stem, the upper portion abruptly narrowed into a subtubulose limb which diverges sharply 90 degrees away from the sheath; leaf lamina very strongly circinate-crisped; alar cells not enlarged. O. wahlenbergii

O. virens (Hedwig) Bridel. Very common in subalpine springs and fens. Both species have goiters at the base of the capsule, but the leaves of this species are never strongly curled, and the angle at which the lamina comes off the sheath is not very wide. The leaves are about twice as long as in the next. The leaves are revolute in the basal half, the distal costa and leaf margins are bistratose and coarsely toothed. While the alar cells are orange, they are not morphologically differentiated from the basal ones.

O. wahlenbergii Bridel. A very common moss of subalpine and alpine snow-melt basins. Sometimes the two species grow together, but they are very easily distinguished. This species is very variable in the height and density of the tufts. In places that dry out in summer, the growth is limited, the tufts are extremely compact, and the leaves small (forma compactus).

O. martiana (Hoppe & Hornschuch) Bridel. This must be one of the rarest mosses in North America. It is known from several localities in the Colorado Front Range, including Mount Evans. It occurs on the wet shore of Summit Lake at 12,800 ft., and on the steep slopes above the north-facing wall of the lake. Actually it is an abundant moss in Colorado in wet alpine situations, but very rarely occurs in fruiting condition. It is easily recognized in the field by its tight mats of a rich golden color, once one has identified it in the herbarium. Actually, it was Kjeld Holmen (see Steere 1975) who came to Mount Evans after seeing the moss a few days before at Lake Peters and North Greenland, and discovered this in fruiting condition!

la. Leaf margin smooth or with a few teeth at the apex; costa smooth; leaves straight or somewhat curved. P. enerve

la. Leaf-margin serrulate; costa rough on the back (2)

2a. Costa ½ or more the width of the leaf base; margins distinctly denticulate; leaves strongly falcate-circinate; plants usually in loose clumps. P. longifolium

2b. Costa only 1/3 the width of the leaf base; margins with small, scattered teeth or nearly entire; leaves less curved, sometimes almost straight, usually in very tight clumps P. sauteri

P. enerve (Thedenius ex Hartman) Loeske. Usually a plant of moist tundra, inhabiting the most optimum wet tundra sites. We have some records from a north-facing slope in the foothills, so the species does occur in compensating environments at lower altitudes. It characteristically forms solid tufts several inches deep and very tightly compacted. The color varies from glossy pale green to greenish black. We have never found it fertile.

P. longifolium (Hedwig) Loeske. This species is ecologically the reverse of P. enerve, being found characteristically in the foothill canyons in cool, north-facing forested slopes along with such common species as Rhytidium rugosum, Orthodicranum rhabdocarpum, and Timmia. At first glance it would appear to be a Dicranum, possibly D. scoparium, but the Paraleucobryum is more distinctly gray-green and the costa less distinct. The leaves are also more finely attenuate than in Dicranum.

P. sauteri (Bruch & Schimper) Loeske. This occurs in the same areas as P. longifolium. Usually the tufts are very dense and the leaves are not very curved. We have noticed that in Paraleucobryum, when leaves are dissected and placed on a slide, those of P. sauteri almost never lie flat, so that one sees the costa and only one marginal area. The marginal areas are about as wide as the costa. In P. longifolium the very wide costa appears to force the leaf to lie flat, so that the entire costa and the very narrow laminal margins are visible.

Technically, Dicranodontium differs from Paraleucobryum in the cross-sectional view of the costa; Dicranodontium has a costa with guide cells and stereid bands. Furthermore, the leaves are extremely long-setaceous, and the lower leaves are fragile, leaving a bare stem. Paraleucobryum has neither guide cells nor stereid bands, and the lower leaves clothe the stem.

According to Frahm (in litt., 15 July 1986), there is a question as to the taxonomic position He writes: "Is Paraleucobryum sauteri a Paraleucobryum or an Orthodicranum, in the sense of O. rhabdocarpum, fragilifolium, flagellare, and others (not scottianum or fulvum). We made a cluster-analysis and P. sauteri was totally isolated from the remaining species of Paraleucobryum. There are, however, some doubts concerning some morphological characters. Now we will make TL-chromatography; perhaps this will give us a help, because Paraleucobryum has no flavonoids."

Rhabdoweisia is a small Dicranoid moss with crisped leaves with smooth cells, erect, straight, bowl-shaped, 8-ribbed capsules. Unlike Dicranella, the peristome teeth are not separate but joined at the base to form a low basal membrane.

R. crispata (Withering) Lindberg. We have a single collection. This was collected and reported by Jamieson (1986b) from San Juan Co.: Summit of Coal Bank Pass, on NE facing base of vertical gneissic cliff, north end of Spud Mt., head of S Fork Coal Creek, T10N R81W, Sec. 32, 10,500 ft., 22 Oct. 1988, Jamieson 13952.

1a. Leaves in two ranks, broadly sheathing at the base, the rough, subulate lamina consisting mostly of the costa, spreading outward. Distichium

1b. Leaves in spiral ranks, not sheathing at the base (2)

2a. Leaves ovate-lanceolate to narrowly lanceolate, not linear; mature capsule strongly ribbed and inclined; weedy species. Ceratodon

2b, Leaves linear; mature capsule not ribbed; mostly alpine plants. Ditrichum

C. purpureus (Hedwig) Bridel. One of the half dozen abundant weedy mosses in the world and no less so in Colorado. It occurs on packed earth, recently disturbed forest soils, burned areas, sidewalk cracks and neglected ground at all altitudes. It should be one of the first mosses learned by a student. When it is fruiting there is nothing that can be confused with it. The capsules are purple-brown, prominently ribbed, with a conical operculum, and the urn is strumose at the base, where the capsule leaves the seta at a definite angle. In the sterile condition, Ceratodon can be mistaken for almost anything, and usually is. The leaves may be broadly triangular-ovate or quite attenuate depending on conditions of soil, light and moisture. It resembles Didymodon particularly, but the rather uniformly quadrate cells in the distal half of the leaf have a characteristic appearance that will be recognized with experience. Nevertheless, sterile Ceratodon is always troublesome.

On limestone cliffs just below Schofield Pass in Gunnison County, a strange form was found that produced a deep turf of very slender shoots that we thought, without microscopic examination, might be Didymodon vinealis. It grew in crevices along with Mielichhoferia. Richard Zander has determined this to be a form with short-ovate leaves and filamentous gemmae. This was described by Zander et al. (1979) Of our collection, Zander writes: "Note that it is very yellow in KOH and no Didymodon has such a yellow hue. The serrulate upper margins and very square upper laminal cells are distinctive."

la. Capsule erect, oblong-cylindric to ovoid-cylindric; spores 17-20 m diameter. D. capillaceum

lb. Capsule inclined, ovoid and somewhat asymmetric; spores over 25 m diameter. D. inclinatum

D. capillaceum (Hedwig) Bruch & Schimper. The genus is unique in its distichous leaf arrangement and need never be confused with any other except Bryoxiphium, which is also distichous, but extremely rare, on sandstone cliffs in southwestern Colorado. The species of Distichium, however, seem to be impossible to distinguish without fruit, and even with fruit, it takes some experience and intuition to decide whether the capsules are really straight or inclined. There does not seem to be any distinctive ecology to separate the two. These are abundant in wet or moist situations in the subalpine and alpine, occurring in fens, on solifluction terraces, and swampy stream sides. D. capillaceum is the common species, while D. inclinatum has been collected only a few times.

D. inclinatum (Hedwig) Bruch & Schimper. The capsule of D. inclinatum tends to be shorter and plumper than that of D. capillaceum and, in most instances the leaves, which usually diverge widely from the stem in D. capillaceum, are more erect, and their distichous arrangement is not so evident. The spore size is evidently a very reliable character. Spores in most instances range from 30-45 m and seldom down to 20.

17a. Stems 1-4 cm long; leaves up to 3 mm, from an ovate-sheathing base sharply contracted to the subula; costa abaxially strongly convex; laminal cells near costa with weakly nodulose longitudinal walls; commonly fruiting. Ditrichum flexicaule (Ditrichaceae)

17 b. Stems to 7 cm or more; leaves from an elongate-ovate base, tapering gradually to the long, slender subula; costa abaxially weakly convex; basal laminal cells with weakly to strongly nodulose longitudinal walls; rarely found fruiting. Ditrichum gracile (Ditrichaceae)

D. flexicaule (Schwägrichen) Hampe. This plant is infrequent and restricted to low altitudes where it forms very compact mats or sods in seepage areas over rocks and has short leaves that rather abruptly narrow to the upper half of the lamina. It might be mistaken for Didymodon. Moffat Co.: Green River, Sandy Canyon, on shaded sandstone ledges, 6,200 ft., Flowers 8861 (B-66780); Boulder Co.: Boulder Canyon, 1.3 mi W of Four Mile Canyon junction, 6,000 ft., Weber et al. 35501. In Colorado material, the leaves are very short, the sheathing base about the same length as the distal lamina, and not long-attenuate. We find occasional stands that have terete julaceous innovations. Here there is never any difficulty in distinguishing the two species.

D. gracile (Mitten) O. Kuntze. A species of the subalpine and alpine zones; a large, lax plant with very long, filiform leaves in which the spiral twisting is very well marked. It often has a golden color. When mixed with Distichium, one has to look closely to see the difference. In Distichium the leaves are green, two-ranked, with a sheath appressed to the stem and the lamina very suddenly narrowed and spreading, but not spirally twisted.

S. glaucescens (Hedwig) Brotherus. This is an example of a species which is better recognized by a metabolic byproduct than by its morphology. Saelania is a relatively nondescript plant with lanceolate leaves and an oblong-cylindric capsule as in Ditrichum or Distichium. The leaves are always a pale blue-green, the color being accentuated by a blue-gray coating on the leaves which "spins off" as a tangled weft of cobwebby material. This material, however, is not an organism but a chemical substance. Such substances occur in the liverwort, Anthelia, and in Lethocolea. In Anthelia, at least, Dr. Siegfried Huneck has identified the threads as a diterpene compound. This very unusual phenomenon must be investigated in more detail.

January 26, 2005

Ref.: Horton 1983

One genus with vegetative appearance very much like Tortula or Syntrichia and difficult for the beginner to distinguish in the sterile condition, although it may be possible to distinguish Encalypta by its having the end walls of the basal cells thickened or colored. Fortunately, one nearly always encounters it with sporophytes, and in fruit Encalypta is the easiest moss to recognize because of its unique cylindrical-mitrate calyptra with an abruptly narrowed tubular apex (resembling a sausage-balloon partially inflated). Because of the resemblance of the calyptra to some old-fashioned candle-snuffers, the plants are called "extinguisher" mosses.

Encalypta usually occurs in small tufts of a dozen or so fruiting stems in rock crevices or over poorly developed flakes of mineral soil on rock ledges or cliffs. It never forms extensive sods like Syntrichia or Tortula (E. procera may be an exception). The basal cancellinae (hyaline windows) of the leaves are seldom as clearly differentiated as in Syntrichia. E. procera is the only species likely to be found without fruit most of the time, and may be separated from similar species of Syntrichia by its production of multicellular, filamentous reddish-green or brown, papillose gemmae on the stems. The leaves of Encalypta are bright green when moist, and never strongly revolute, as they are in many Syntrichia species.

la. Plants relatively robust, usually sterile and commonly bearing abundant reddish green to brown, filiform gemmae on the stems; capsules spirally furrowed. E. procera

1b. Plants small, usually with capsules, lacking gemmae; capsules smooth or merely straight-ribbed (2)

2a. Leaves gradually narrowed, acute, the apex stoutly apiculate. E. alpina

2b. Leaves oblong, obtuse or very broadly acute (3)

3a. Calyptra fringed, the teeth clearly differentiated as narrowly triangular, often darker units, their bases marked by rows of small quadrate cells. E. ciliata

3b. Calyptra not fringed at the base (4)

4a. Seta very short, the calyptra resting on the uppermost leaves. E. intermedia

4b. Seta longer; calyptra elevated beyond the leaf apices (5)

5a. Capsule distinctly ribbed; peristome present. E. rhaptocarpa

5b. Capsule smooth; peristome absent E. vulgaris

E. alpina J. E. Smith. Horton (1983) says, "E. alpina is differentiated from all other species of Encalypta by the narrowly acute and stoutly apiculate apex of the leaves." The capsule does not appear to have raised ribs, although lines of darker cells are visible under the microscope. It is exclusively alpine. We have two collections, one from Park Co.: Hoosier Pass, 12,500 ft., Weber & Holmen B-4417, and Gunnison Co.: North Italian Peak, 11,000-13,000 ft., 25 July 1955, Weber& Langenheim B-18775.

E. ciliata Hedwig. Frequent in the foothill canyons and up to the tundra, nowhere common. The calyptra has a regular fringe of narrowly triangular segments. The capsule is smooth but fragile.

E. intermedia Juratzka. According to Horton (1983), this species is almost always found in protected overhangs on calcareous outcrops. We have no specimens annotated by her, but she maps it (Fig. 219) as occurring in the western arid tier of counties and evidently some eastern Front Range foothill cuestas. Aside from the short seta, she says, "The opaque calyptra is golden and more or less densely papillose with a short rostrum, and the base of the cylinder is entire. Capsules lack a peristome and are furrowed, with pale orange or orange ribs. Around the mouth of the capsule there is a relatively broad, stiffened band and the cylindric shape of the mouth does not change with age. In almost all the North American populations the leaves are muticous with a rather inconspicuous costa ending well below the apex." A special effort should be made to collect all specimens of the genus in the canyon-lands.

E. procera Bruch. Fairly common but restricted to the San Juan Mountains. Distinguished from all others in having filamentous gemmae in the leaf axils. It is rarely fruiting, but a few collections from the San Juans have sporophytes. The stems are usually elongated compared to the other species. Although it is supposed to be autoicous, our material is almost always sterile. The principal difference between this and E. streptocarpa, often reported, is in the sexuality, but Crum and Anderson maintain that all American collections belong to E. procera. This species differs from all of our other ones in having distinctly elongate stems. The leaves have a long reddish point. The calyptra tapers gradually rather than suddenly to the apex. The basal cells of the "window" have orange walls, especially the thickened end-walls, and the bases of the leaves have inter-tangled slender rhizoids associated with them.

E. rhaptocarpa Schwägrichen. Montane and subalpine, possibly largely replacing E. vulgaris on limestone at high altitudes. Summit Co.: Blue Lake Dam, Monte Cristo Creek valley, on limestone, Weber & Wittmann B-111125. We find it often very difficult to determine whether the capsules of the respective "species" are really ribbed or not. Horton also is equivocal about whether these are species or two subspecies which show tendencies toward each other; the question is still open. E. vulgaris is the older name, and perhaps E. rhaptocarpa is best regarded as a subspecies.

E. vulgaris Hedwig. Common in rock crevices in the foothills canyons and going over to bare ground in the tundra. Mature capsules may be somewhat wrinkled, but do not have strong, reddish ribs. And, while the calyptra may be somewhat ragged at the base, it is not really ciliate. Horton says that the true E. vulgaris in America is restricted to California and Arizona. Colorado populations are either true E. rhaptocarpa or intermediates.

June 19, 2003

A rather heterogeneous group of genera sharing the following characters: leaf cells elongate-linear, costa short, double or lacking; alar cells often well-developed. The technical characteristics, exclusive of the sporophyte, suggest a close relationship with Pleurozium!

1a. Stems not complanate, julaceous; alar cells forming a conspicuous, bulging bistratose group at the more or less constricted leaf base; plants of alpine tundra. E. concinnus

1b. Stems complanate; alar cells unistratose, forming a clear area of quadrate-rectangular cells; plants of outer foothills. E. cladorrhizans

E. cladorrhizans (Hedwig) Müller Hal. We have one collection, from Boulder Co.: Boulder Mountain Parks; Bear Canyon Trail, 2 Dec. 2001, Weber, Wittmann, & Mazurek B-112798. This is an eastern woodland species with extensions in northwest America. The closest station in the western United States apparently is in Arkansas.

The similarity of the vegetative parts with those of Pleurozium is striking although the species are easily distinguished. In Entodon cladorrhizans the color of the plant is a bronze yellow; the stem is not red, and has scattered stiff tufts of rhizoids; the alar cells are in several rows (about 8 across and ten or more high), quadrate and little thickened. The stem leaves are closely imbricate and do not expose the stem. In Pleurozium the plants are green, lack rhizoids, and the leaves are rather transparent. The alar cells are few (about 3 across and high), with very thick, yellow walls. The stem leaves are more distantly attached and spread out, revealing the red stem.

E. concinnus (De Notaris) Paris. An alpine species, very rare in western North America. Usually occurring mixed with other mosses in optimum moist, mossy tundra sites. The plant, always sterile, is recognized by its yellow-green color and rather julaceous stems with rounded leaf-apices. The green alar cells stand out conspicuously because of their bulging, densely bistratose tissue. For discussion of the American distribution of these two species, see Steere (1975). I have collected this on Mount Wilhelm, in Papua New Guinea!

F. ciliaris Bridel. Fabronia is the tiniest and most delicate of our pleurocarpous mosses. The creeping stems, leaves and all, are only about 0.3 mm wide when dry (to about 0.5 mm when wet). The plants can be recognized very easily under the microscope, however, because of the large teeth which project outward from the margins of the leaves. These teeth are commonly hyaline, contrasting with the green cells of the leaf lamina. The leaf-tip is also hyaline, abruptly long-attenuate from the ovate blade, and frequently forked at the apex. This very unusual serrate or even fimbriate condition is unique among our mosses. Because of the ciliate margins and the long, slender, often colorless leaf-tips, the branches often appear fuzzy under the hand-lens. Fabronia almost always grows in the innermost recesses of rock crevices of cliffs in the foothill canyons. Our collections are all from siliceous rocks. The tufts are loosely attached to the substrate, sometimes almost lying free, and they range in color from bright green to pale yellowish-green. The capsule is goblet-shaped on a well-exserted seta; the peristome is like that of Orthotrichum.

March 14, 2005

Ref: Pursell ined., in BFNA

The leaves of the Fissidentaceae are in two ranks, lying in a single plane and thus giving the shoot a frond-like appearance. The leaves are unique among our mosses. The lower half of the leaf has a large, single lamella extending from the costa toward the stem, forming in effect a second leaf-blade lying over and partly covering the inside half of the lamina. This smaller "half-leaf" is called the dorsal lamina. The "sheath" formed by this structure is called the "vaginant lamina." The sporophytes are minute, with a well-developed seta and a cylindrical urn that is constricted below the rim; the operculum is rostrate. The species of Fissidens, our only genus, are limited to wet ground, dripping cliffs, and shelving streambanks and road-cuts of wet clay. A few species even live submerged in pools and streams.

la. Completely submerged in pools or pot-holes in sandstone stream-channels; plants slender, with long, narrow, rather distant leaves. F. fontanus

1b. Terrestrial but often in very wet places; leaves not as above, usually touching each other and forming a flat frond-like shoot (2)

2a. Large plants several centimeters long; leaves ribbon-shaped, multistratose, the vaginant lamina similarly shaped, running along the and not easily seen except along the margin. A plant of waterfalls. F. grandifrons

2b. Minute plants less than a centimeter long; leaves not as above (3)

3a. Leaves not bordered by narrower elongated cells. F. osmundoides

3b. Leaves wholly or in part bordered by a band of narrow elongated and colorless cells (4)

4a. Leaf apex rounded or obtuse, not apiculate, F. obtusifolius

4b. Leaf apex ending in a short apiculus (5)

5a. Leaves crispate when dry. F. crispus

5b. Leaves flat or nearly so when dry (6)

6a. Leaves lanceolate to ligulate, acute; apiculus poorly developed. F. bryoides

6b. Leaves ovate to lanceolate, rounded to obtuse-apiculate; apiculus a single sharply pointed cell. F. sublimbatus

F. bryoides Hedwig. This species has stems with closely overlapping leaves, flat when dry, the limbidium is very thin and often disappears near the leaf apex, the laminal cells are up to 16 microns, and the their walls are are distinctly squared (angular). Capsules are usually present. In the other species, capsules are rarely produced. Frequent, but very inconspicuous, on moist banks from the foothills through the subalpine. This species is not substrate-restricted, occurring on soil over seeping granite rocks. Our collections are from Boulder, Gilpin, Grand, Jackson, Larimer, and Mesa counties.

F. crispus Montagne. In this species, the leaves are longer and more slender, less closely overlapping than in F. bryoides, and curl downward rather than lie flat when dry, the cells are small (6-10 m), rounded, and deep, and the limbidium is strong and broad, developed on all laminae. Pursell described the habitat as follows: Moist, shaded soil, over rocks and streams, infrequently submerged by fluctuating water levels. Collections: Boulder Co.: Boulder Canyon, Sayre 212a (B-191887), Weber B-80111; Jackson Co.: Big Creek Falls, 2.5 mi SW of Big Creek Lakes, 2800 m., Rolston 81120 (COLO B-93577).

F. fontanus (La Pylaie) Steudel. A very distinctive species, forming frondose tufts in pools of intermittent streams. The leaves are long and narrow, up to 3-4 mm, and except for the distichous arrangement, might be thought to resemble a small Fontinalis rather than a Fissidens. In fact it is made a special genus called by some Octodiceras julianus. We have no Colorado record, but F. fontanus has been collected in the Canadian River Canyon in Mora County, N.M., and should be expected in the drainage of the Purgatoire or Apishapa in similar situations in south-eastern Colorado.

F. grandifrons Bridel. This extraordinary species is common in the canyon country of Utah. The fronds are quite long and pendent on seeping canyon walls, and the leaves are multistratose. We have one collection that occurs within easy walking distance of the Colorado state line: Uintah Co.: Jones Hole Creek Campground, Yampa River, in the direct spray of a waterfall, Lehr 2036 (B-113306). The species must occur in Colorado, but most likely will be discovered by a boating party.

F. obtusifolius Wilson. The species is easily distinguished by the obtuse or rounded leaf apices. Pursell describes the habitat as follows: "Typically on limestone and limestone-bearing sandstone along streams and waterfalls, often near the high water level, sometimes inundated." Collections: Mesa Co.: Unaweap Canyon between Whitewater and Gateway, Weber et al. B-43555; Montezuma Co.: Beaver Creek, 10 mi NNW of Dolores, 7,500 ft., Pursell 3247.

F. osmundoides Hedwig. Easily recognized by the lack of a leaf border and the crenulate margin caused by the protruding corners of the marginal cells. F. osmundoides is not uncommon but is found only on peaty banks in the alpine and subalpine. Our records are from Boulder, Clear Creek, Grand, Jackson, Lake, and Summit counties.

F. sublimbatus Grout. Pursell describes the habitat of this as follows: "Soil in arid areas where plants are often partially buried, often under overhanging rocks and boulders and in the shade of trees and shrubs." Collections: Boulder Co.: Hall Ranch Open Space, Wittmann & Weber (B-114729), Steamboat Mt., N of Lyons, Wittmann & Lehr (B-114761).

July 8, 2003

The Fontinalaceae are large mosses attached to rocks in mountain streams. The leaves are tristichous (no other mosses in Colorado are tristichous) and the branching is irregularly pinnate. The sporophytes are borne on short lateral shoots and the capsule is almost hidden by the closely enveloping perichaetial leaves. The peristome of the Fontinalaceae is also unique, consisting of an outer row of 16 teeth and an inner row of 16 cilia variously united to form a cone-like trellis.

la. Leaves without a costa, dark green to blackish, plane or keeled, not falcate-secund. Fontinalis

lb. Leaves with a costa, always keeled and folded, reddish- or golden-brown, strongly falcate-secund. Dichelyma

Dichelyma is a large, semi-aquatic moss of the size of Fontinalis, but the strongly falcate leaves suggest Scorpidium; the leaves, however, are keeled, folded, and three-ranked.

1a. Leaves strongly falcate, folded lengthwise (conduplicate), the costa merely percurrent; the stems somewhat wiry; in swiftly flowing water. D. falcatum

1b. Leaves straight or very slightly falcate, not folded, the costa long-excurrent; the stems flaccid; in quiet water of fens. D. uncinatum

D. falcatum (Hedwig) Myrin. Attached to stones in cold, rocky streamlets on slopes in the subalpine spruce and aspen forests. Our collections are from Rocky Mountain National Park and in the Park Range of Jackson Co.

D. uncinatum Mitten. Evidently restricted to quiet water of fens. Our single collection is from the San Juan Mountains, San Juan Co.: Chattanooga Fen, just off Colorado Hwy. 550 near south base of Red Mountain Pass, near historic town of Chattanooga, Cooper B-110449.

la. Leaves (examine the median cauline leaves) usually keeled or keeled and folded along the keel (2)

lb. Leaves usually plane. F. hypnoides

2a. Ends of leafy stems and branches conspicuously elongated, triangular-pyramidal in shape, with closely imbricate leaves, the branch clearly triangular in gross form; leaf keel straight or slightly curved, leaf apex acute, entire; perichaetial leaves apiculate. F. neomexicana

2b. Ends of leafy stems and branches not as above; keels moderately to strongly curved (from base to apex), leaf apex blunt to broadly obtuse; perichaetial leaves obtuse. F. antipyretica

F. antipyretica Hedwig. Common in the subalpine zone, but also occurring at lower altitudes. In my experience this seems to prefer slow-moving water, while F. neomexicana prefers swift water. Field study is needed to determine what precise ecological distinctions there may be in the respective habitats.

F. hypnoides Hartman. This species prefers slow-moving water of ditches and swamps. F. duriaei and F. hypnoides are so-called "critical species," and may in fact not be distinct from each other. If one chooses to consider the complex a single species, then the older name which should be used is F. hypnoides. In her recent monograph, Winona Welch (1960) wrote: "Plants of Fontinalis hypnoides show great variation in vegetative and fruiting structures. On some plants in the same collection or in different collections, all leaves are typical of F. hypnoides. Some branch leaves on plants of F. duriaei resemble median cauline leaves of F. hypnoides. It is very important that median cauline blades of well-developed or mature plants be used for accurate determination. Occasionally it is difficult to name the species with certainty. However, plants which are distinctly F. hypnoides and those which are F. duriaei without question have leaves which are definitely different and give cause for retaining the two species." In Colorado both species have been collected in the same localities by the same collectors on the same day, suggesting that here at least what we are dealing with is variability of a single taxon. There seems to be no evidence of separate ecologies for the two, and not enough field observation has been made in the light of Dr. Welch's notes. The F. hypnoides phenotype is the more infrequent of the two. In Europe two species may very well be involved, since there F. duriaei has a more southerly distribution than F. hypnoides, being absent from Scandinavia but going into North Africa and Iraq, while F. hypnoides occurs widely in Scandinavia and reaches Italy but goes no farther south. Both species occur widely throughout the rest of Europe.

F. neomexicana Sullivant & Lesquereux. Common in swift-moving water of streams in the foothills and subalpine. This is the easiest of the group to recognize, since the strongly triquetrous and dense terminal shoots are usually quite evident especially in the fresh condition. This species has an unusual disjunct distribution, mostly Rocky Mountain but occurring on the Upper Peninsula of Michigan!

August 3, 2004

Ref: Fife (1970), Smith (1994)

la. Capsule elongate, asymmetric and curved, the mouth oblique instead of perpendicular to the axis. Funaria

1b. Capsule straight, goblet-shaped, erect, smooth, with the mouth perpendicular to the axis (2)

2a. Capsules urceolate and short-necked. Physcomitrium

2b. Capsules globose-pyriform and long-necked Entosthodon

E. sonorae (Cardot) Steere. A minute, evidently rare, moss on eroding sandstone cliffs. The leaf rosette is only a few mm high, the leaf cells large and hyaline, the costa ending very near the leaf apex. When dry, the leaves are contorted. The leaf cells are hyaline and irregularly hexagonal to rectangular; the cells of the margin are distinctly longer and narrower. The seta is very slender and elongate. The capsule is oblong, with a long neck, and the peristome is absent. The distal leaf margin is coarsely serrate by the blunt ends of protruding marginal cells, or serrulate by the bulging short tips of the cells. This species is otherwise known from Arizona and Mexico. The southwestern corner of Colorado is evidently the farthest north it comes in North America. Our only collection is from Montezuma Co.: on soil beneath a large overhanging boulder, mouth of Yellowjacket Canyon at the junction of Yellowjacket and McElmo creeks, ca. 30 mi W of Cortez, E-facing slope, 4,800 ft., 10 June 1958, Pursell 3224 (as E. tucsonii). The specimen is mixed with Weissia ligulifolia.

1a. Capsules sulcate or ribbed when dry, the mouth very obliquely oriented; revoluble annulus present (2)

1b. Capsules smooth or slightly wrinkled when dry, the mouth less strongly oblique, sometimes almost transverse; annulus absent; minute plants less than 1 cm tall, infrequent in desert-steppe areas. F. americana

2a. Endostome very short and truncate; plants minute; rare on wet tundra. F. microstoma

2b. Endostome of lanceolate segments, at least half as long as the exostome; common large (over 1.5 cm tall) weedy plants especially in burned areas, at any altitude; also abundant in greenhouse pots. F. hygrometrica

F. americana Lindberg ex Sullivant. We have two certain records: Mesa Co.: on arid benches above East Creek N of Gibbler Gulch, in deep recess, on calcareous soil at base of overhanging sandstone rock, 1 June 1973, Weber, Kunkel & LaFarge B-43493. and from Delta Co.: Unaweap Canyon. An old report of "Funaria hybernica Hooker" (Porter, 1876) may have referred to this species but the specimen has not been found. Recently, the Delta County specimen was annotated by D. K. Smith as F. muhlenbergii, but that species is European only.

This species, according to Crundwell & Nyholm (1974) has been called F. muhlenbergii, a European species which probably does not occur in America. Most or all specimens so named are misidentified. The authors say: The nomenclature of this moss is a confused one. The name Funaria muhlenbergii was apparently used by Hedwig fil. in manuscript, and applied to a moss collected by Muhlenberg in Pennsylvania. Early European authors considered that European material belonged to the same species." Our species is treated as Entosthodon americanus by Fife, J. Hattori Bot. Lab. 58:192. 1985, but this disposition evidently needs more study.

F. hygrometrica Hedwig. A very abundant moss of sandy soil along streams and in burned areas in forests, where the burned wood and depressions where trees once grew provide seasonally moist sites. The offset operculum is unmistakable. Hygrometrica evidently refers to the seta, which was seen to twist and untwist with changes in relative humidity. It is almost always to be found in flower pots in greenhouses and nurseries.

F. microstoma Bruch & Schimper. A very small plant, rarely occurring on the tundra. Fife cited a specimen collected by Schantz from the vicinity of Pikes Peak, Schantz 47 (FH), evidently collected on the first International Phytogeographic Excursion in 1914. We have it also from Summit Co.: Blue Lake, Weber, Wittmann & Spribille B-11112. Better specimens are needed.

This genus is very easy to recognize by its small size, broad, translucent leave with large Bryum-like cells, and its small, erect, goblet-shaped capsules on a short or longer seta. The operculum is flat when attached, and has a distinct erect rostrum. The plants are evidently annual, and is so small and so often mixed with common species (weedy Bryum, for example) that they are only collected by mistake.

1a. Annulus present, large, revoluble, of two or three rows of bloated cells. P. hookeri

1b. Annulus remaining attached to the capsule mouth, consisting of a row or two of small, brown cells. P. pyriforme

P. hookeri Hampe. We have no modern collections. P. coloradense E. G. Britton (1894) was described from Colorado based on a type collection of Brandegee, April 13, 1877, probably from the Arkansas River basin. Crum & Anderson (1955) saw a possible fragmentary type specimen of this on a Grout slide at DUKE. They concluded this taxon is synonymous, and that "seta length is of no great importance in P. hookeri, where the seta ranges from 2 to 4 mm in length, so that the capsules may be either emergent or exserted." This is evidently a moss that is most common on the high plains and should hardly be expected to occur in high mountain wetlands. We have no records of this since Brandegee's day, but the plains of Colorado have not been explored bryologically.

P. pyriforme (Hedwig) Hampe. The only relatively common species, P. pyriforme has been found only a few times, once in a chain of beaver ponds in Routt County, and once in a sandy flood plain near Boulder. It frequently occurs in greenhouse pots, so is likely to escape into adjacent wet areas almost anywhere. In this species the annulus is small, not revoluble, consisting of a single row of thick-walled, orange cells, remaining attached to the capsule; the capsule mouth is bordered with about 6-8 rows of horizontally elongate cells. Collections are from Boulder Co.: Rocky Flats, floodplain of South Boulder Creek, 7 mi S of Boulder, June, 1967, Weber B-39084, June, 1973, Kunkel & Shultz B-46180.; Lake Co.: Twin Lakes, 1873, Wolf & Rothrock (as P. latifolium = hookeri, but the annulus is not large and revoluble). Routt Co.: Diamond Park Road, 28 mi N of Steamboat Springs, 18 July 1951, Weber B-23608, Hinman Park, 8,000 ft. alt., on road bank, 5 July 1971, Flock B-96023.

P. immersum Sullivant T. P. James (1878), reporting on the collections of Wolf and Rothrock at Twin Lakes, listed Aphanorrhegma serratum (obviously a misidentification of a Physcomitrium, possibly P. immersum Sullivant). This is a plant of the Great Plains and is unlikely to occur in the high mountains. The Twin Lakes lie in the upper dry, although high, valley of the Arkansas River. The locality is now destitute of mosses, but more than one Physcomitrium might once have occurred on the gently sloping sandy shore, which is now greatly changed by drainage, development, and seasonal recreation. Further serious search for specimens of Physcomitrium needs to be made along the floodplains of eastern Colorado.

Pyramidula tetragona (Bridel) Bridel was reported by Porter (1876) from "moist, sandy soil on the plains, Hall." The specimen, in the T. P. James Herbarium at Harvard is from "moist sandy soil on the Platte, E. Hall." This probably came from the Scotts Bluff area of Nebraska, but the eastern tier of Colorado counties has never been explored for mosses and some of the ephemeral spring-growing species are very likely to be found here. It is a common weedy moss in the eastern and midwestern United States.

October 13, 2004

Ref.: Muñoz (1983-1999), Greven (2003). Ireland (2004)

la. Stem with short lateral branchlets arising along its length and creating the impression of a main stem with horizontal side-branchlets; basal and usually median leaf cells with sinuose-nodulose walls; never fruiting in our region; plants of moist or wet depressions in the subalpine or alpine. Racomitrium

1b. Stems simple or branched more or less dichotomously, the branches all ascending together; leaves without elongate sinuose-nodulose cells or with this character not strongly developed; most species found fruiting here; plants of various sites (2)

2a. Leaves distinctly biplicate distally, in adaxial view the costa appearing to be depressed on each side under the adjacent lamina; hair points always very well developed. Coscinodon

2b. Leaves not biplicate; hair points various. See the following key to Grimmia, Schistidium, and Jaffueliobryum

With a little practice, the various genera presently segregated out of the old Grimmia may be recognized in the field. Coscinodon forms neat hoary tufts with long hair-points. It is easily recognized when it fruits in early autumn, by the large mitrate calyptra. Jaffueliobryum grows in very small tufts in crevices of sandstone cliffs or on bare vertical faces of bad-land "fingers" in the desert-steppe. It has broad, thin, light green leaves. The Grimmia anodon group with its two species have immersed capsules and enlarged perichaetial leaves; they form low, short-stemmed dense tufts with variable hair-pointed leaves, and usually grow on sandstone or limestone in the desert-steppe. Schistidium grows on various rocks, sometimes in or near shallowly flowing rills. The plants are nearly always fruiting, and have immersed capsules with a red operculum. In size they range from a few millimeters high tufts to some several centimeters long.

1a. Leaves concave, obtuse, oblong-lanceolate, lacking hair-points; lamina unistratose throughout, margins incurved; costa narrow, ending below apex; leaf cells rather uniform, quadrate to short-rectangular with smooth thin walls throughout; rare, forming dull green tufts in icy water from melting snow-banks in the alpine region. G. mollis

1b. Leaves not as above, if broad then acute and/or with hair-points, or with revolute margins, and in different habitats (2)

2a. Leaves muticous, with gemmae on their distal portions. G. anomala

2b. Leaves not muticous, or if so, lacking gemmae (3)

3a. Leaves, even when dry, widely spreading and contorted, or spirally twisted around the stem; hair-points very short or absent (4)

3b. Leaves never wide-spreading or crisped or spirally twisted around the stem when dry; with or without hair-points (7)

4a. Leaves, when dry, spirally twisted around the stem, the tips often coiled into a circle, erect-spreading when wet; upper leaves sometimes with a short hair-point; gemmae often present on the back of the costa of the upper leaves; rare, forming continuous sods on cliff faces, almost always associated with Amphidium. G. torquata

4b. Leaves crisped and contorted when dry, linear-lanceolate. (5)

5a. Leaves ca. 2 mm long, elongate-lanceolate, tapering and gracefully cygneous when wet, yellowish, the older ones brown; hair points well-developed. G. trichophylla

5b. Leaves up to 1.5 mm long, abruptly narrowed , generally blackish; hair-point very short or lacking (6)

6a. Median leaf cells rectangular-quadrate, the walls with nodular thickening but not strongly sinuose (accordion-pleated); in small tufts on protected overhangs of alpine fell field boulders; rare. G. incurva

6b. Median cells rectangular quadrate, conspicuously sinuose-incrassate (accordion-pleated). Very rare, in similar situations. G. elongata

7a. Stems very slender, less than 0.5 mm wide, about 2 cm long, almost unbranched; leaves 1 mm long or less, the upper ones with a spinose hair-point; always sterile here, on vertical cliffs wet with seepage water (8)

6b. Stems stouter, usually more or less branched, with longer leaves (9)

8a. Hair-point coarse and finely denticulate, sometimes without a hair-point. G. teretinervis

8b. Hair point weak, often flexuose, very short and strongly spinulose. Schistidium tenerum

9a. Leaves plane or concave, dark green or blackish, bistratose, margins not recurved, apex with a long, flat hair point; costa lying in the plane of the lamina, hardly visible. (10\

9b. Leaves not as above in all respects (11)

10a. Leaves ovate-triangular; basal marginal cells oblate (transversely elongated), hair points stout, strongly denticulate and decurrent. plants forming very low tufts which separate into individual stems upon handling, the leaves immediately spreading widely upon wetting; on sandstone ledges in southern Colorado. Grimmia laevigata

10b. Leaves oblong-ovate, the basal marginal cells quadrate-rectangular with thickened end walls; hair points only slightly decurrent. G. ovalis. Common on sunny granite outcrops in the foothills of the Front Range, and on rhyolite boulders, subalpine forest flood-plains, San Juan Mts.

11a. Robust; stems up to 10 cm long, in sods with the stems usually trending downslope; leaves 3-5 mm long, linear-lanceolate, elongate, revolute; lower leaf cells rectangular, with thickened sinuose walls; capsules uncommon, on short side branches, strongly longitudinally ribbed; abundant on massive granite outcrops, foothill canyons to alpine. G. elatior

11b. Stems shorter (usually up to 2.5 cm); leaves shorter; basal cells usually not sinuose-walled; usually fertile (except G. pilifera). . . . (12)

12a. Seta shorter than the capsule, which is therefore immersed (13)

12b. Seta equaling or longer than the capsule, which is exserted (17)

13a. Leaves 2 mm long; hair point often very long (see discussion under G. elatior). G. pilifera

13b. Leaves about 1mm long; hair points short or nearly lacking (14)

14a. Columella falling with the operculum; leaves revolute on one edge, bistratose distally; plants brownish or black; hair-point lacking or short; tufts small, often loose; usually on granitic rocks (15)

14b. Columella not deciduous; leaves plane or in-rolled, unistratose; plants in low, dense tufts or sods, very short, and often filled with soil or sand, the upper leaves at least with a distinct hair point (except sometimes in Grimmia plagiopodia); on sedimentary rocks at low altitudes. See Schistidium

15a. Calyptra mitrate, covering most of the capsule; capsule symmetrical, on a short, straight seta; hair-point long; leaves very broadly ovate or obovate, erose-denticulate above the middle, with long hair-point; plants green; perichaetial leaves not conspicuously larger than the others; crevices of sandstone outcrops or bare vertical surfaces of "badland" formations. See Jaffueliobryum

15b. Calyptra mitrulate, covering only the upper end of the capsule, irregularly split; leaves not erose-denticulate; hair-points developed on upper leaves, but often short; plants usually brownish or black; perichaetial leaves conspicuously larger and broader than the others; on horizontal surfaces of sedimentary outcrops (16)

16a. Leaves keeled distally, unistratose; peristome well-developed, cribrose; hair-points poorly developed. Grimmia plagiopodia

16b. Leaves concave distally; bistratose on the margins and toward the leaf apex; peristome absent or rudimentary; hair-points strongly developed on perichaetial leaves; peristome absent; plants autoicous. G. anodon

17a. Tufts low, very dense, blackish, the leaves with plane or incurved margins, usually less than 2 mm long (18)

17b. Tufts taller, loose, green or brownish-green; leaves usually over 2 mm long, with the margins usually revolute along one edge (21)

18a. Leaves short, broad, broadly keeled, curved toward the stem when dry; seta cygneous or simply curved. G. bicolor

18b. Leaves elongate, narrow, not keeled or curved toward the stem when dry; seta straight. (19)

19a. Basal half and margin of the leaf with elongate rectangular clear cells, not thickened transversely; costa strongly raised abaxially. G. donniana

19a. Basal and marginal cells otherwise (20)

20a. Leaves short and stiff, not at all flexuose when moist. G. montana

20b. Leaves not short and stiff, usually flexuose when moist; costa weak proximally, channeled distally and raised I-beam-like on the abaxial surface (21)

21a. basal marginal leaf cells quadrate; leaf margins always plane; capsules smooth, brown, with thick-walled exothecial cells. G. alpestris

21b. Basal marginal leaf cells rectangular; one leaf margin sometimes slightly recurved; capsules striolate, yellowish, with thin-walled exothecial cells. G. sessitana

22a. Leaf lamina distally unistratose; seta curved downward; leaves broad, abruptly narrowed to the broad-based hair-point; green apex of leaves distinctly rounded or even emarginate; hair points long, making the tufts hoary-looking; calyptra mitrate or mitrulate. G. pulvinata

22b. Leaf lamina bistratose; seta straight; Leaves narrow, the green apex acute and grading into the narrow-based hair-point; hair-points evident but relatively short and inconspicuous; leaf lamina bistratose; calyptra cucullate (23)

23a. Leaf-margins plane or incurved; leaves channeled above, thickish and waxy-looking; costa broad, not terete; leaves not keeled; plants dioicous. G. ovalis

23b. One or both leaf margins revolute; costa terete; leaf keeled; plants autoicous. G. longirostris

Grimmia and its related genera form one of the larger groups of mosses in our flora and the species have been notoriously difficult to identify. However, they do sort out quite well into rather distinct ecological situations, and the following list of sites, with their respective species, may help in instances where the key may be equivocal.

1. Wet gravels of snow-melt streams (plants not anchored firmly to rock): Grimmia mollis, Schistidium.

2. Vertical siliceous cliff faces: Grimmia elatior, G. arizonae, G. piliferum, Schistidium tenerum, S. apocarpum group.

3. Sloping siliceous rock faces, irrigated periodically: G. elatior, Schistidium occidentale, S. alpicola, S. agassizii.

4. Siliceous boulders, not permanently wet by seepage: Coscinodon calyptratus, C. cribrosus Grimmia alpestris, G. bicolor, G. donniana, G. longirostris, G. anomala, G. elongata, G. incurva, G. montana, G. ovalis, G. pulvinata, G. trichophylla, Schistidium apocarpum group.

5. Crevices of horizontally layered sedimentary rocks: Jaffueliobryum wrightii.

6. Surfaces of sedimentary rock formations: Jaffueliobryum raui, Coscinodon cribrosus (vertical), Grimmia anodon, G. laevigata, G. plagiopodia, G. pulvinata, G. sessitana, G. teretinervis (also in granite crevices).

With experience, one may learn to recognize many of the species without capsules, but there are a few that evidently cannot be separated. Capsules, fortunately, will be found eventually, unless either one or the other sex simply does not occur here.

A calyptra, when present, is mitrate in Coscinodon, Jaffueliobryum, G. anodon, G. plagiopodia, G. poecilostoma, and G. pulvinata. In all other Grimmiaceae the calyptra is cucullate.

Grimmia almost always found with many capsules: G. alpestris, G. bicolor, G. donniana, G. pulvinata, G. longirostris, G. sessitana.

Grimmia very rarely found with capsules: G. elatior, G. laevigata, G. ovalis, G. mollis, G. montana, G. pilifera.

Grimmia never found with capsules: G. anomala, G. teretinervis, G. incurva, G. torquata, and Schistidium tenerum.

Coscinodon varies: capsules are present only in certain seasons, but when present are quite abundant.

There are three types of opercula: mammillate or conical, rostellate, and rostrate. Rostellate stands intermediate between mammillate and rostrate. Recognizing these when they are present may be helpful in eliminating some species from consideration:

Mammillate or conical: alpestris, anodon, plagiopodia, sessitana

Rostellate: mollis, montana

Rostrate: anomala, elata, incurva, laevigata, ovalis, longirostris (variable), pulvinata, torquata

Greven (2003) recognizes four growth forms. Our species fall into the following:

Cushions (stems tightly clustered and rounded): alpestris, anodon, bicolor, incurva, laevigata, montana, orbicularis, pulvinata, sessitana; Coscinodon spp.

Tufts (stems caespitose, dense or loose): laevigata, longirostris, ovalis, sessitana, teretinervis, torquata.

Patches (rather loose and irregularly formed): anomala, elatior, mollis

Mats (extended flat colonies): alpestris, montana, plagiopodia

Ref: Hastings (1999).

1a. Leaves 1.1-1.9 mm long, bistratose distally; margins involute above; capsules immersed. C. cribrosus

1b. Stem leaves 1.4-2.4 mm long, unistratose or patchily bistratose above; margins plane or revolute on one side; capsules exserted. C. calyptratus

C. calyptratus (Hooker ex Drummond) C. Jensen in Kindberg. Very abundant and characteristic of granitic boulders in the outer foothills in the ponderosa pine stands, but less commonly occurring up to the highest elevations, on both sides of the Continental Divide. Fruiting abundantly in early autumn and easily recognized in the field by the hoary aspect, exserted capsules, and large mitrate calyptra. The species also occurs in central Asia (Muñoz 1998).

When sterile, one must be aware that there are Grimmia species with hoary tufts and long hair points. The leaves of C. calyptratus are plane, with thickened, bistratose but nor recurved margins. The distal lamina is partially bistratose, while the rest of the lamina is unistratose with rounded-quadrate cells, and quite translucent. The basal juxtacostal cells are elongate-rectangular, those more toward the margin short-rectangular, and the few marginal rows elongate-rectangular and hyaline. The leaf cells are not thickened or sinuose-walled.

C. cribrosus (Hedwig) Spruce. Hastings says: "Its biplicate, bistratose leaves with strongly incurved margins and immersed capsules with distinctly perforated peristome teeth separate C. cribrosus from other species in the genus." This is a distinctly smaller plant than G. calyptratus. The species is evidently minerotrophic, restricted to a particular rock formation associated with mining areas. Collections: Chaffee, Conejos, Gunnison, Hinsdale, Ouray, Summit, and San Juan counties. A Flowers specimen (3593) from San Juan Co., Utah was collected as Grimmia alpestris.

Sterile tufts of Coscinodon , especially C. cribrosus, will be mistaken from Grimmia species, such as G. alpestris and G. sessitana. They have a shaggy appearance because the hair points are often as long as the leaves; the leaves do not spread out arcuately when wet, as they do in the common G. longirostris, and the leaf margins are not recurved. The leaf anatomy is critical. In Coscinodon the hair point is commonly very long, slender, flexuous, and practically without teeth. The distal leaf lamina is "crimped" (narrow and biplicate) and its leaf cells are small and irregularly rounded-quadrate and often crimped in the middle (sinuose). The median cells are quadrate-rectangular, thin-walled, and the basal cells are rectangular, the basal marginal ones more elongate-rectangular. The costa is commonly distally grooved.

Ref.: Greven (2003)

The subgenera of Grimmia: Although these groups are based largely on the sporophyte, they do hang together fairly well vegetatively and it might be useful to have this analysis.

Gasterogrimmia includes G. anodon, G. plagiopodia, and G. poecilostoma. The group is uniquely characterized by the sporophyte, in which the seta is very short and asymmetrically attached to the urn. Unfortunately, capsules are only infrequently present.

Litoneuron includes G. laevigata and G. ovalis. Here the seta is straight to sinuose, the leaf is concave and the costa is not raised above the level of the laminal cells. The seta is straight to sinuose and the lamina is -3-stratose

Guembelia includes G. alpestris, G. donniana, G. elongata, G. longirostris, G. montana, G. pilifera, G. sessitana, and G. teretinervis. Here the seta is straight to sinuose, the leaf is keeled and the costa prominent.

Rhabdogrimmia includes G. anomala, G. elatior, G. incurva, G. pulvinata, and G. trichophylla. Here the seta us usually arcuate or cygneous, the leaves are unistratose distally. Probably G. elatior really belongs in Guembelia.

G. mollis stands alone in having a costa not reaching the leaf apex, having broad concave, rounded leaves, and occurring almost totally submerged in running melt-water of show patches at bvery high altitudes.

G. alpestris (Weber & Mohr) Schleicher. This somewhat resembles G. montana, but is always distinctly glaucous-green, with rather long hair-points. It also forms a softer tuft, and is never the deep black color that one associates with G. montana. It closest relative is G. sessitana, which differs in its rectangular basal marginal cells. Collections: Gunnison, Montrose, San Juan, and Summit counties. In Montrose Co. it occurs in Paradox Valley, between Naturita and Bedrock, on a gypsum-salt dome in center of the valley in an area vegetated chiefly by lichens and small black mosses, 5,230 ft. alt., !H. Greven. Otherwise it can occur at very high altitudes in the montains. Some collections are quite sterile and have extremely small leaves, almost too difficult to dissect; these are evidently male plants.

G. anodon Bruch & Schimper. G. anodon and G. plagiopodia are dark green or brownish rather than green mosses forming very low, tightly attached tufts on sedimentary and calcareous rocks at low altitudes. The only good technical distinguishing feature for them are the absence and presence of a peristome, respectively. The immersed capsule and the enlarged perichaetial leaves serve to mark the group. Plants low, the stems decumbent, "brushed" to one side, the perichaetial very broadly oblong; copper- colored distally, green below, hair point broad at the base, flexuose and serrulate; distal laminal cells rounded-quadrate; margins nearly plane, bistratose; basal marginal cells quadrate, the inner rectangular (2:1), not sinuose. Capsule immersed, globose, operculum conical, immature

calyptra mitrate, attenuate. A short seta is present, but it is attached to one side of the capsule base, and is bent sideways and upward.

Schistidium flaccidum may be mistaken for G. anodon. See discussion under that species.

Greven states (in litt.) that sterile plants can be distinguished as follows: