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Historical references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.