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Colorado Flora: Eastern Slope, Fourth Edition A Field Guide to the Vascular Plants
Colorado Flora: Eastern Slope, Fourth Edition A Field Guide to the Vascular Plants
Colorado Flora: Eastern Slope, Fourth Edition A Field Guide to the Vascular Plants
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Colorado Flora: Eastern Slope, Fourth Edition A Field Guide to the Vascular Plants

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Colorado Flora: Eastern Slope describes the remarkable flora of the state, distinctive in its altitudinal range, numerous microhabitats, and ancient and rare plants. Together with Colorado Flora: Western Slope, Fourth Edition, these volumes are designed to educate local amateurs and professionals in the recognition of vascular plant species and encourage informed stewardship of our biological heritage.

These thoroughly revised and updated editions reflect current taxonomic knowledge. The authors describe botanical features of this unparalleled biohistorical region and its mountain ranges, basins, and plains and discuss plant geography, giving detailed notes on habitat, ecology, and range. The keys recount interesting anecdotes and introductions for each plant family. The book is rounded out with historical background of botanical work in the state, suggested readings, glossary, index to scientific and common names, references, and hundreds of illustrations. The books also contain a new contribution from Donald R. Farrar and Steve J. Popovich on moonworts. The fourth editions of Colorado Flora: Eastern Slope and Colorado Flora: Western Slope are ideal for both student and scientist and essential for readers interested in Colorado's plant life.

LanguageEnglish
Release dateApr 6, 2012
ISBN9781607321415
Colorado Flora: Eastern Slope, Fourth Edition A Field Guide to the Vascular Plants

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    Colorado Flora - William A. Weber

    Colorado’s Major River Systems

    Some of Colorado’s Major Topographic Features

    Colorado

    Flora

    EASTERN SLOPE

    Bill Weber holding Ptilagrostis cf. porteri and Saussurea sp., Yabagan Pass.

    Altai, 1978. Photo by Ivan Krasnoborov, W. A. Weber collection

    Colorado Flora

    EASTERN SLOPE

    FOURTH EDITION

    A Field Guide to the Vascular Plants

    William A. Weber

    Fellow of the Linnean Society of London,

    Professor Emeritus,

    University of Colorado Museum of Natural History

    Ronald C. Wittmann

    Associate, University of Colorado Museum of Natural History

    with the assistance of

    Linna Weber Müller-Wille

    UNIVERSITY PRESS OF COLORADO

    Boulder

    © 1990, 1996 by William A. Weber

    © 2001, 2012 by William A. Weber and Ronald C. Wittmann

    Published by University Press of Colorado

    5589 Arapahoe Avenue, Suite 206C

    Boulder, Colorado 80303

    Fourth edition

    All rights reserved

    Printed in the United States of America

    The University Press of Colorado is a cooperative publishing enterprise supported, in part, by Adams State College, Colorado State University, Fort Lewis College, Metropolitan State College of Denver, Regis University, University of Colorado, University of Northern Colorado, and Western State College of Colorado.

    This paper meets the requirements of the ANSI/NISO Z39.48-1992 (Permanence of Paper).

    Library of Congress Cataloging-in-Publication Data

    Weber, William A. (William Alfred), 1918–

      Colorado flora : eastern slope / William A. Weber, Ronald C. Wittmann. — 4th ed.

           p. cm.

      Includes bibliographical references and index.

      ISBN 978-1-60732-140-8 (pbk. : alk. paper) — ISBN 978-1-60732-141-5 (ebook) 1. Plants—

    Colorado—Identification. I. Wittmann, Ronald C. II. Title.

      QK150.W38 2011

      581.9788—dc23

                                                               2011021430

    Design by Daniel Pratt

    21   20   19   18   17   16   15   14   13   12            10   9   8   7   6   5   4   3   2   1

    For their true understanding and unwavering support

    during decades of work in making the Colorado Flora

    I dedicate this edition

    to my late wife Sammie (1917–1996),

    my daughters Linna, Heather, Erica, and their families,

    and to all my friends, mentors, colleagues and students,

    who have shared learning and teaching with me.

    Bill Weber

    For my wife Judy and son Matthew,

    and the many dear friends

    who have patiently waited for me as

    I wandered off the well trodden path contemplating weeds.

    You have encouraged and supported our botanical enterprises

    and shared your love of the natural beauty of Colorado.

    Ron Wittmann

    CONTENTS

    Preface to the Fourth Edition

    Acknowledgments

    Background of Floristic Work in Colorado

    Books to Inspire

    A Vade Mecum for the Field Botanist

    Key to the Families

    Ferns and Fern Allies

    Gymnosperms

    Angiosperms

    Figures

    References

    Index

    Glossary

    Illustrated Plant Structures

    PREFACE TO THE FOURTH EDITION

    The field guides of the Colorado Flora began in 1949 and continue to be works in progress. We learn more and more every season and I, myself, feel very lucky to have lived long enough to have made a good start. Ours is a most interesting flora, unique in its remarkable altitudinal range, special climatic events, and numerous microhabitats. Although having written extensively on vascular plants, lichens, and bryophytes, we have only scratched the surface. Twenty years ago I (Bill) rashly pronounced that I had seen nearly all of the flora in the field and that there was not much left to do. With the wisdom of age and experience, I now am convinced that the flora is so vast and complex that our work is still in the exploratory stage, even after our combined efforts of more than 65 years (Weber) and 30 years (Wittmann).

    Currently, taxonomy is in a state of upheaval, largely due to evidence introduced by the comparative analysis of DNA. Some of this molecular evidence supports the traditional classification, but much of it seems to conflict with morphological ideas that have ruled taxonomy since the time of Linnaeus. Although we feel that it is premature to adopt the new taxonomy, lock, stock, and barrel, we often mention at least the proposed changes. In any case, professional and amateur field botanists require a practical classification scheme that is based primarily on morphology and ecology, and that de-emphasizes characters that cannot be easily observed. We hope that local floras and field guides such as this one will serve their needs.

    If nothing else, the present, often heated, taxonomic debates illustrate that botany is a vibrant, dynamic science. Plant names will continue to change to reflect our best understanding of phylogeny. There is no end in sight. According to Dominguez and Wheeler (1997), Taxonomic stability is ignorance. Amateurs, who are often confused by all of the new names, should keep in mind that in our science the names we use are concise statements of our opinions. To paraphrase Gilbert & Sullivan’s Nightmare Song (Iolanthe), We conceive you may use any names that you choose to indulge in without impropriety. In this edition we have provided a number of references in support of our positions.

    While we were preparing this fourth edition of the Colorado Flora, a massive project has been going on to produce a complete encyclopedia of the vascular plants and bryophytes of the United States and Canada. The Flora of North America North of Mexico (abbreviated FNA in our text) is well under way, but is only half completed. We have attempted to bring the Colorado Flora into harmony, following the FNA in some instances and noting differences of opinion in others.

    The FNA promises a wealth of information (in 30 volumes), including exhaustive descriptions for all of our taxa, but it is in no way a replacement for local floras. There are several disadvantages: (1) Because of the larger number of species, FNA keys tend to be complex and technical. (2) There are many errors (omissions and inclusions) regarding the occurrence of plants in Colorado. A lack of documentation makes it nearly impossible to settle distribution issues cleanly. (3) Distributional information is too general. (4) Habitat data are slighted. (5) Synonymy is scanty, making it difficult to relate the FNA to other floral treatments of Colorado. (6) Authors often have no direct knowledge of the Colorado flora. (7) The FNA is expensive and certainly cannot be carted into the field.

    Our field guide offers several advantages: (1) In a local flora the keys can be simpler. Taxa tend to be better defined (less problematic) locally. (2) Occurrences are supported by documented records that are available in herbaria and online databases. (3) Distribution of uncommon taxa is frequently given by county. (4) Habitat is a major point of emphasis. (5) We account for names used in other Colorado treatments, including the FNA. A published Catalog (Weber & Wittmann, 1992) provides even more complete synonymy. (6) Local authorities have more intimate knowledge of the flora in the herbarium and in the field. (7) Local floras are relatively inexpensive and they can be carried into the field.

    Readers will find invaluable the points of view offered by other local and regional floras. The University of Colorado Museum’s website is a particularly useful resource. Here will be found the specimen database of Colorado vascular plants, various county checklists, Catalog of the Colorado Flora (electronic), and links to other herbaria and botanical sites. When citing specimens we use standardized herbarium abbreviations: COLO (University of Colorado), CS (Colorado State), RM (Rocky Mountain). Other abbreviations may be decoded at the Index Herbariorum website: http://sweetgum.nybg.org/ih/.

    We believe that those who use field and local floras should become serious about their hobby and try to really learn about their subject, including the history of classification, the rules of nomenclature, and the fundamentals of growth and reproduction. The ultimate purpose of our Flora is to educate local amateurs and professionals in the recognition of plant species so that they can be better stewards of our priceless and irreplaceable biological heritage.

    W. A. WEBER (BILL)

    R. C. WITTMANN (RON)

    February 2012

    ACKNOWLEDGMENTS

    Our exploration of the Colorado flora has been a cooperative enterprise, involving both field and herbarium work and spanning more than half a century. So many of our students, foreign visitors, and specialists have given invaluable aid that it is impossible to name them all. We have learned from each other, and it is difficult to sort out the mentors and the mentored. We are indebted to those who have alerted us to new specimen records and who have reported errors in earlier versions of our books and papers. Thank you all.

    David Cooper, wetland ecologist of Colorado, the western USA, and the Peruvian alpine, has opened our eyes to the riches of calcareous and iron fens that are yielding new records. That was a neglected phase of Colorado’s flora, and his work alerted us to the rich bryophyte flora as well as vascular plants.

    Those specialists who have contributed their keys or whose excerpted ones we have used are acknowledged in the text. Special thanks are extended to George Argus for his help on Salix, Rich Scully for Potentilla and Lamiaceae, Steve O’Kane for his keys to Physaria, and the moonwort team of Don Farrar and Steve Popovich for providing the latest treatment of Botrychium. We also thank Tim Hogan and Nan Lederer, who for the past ten years have been indispensable managers of the COLO herbarium. Their development of the specimen database of the COLO collections and county checklists is much appreciated.

    A special kudos to my junior author, Ron Wittmann, whose career has been in Boulder as a physicist with the National Institute for Standards and Technology. In the 1970s Ron came to the herbarium as an amateur botanist and gardener with a few plants that needed identification. He had taken a course in plant taxonomy under C. Leo Hitchcock at the University of Washington. Since that beginning, Ron has developed a unique skill in recognizing the Colorado flora in all of its aspects—vascular plants, lichens, and bryophytes. In my declining years he has become my arms and legs, daring to scale steep slopes, scorning cold-weather clothing, casting aside his hand lens and using his twenty-power eyes to discover rare bryophytes. For the present edition Ron has been the computer guru. Bill Weber has done the leg work in the herbarium and handled much of the correspondence. He was not allowed to touch the keys to use two fingers to type the manuscript. Our joint effort has included some crucial discussions, Ron approaching problems with the precision of a professional physicist against Bill’s more romantic approach and taxonomic experience. We never came to blows, and our partnership has been truly fruitful.

    In the final stages of preparing the manuscript my daughter Linna Weber Müller-Wille, my son-in-law Ludger Müller-Wille, and my grandson Ragnar Müller-Wille offered editorial help and made constructive comments on the introductory sections of the flora.

    W. A. WEBER (BILL)

    R. C. WITTMANN (RON)

    February 2012

    BACKGROUND OF FLORISTIC WORK IN COLORADO

    William A. Weber

    Herbaria

    Our knowledge of floras have their beginnings with the collecting of botanical specimens. Fortunately the habit of establishing herbaria, collections of dried plants, began long ago, perhaps in Italy. Reports not backed up by specimens in an herbarium are useless hearsay. These collections must be guarded from abuse, carelessness, and destruction by wars, for the very basis of our knowledge of plants rests on these. The actual specimen upon which a plant name is based is called a type specimen. Linnaeus’ type specimens are deep underground in a bombproof vault in London. At the very end of hostilities in World War II the specimens of monocots that were carefully saved in caves by the curators of the Vienna Museum herbarium were discovered by American soldiers. It was a day for celebration, so those priceless specimens, thousands of them, were burned. Perhaps the greatest tragedy was the destruction of the Berlin herbarium during a bombing raid in March 1943. Accidents will happen, but as scientists we are bound to try to preserve what we can.

    The local herbarium should be available to amateurs, government workers, and conservationists as a source of reliable information. Serious students should be encouraged to use the facility and volunteer their services.

    Documentary collections should be considered vital archival materials, and must not be allowed to deteriorate even if their use decreases as emphasis on other disciplines increases. Detailed information should be maintained for local floras, and the label data should be put online so that it is practicable to trace state records to specimens. The University of Colorado Herbarium maintains a database providing the complete label data and a checklist for each of the 67 Colorado counties. The latter is an indispensable tool for the fieldworker, and, if space is at a premium, county lists will prevent cluttering the herbarium collections with superfluous common specimens. Many herbaria are available online, but as yet there is no central database for all herbaria.

    The University of Colorado Herbarium (COLO)

    It was many years before there was a large enough herbarium in Colorado to be considered important. In the early 1900s Colorado State College and the Colorado Historical Society each had a small herbarium. The University of Colorado Herbarium was established in 1946. It now contains over 300,000 vascular plant specimens, 112,000 lichens, 118,000 bryophytes, and smaller collections of algae, fungi, and slime molds, and is well known internationally.

    When I came to Boulder, I immediately made my first visit to the University of Colorado Museum of Natural History. When the director, Dr. Hugo Rodeck, asked me what I wanted to do at Boulder, I replied, I would like to build an herbarium. His answer was Do we need one? Herbaria in those days rarely found a home in a museum but belonged to a teaching department of botany. Colorado had no botany department. At the time, the attic of the museum building did contain an herbarium, but it was the personal property of Joseph Ewan, who had left Colorado during World War II to join other botanists seeking new sources of quinine in South America. Because Ewan didn’t receive the PhD, he was not invited back to Boulder, but moved to Louisiana, where he became the century’s most renowned historian of American botany. He shipped his herbarium back to Tulane University.

    The Colorado Historical Society donated their small herbarium of collections made by Alice Eastwood in the 1880s. She had been a teacher of classics at a Denver high school. Those precious specimens formed the tiny nucleus of what would become the University of Colorado Herbarium in 1946.

    It was obvious that several things had to be done. For teaching purposes we needed to have an herbarium housed in the Museum and to ensure continuous curatorship and growth. Francis Ramaley, Professor of Biology at Boulder from 1898 to 1942, had collected ecological specimens; many of them were poor in quality, the collection was limited in scope and contained many duplicates. These were temporarily housed in the Museum by the Biology Department, but did not constitute an herbarium.

    The Museum had no tenure-track faculty. The staff consisted of a director, the departmental secretary, and the preparator. Anyone else who worked in the museum did so on a volunteer basis. I was an instructor in the Biology Department and became a faculty member of the museum only in 1962 when the administration of CU President Quigg Newton collapsed and its provost Oswald Tippo moved me into the museum as a full professor. This freed me to teach what I wanted and to have time for botanical travel and research.

    There was hardly any money to do all of the things necessary to build an herbarium. There was mounting, classifying, and label writing. Luckily I had learned to type with just my index fingers while typing briefs in a patent law firm in New York. Single labels were easy, but how to create masses of duplicate labels for exchange specimens? I started with a manual typewriter, then on to an electric one with balls of type, later picking up discarded machines from Norlin Library—tape machines such as the Flexowriter and Edityper. When we got into lichens and wanted to send out exchange sets of sixty duplicates, we needed something better, so we bought a small printing press, complete with a tray of type, and set the labels by hand.

    A strong program of field collecting in Boulder County, followed by intense collecting throughout the state, was needed to build the collections. Many reports of the species in the state flora had to be verified. These were mostly to be found among the collections of the early transcontinental expeditions, housed in the few great herbaria of the eastern US: the New York Botanical Garden, Harvard University, the Philadelphia Academy of Science, Iowa State College (C. C. Parry), the Missouri Botanical Garden, and a few major herbaria of the West Coast (Pomona, UC Berkeley, Stanford, and UCLA). The herbaria at Colorado College and Colorado State University were relatively small, although they were of great importance. For the University of Colorado herbarium I had to create a new and useful one that would aim at completeness. It would be necessary to borrow critical specimens to establish the validity of the identifications.

    So collect I did. Naturally I began with the Boulder area, but soon began a project that would ensure the coverage of the whole state. I began with the corner counties: Baca, Montezuma, Moffat, and Sedgwick. Most of my fieldwork for the first few years was done in Boulder County and vicinity, but I took my students to the Western Slope for a long weekend every spring and began to fill the herbarium with much new material. During the summer I taught field botany at Science Lodge, now the University Mountain Research Station, and the Rocky Mountain Biological Laboratory at Gothic, north of Gunnison, and began serious collecting of the alpine flora. I also began to fill in the gaps in the collections at Mesa Verde National Park and established an herbarium at Dinosaur National Monument and Colorado National Monument. My students were all-important; we learned together and many of them have gone into professional teaching and research positions. My connection with them has never ebbed.

    Early Floristic Efforts

    Before Colorado began to be explored, Colorado plant species were being collected and described, but not from specimens collected in Colorado. Some were collected in colonial America, some by Lewis and Clark in Wyoming, some in Mexico by Sessé and Mociño, some in Europe, some in the Altai mountains of Siberia. The type specimens are scattered in great herbaria in different parts of the world. Most early collections of Colorado species are in the herbaria of Harvard University, the New York Botanical Garden, the Philadelphia Academy of Science, the Missouri Botanical Garden, and the Royal Botanical Gardens at Kew.

    Here are a few of those botanists who paved the way to the publication of Colorado Flora.

    Edwin James (1797–1861)

    The first important plant collections in Colorado, about 700 species, were made by Edwin James on Major Stephen H. Long’s Expedition, 1820, which trekked along the South Platte River, down along the base of the Front Range, to the base of Mesa de Maya near the present village of Branson, thence on to New Mexico, Oklahoma, and Texas. James collected the first alpine species in Colorado on a hike up Pikes Peak. Aquilegia coerulea, the state flower of Colorado, was collected on this historic expedition. George Goodman, my major professor at Iowa State, and his colleague Cheryl Lawson, retraced James’ steps and brought together all aspects of the Long Expedition with critical evaluations of the collections and their histories (Goodman & Lawson 1995). Except for its lack of keys to the species, this book could serve very well as the first flora of Colorado, since it deals with the plants, their habitats, the campsites where they were collected, and the taxonomic changes that the names have undergone over almost two centuries.

    Thomas C. Porter (1822–1901) and John M. Coulter (1851–1928)

    A Synopsis of the Flora of Colorado, by Thomas C. Porter and John M. Coulter, 1874, was an outgrowth of the Hayden Surveys. It was the first of a planned series of publications aimed at introducing the flora to students and scientists who were beginning to discover Colorado: a detailed catalog of the flowering plants, with descriptions but no keys, and short lists of the bryophytes (by Leo Lesquereux), lichens (Henry Willey), and fungi (Charles H. Peck).

    Alice Eastwood (1859–1952)

    Alice Eastwood, born in Canada, came to Denver in 1873 and graduated from East High School. After graduation, she taught at the same school for about ten years. She never had any further schooling but earned enough to retire from teaching and devote her life to botanical studies. In 1889, upon hearing that T.D.A. Cockerell was living in Westcliffe, she visited him there and botanized with him. Cockerell (then 23 years old) left Colorado for England in 1889 after designating Miss Eastwood (then 29) secretary of his Colorado Biological Association. A letter from May 30 reads: In case I do not again communicate with you before your departure I wish you a most pleasant journey and great happiness and prosperity on the new journey which you begin at the end of this. I have learned much from you, in some respects more than from anyone else. I do not hope to be able to return the obligation to you but perhaps I can, to my fellow man (Weber 2004, p. 548). Alice Eastwood went on to California to become curator of the herbarium of the California Academy of Science in San Francisco, and eventually became America’s most famous woman botanist. She became a heroine of the science when she thoughtfully segregated the type specimens and carried them out of the collapsing building during the great earthquake and fire of 1906.

    The only flora available to Eastwood was Coulter’s Manual of the Botany of the Rocky Mountain Region (1885). She felt that she should try to meet the needs of the local population, so she published A Popular Flora of Denver, Colorado (1893). She evidently was well enough known in Denver that when Alfred Russel Wallace came to Colorado he sought her out and they made a botanical excursion to Gray’s Peak. A small number of her collections came to the University of Colorado Museum when the Colorado Historical Society broke up its herbarium in the 1930s.

    Per Axel Rydberg (1860–1931)

    Per Axel Rydberg’s Flora of Colorado (1906) is of more than historical interest. Collections on which the book was based were made by Professors James Cassidy (1881–1889) and C. S. Crandall and a student, J. H. Cowen, who became professor following Crandall’s retirement. Cowen died before he assumed his position. Since the college, now Colorado State University at Fort Collins, was unable to carry out further studies of the plants, an arrangement was reached with the New York Botanical Garden to have Dr. Rydberg, who had field experience in Utah, Montana, Nebraska, and Wyoming, develop the eventually published flora.

    Rydberg did fieldwork in Colorado only once, in July and August of 1900, with F. K. Vreeland, to the Sierra Blanca (Blanca Peak). Some sites visited were Turkey Creek and Indian Creek Pass, in Huerfano County, where they collected five to six thousand specimens, now at the New York Botanical Garden. Aside from the small herbarium that had accumulated at Fort Collins, Rydberg relied on studying T. C. Porter and J. M. Coulter’s Synopsis of the Flora of Colorado; J. M. Coulter’s Manual of the Botany of the Rocky Mountains; T. S. Brandegee’s account of the Flora of Southwestern Colorado; John Torrey’s report on the Edwin James collections made on the Long Expedition; Asa Gray’s reports on the collections of C. C. Parry, Elihu Hall, and J. C. Harbour; E. L. Greene’s various publications in Pittonia, Plantae Bakerianae, Leaflets of Western Botany, and the publications of Aven Nelson, M. E. Jones, George E. Osterhout, and Alice Eastwood; and the journals Zoë, Erythea, and the Proceedings of the California Academy of Science.

    Alice Eastwood and T.D.A. Cockerell were the only resident botanists in Colorado in Rydberg’s time. Most of the species known then had first been discovered in other states by transcontinental expeditions, such as Lewis and Clark, and the Hayden Surveys. There was little information about the habitats of Colorado plants. The most Rydberg could do in his flora was to make keys to the genera and the species, and list the known localities from the briefest statements on the specimen labels. The only field information Rydberg could use was from his own experience in neighboring states or territories.

    A few of the striking things about his book may be mentioned here. The Flora of Colorado was the first place in which Rydberg attempted to make sense of the enormous genus Astragalus and to break this terribly unwieldy mess into, easily, eighteen smaller genera. These have never been accepted, but it was a sincere and justified attempt. He divided Gentiana (a genus that is, in the strict sense, exclusively Eurasian) into four genera. He divided Aster into twenty groups, Senecio into 17 groups (these still wait to be accorded generic status), and Cirsium (as Carduus) into nine. These remain the most difficult taxa in the Colorado flora.

    Rydberg’s reputation of being a great splitter has been exaggerated. It is true that his lack of field experience led him to name many herbarium sheet species, but in the long run, scientists often find merit in his observations. As my friend Áskell Löve said to me, It is better to split than to lump, because the lumper tends to lose valuable information. We should remember that in Rydberg’s time, genetics was an infantile science, polyploidy and apomixis were unknown, and of course a hundred years of advances in our knowledge lay ahead.

    T.D.A. Cockerell (1866–1948)

    From age twenty-one Theodore Dru Alison Cockerell spent three years, 1887–1889, in Westcliffe, Wet Mountain Valley, because of a mild case of tuberculosis. He never had any postgraduate training, but he was already an accomplished naturalist in England, specializing in molluscs. Aside from making a bare living from doing chores, in Colorado he began to collect the entire biota of the Sangre de Cristo area, sending specimens and information to specialists all over the world, answering their questions about Colorado’s flora and fauna, which were virtually unknown abroad. He encouraged his neighbor, Mrs. M. E. Cusack, to collect plants. Her collections eventually ended up in several herbaria of England, including the Royal Botanic Garden at Kew. Cockerell returned to England, where he did research that helped Alfred Russel Wallace in his revision of Island Life. After a decade or so in Jamaica and New Mexico, he returned to Colorado and in 1904 came to the University at Boulder, where he spent the rest of his life. He became the world authority on wild bees and wrote a monograph on the plant genus Hymenoxys and a book on the zoology of Colorado. In 1906 he began serious excavations of the Florissant Fossil Beds, which eventually became a national monument. Cockerell was certainly the most famous resident naturalist of Colorado and did much to introduce Darwinian theory to the United States. He was also a humanitarian and a life-long socialist. For more detailed biographical information on Cockerell, see Weber 2000, 2004.

    Leon Kelso (1907–1982)

    The story of Leon Kelso is that of a potentially great opportunity missed. Kelso dearly wished to have a chance to write a flora of Colorado. He came to Colorado from Kansas to attend the University, attended the University of Denver and then the University of Colorado, where he planned to do graduate work in plant taxonomy. He was intensely interested in willows, Carex, and grasses, and knew a lot about the exciting geographic distributions of the alpine flora. He was told by the department head, Who are you to work on Rocky Mountain plants, when Dr. Rydberg and Dr. Greene worked on them for 50 years? You, putting your ideas beside theirs; it is absurd. . . . [I] would have no majoring in systematics here. Kelso wrote, Taxonomic work in western botany has been obstructed by the closed shop policy that has existed in systematic natural history for the past thirty years. He left Colorado, broke all of his ties with our institutions, and began to publish his own journal, a flora of Colorado called Biological Leaflets. He went to the U.S. Biological Survey in Washington DC, where he became a specialist in identification of the stomach contents of small game birds and animals, earned a master’s degree at Cornell, and published many papers on the owls of the world. He taught himself Russian and wrote reviews of Russian ornithological literature, finally becoming interested in electrostatic and bioelectronic phenomena concerning birds. In my mind, there is no doubt that if he had been encouraged, he had the potential to write a flora of Colorado.

    Leon Kelso’s educational experience was a classic example of the damage that can be done by established scientists who, for whatever reason, decide that a young person desiring help and counsel is unworthy or incapable of doing significant work. I wrote an account of Leon Kelso’s life and his list of publications for the Colorado Native Plant Society’s Biographies of Forgotten Botanists series (Weber, n.d., unpubl. ms.; publication is available on request). Many of his caustic comments on the state of the science in his time have significance to us today. I think it is important for Colorado botanists to know who this man was, because, like Rydberg and Greene, he was snubbed by the establishment, but many of his observations show that he was an astute observer and had great potential for Colorado botany that unfortunately was never realized.

    Harold D. Harrington (1903–1981)

    Up until the mid-1940s there was no flora of Colorado that was either complete or that contained keys and detailed descriptions. The only volume in use when I arrived in Colorado in 1946 was Coulter & Nelson’s Flora of the Rocky Mountains, even then a rare and out-of-print book. It covered a number of the states and had both keys and short descriptions. Had it been available, I would have used it in my field courses.

    Dr. Harrington was brought to Colorado in 1943 from the University of Iowa for the express purpose of writing a standard flora of Colorado. There was, of course, the historical link to Rydberg’s effort that was in some way sponsored by the Experiment Station of Colorado State University. I believe this project was his major responsibility.

    Harold was a large person, very quiet and even-tempered, not terribly outgoing, but someone one could be comfortable with. His face reminded me of Babe Ruth. He worked in a large room in a small building that evidently was once a gymnasium, situated on the main thoroughfare. The herbarium was not very large, and bundles of specimens, mounted and awaiting mounting, were always lying on a number of tables. I don’t think he worked with a typewriter, but put down all of his notes in longhand. The University of Colorado has his original handwritten manuscript, which dealt with one species each on 8½ × 11–inch manila cards. On one side he wrote the descriptive information, which seems to have been used unaltered in the final manuscript. On the reverse side of each card there is a county map of Colorado.

    The cards, especially the data on the maps, are a valuable archive. Harold did not intend to map the complete distribution of common species on his cards. Rather, he placed a spot on the map marking where a specimen of a poorly known or rare species had been seen, and used a series of abbreviations that indicated in which herbarium he had seen that specimen. This was a remarkably useful device, which proved crucial in several instances in our current revision.

    Harrington didn’t have much time for fieldwork, and I believe that his health prevented him, at least in his last years, to do much collecting in the alpine. I accompanied him on a few days’ trip to Baca County, and was told that he had botanized in Browns Park, Moffat County, with C. L. Porter, curator of the Rocky Mountain Herbarium at Laramie. Porter told me about a trick they pulled on Harold on that trip. It was evidently customary on these trips to shoot some small game for meat; Harold did not participate. As they were cooking the meal they chatted about their favorite small game. Prairie dog was mentioned. Harold demurred at the idea of eating such a rat-like animal, but Porter told me they made sure Harold not only ate, but actually raved about the anonymous piece of prairie dog they put on his plate of beans.

    Whenever Harold had accumulated enough questions to necessitate a visit to Boulder or Colorado Springs, he would spend a few days with me or with Bill Penland. These trips were important because, while the herbaria we had were still very small, we were actively collecting and adding more species to the state list.

    One of the main problems in writing a flora was that the state of Colorado did not have its own herbarium resources. Most of the critical specimens had been collected on the historical expeditions or by private collectors, like Greene, Jones, and Baker, from California. The historical collections were at Gray Herbarium or the Philadelphia Academy of Science. It is doubtful that Harrington had sufficient funds to borrow many specimens. His book contains many statements like Reported from Colorado but I have seen no specimens.

    Harrington’s Manual of the Plants of Colorado was published in 1954 by the small Sage Press of Denver. The massive book was printed from a typewritten copy. Nevertheless, it was the very first and is still the only flora of Colorado that contains keys and adequate descriptions for the vascular plants of our state. My original copy has stood the test of time and, although the pages are dog-eared and the binding is loose, it must be referred to constantly. Harrington deserved much more credit than he has been given. Completing the task that was given him, working essentially alone and without a large appreciative audience of users, has been the common lot of naturalists on this frontier of North America.

    It was a standard flora of its time, and its use was not intended for amateurs. It remained for someone else to write a field guide, and this is the next story that needs to be told.

    This Handbook’s History

    The need for a book on local flora was evident in 1946 as soon as I found I would be teaching a field course on this subject. No useful text was available. Coulter & Nelson’s Flora of the Rocky Mountains was long out of print; Harold Harrington’s Colorado Flora was far from being completed. The only account of the flora of the Boulder region was a list of the plants of Boulder County, published at the University of Missouri, by Francis Potter Daniels, a visiting professor of classics who spent a few summers in Boulder. His collections went to the University of Missouri (donated finally in 2009 to Herbarium COLO).

    In 1949 I was ready to prepare a preliminary Flora of Boulder County, which I produced on mimeograph. At the time, the Dean of Arts and Science had one secretary who volunteered to help professors do stenographic work in her spare time. I cut the stencils (no erasures!), and Mrs. Suma Service ran them off. Dr. Gordon Alexander donated funds to print a hundred copies. My little daughter Linna helped me collate all the 200-plus pages, which I bound in two black three-ring binders. The work contained keys to the families, genera, and species, and the text contained copious specimen citations. I followed Rydberg’s practice of constructing the keys with left justification rather than the traditional indented keys that are so wasteful of space. We worked on a shoestring budget. These copies were distributed free of charge. There are only a few left, keepsakes in former students’ hands. If one examines the treatment now, it is obvious that at the time my nomenclature was conservative. When I used the name Anemone patens rather than Pulsatilla, Dr. Edna Johnson said, My, you are a lumper, aren’t you? How times have changed.

    George Kelly, a Denver nurseryman and leader of the Colorado Forestry and Horticulture Association, had an office on Bannock Street, and was sitting on a mass of unidentified and unmounted herbarium specimens. He and I gathered a number of lady volunteers and taught them how to mount specimens. One of them was Katherine Kalmbach; the herbarium at the Denver Botanical Garden now bears her name. Dr. E. H. Brunquist, a professor at the Medical School, was an amateur botanist who joined me on some field trips. Moras Schubert, of the University of Denver, was the local plant ecologist who had studied under Dr. George Aikman at Iowa State College. These constituted the leaders of the Denver Botanical Club, and were the spiritual descendants of the young Denver schoolmarm Alice Eastwood.

    George Kelly talked me into turning the mimeographed Boulder Flora into a book, Handbook of Plants of the Colorado Front Range, published in 1953 by the University of Colorado Press, price $4. This was a bare-bones volume with very few illustrations for the glossary. It was printed locally by Johnson Publishing Company, and I visited the place many times to see the pages emerging from hot type. It was a real book. Gone were the specimen citations; the locality data were more general, but there was an adequate introduction and a list of useful reference books. My first royalty check was for 40 cents! A revised, up-to-date second edition with eight more pages was published in 1961. The flyer gives its asking price as $4.00 hardbound and $2.50 paperback. By that time I had begun work on lichens and spent a year in Sweden.

    Acknowledging the success of these books, J. K. Emery, the new director of the press, suggested a new edition that would cover more territory and species and be more attractive. Its binding was blue, with black lettering and a drawing of Jamesia americana on the cover. For illustrations I was able to borrow those made by Charles Yocom, an old classmate at Ames and Pullman. Emory insisted the book must have a name that indicated its broader scope despite the fact that it still covered only the Front Range. Rocky Mountain Flora was a good-looking book, but the paper was much too stiff and it was more of a stay-at-home textbook than a field guide. It was published in 1967, 437 pages, with metric system tables on the inside cover, and an index. I did not like the new title because it did not properly describe the contents.

    The old University of Colorado Press was never supported financially and hardly was more than a small house organ. It barely survived to be overhauled, and was renamed the University Press of Colorado, now a consortium of several universities. John Schwartz became its director, and despite continued low funding, turned the press into a thriving one. John was an old-fashioned publisher who could move mountains with only the help of ants. He turned Rocky Mountain Flora into a real field guide.

    Rocky Mountain Flora of 1972 was now the fourth edition, revised, of the book originally published by the University of Colorado Press in August 1953. It was 438 pages, hardbound, with a color plate of Calypso bulbosa on the cover and the corners neatly rounded. This edition was by far the most successful one, since as the book became more bulky, the fonts became smaller, and the cost prohibited the continued use of rounded corners. This is the edition preferred by older folks, who like the large print and fewer new names to remember.

    The fifth edition of 1976, 479 pages, was outwardly similar to the fourth, but the front cover was graced by a color plate of Primula parryi and the text illustrations were replaced by those of Anne Pappageorge. Added were 32 color plates.

    Pressure built for me to produce a flora for all of Colorado. Fortunately, at this juncture a physicist, Ron Wittmann, came to the herbarium with an interest in floras that was kindled by a course he took under C. Leo Hitchcock at the University of Washington. Ron appeared in the 1970s, and soaked up all he could about the vascular flora, the lichens, and the bryophytes. He has become my field companion, computer guru, proofreader, climbing monkey, general man-of-all-work, and co-author of all the subsequent volumes.

    We accepted the challenge, but decided that it would no longer be a field book if we took on all of Colorado. Instead we decided to write two books, their floras separated by the Continental Divide: Colorado Flora: Eastern Slope (1990) and Colorado Flora: Western Slope (1987). The first edition of the former featured 64 colored plates and 115 line drawings; the latter had a like number of plates and drawings. That pair of books finally arrived at a most satisfactory arrangement of the material.

    The second edition of the two floras—Eastern Slope, 1996, 524 pages, 64 colored plates, and Western Slope, 1996, 496 pages, 64 colored plates—moved all of the line drawings to the end. The third edition—Eastern Slope, 2001, 521 pages, Western Slope, 2001, 488 pages—lacks the colored plates because the press was unable to finance them.

    My Emergence as a Botanist

    If anyone thinks that writing a book entails just setting to work with pen and paper (or computer)—forget it! It takes years of preparation, even before you come on the scene. Colorado Flora for me began in 1930 in New York City, where I was a student at Evander Childs High, which had teachers who took an interest in me. It was also in the days of the birth of recreational bird study, and my field comrades included my own Sialis bird club, and my mentors were folks like Joe Hickey, Ernst Mayr, Bill Vogt, Alan Cruikshank, and Roger Tory Peterson. Here is where I learned the elements of plant and animal taxonomy with the experts in the field, and in the halls of the American Museum of Natural History, the New York Botanical Garden, and the Linnean Society of New York. My most important mentor was my cousin F. Martin Brown (Brownie), the famous lepidopterist, whose career embraced Fountain Valley School of Colorado Springs and Florissant Fossil Beds National Monument. Brownie brought me my first primitive microscope at age five and showed me how to grow one-celled animals in stagnant water. He got me excited about Colorado, but I never dreamed that I would ever see the place.

    I became a botanist at age twelve in New York City and learned the flora with the help of a mentor. My first field guide was Professor M. L. Fernald’s enormous Gray’s Manual of Botany, 7th edition (given me by a high school teacher who had to have it in college but had never cracked it). I became a nature counselor in a Scout camp in the Catskills and the Maine woods. I made an herbarium of the plants of Long Island for a private nurseryman. I was an ardent ornithologist and went on field trips to interesting botanical areas.

    After I escaped from New York during the Great Depression, I had experience with the flora of Iowa, where I worked in the herbarium at Iowa State College, typing herbarium labels (up to forty replicates, no carbons, letter-perfect), for Dr. Ada Hayden under Franklin Roosevelt’s National Youth Administration work-study program. I compiled a checklist of the plants of Story County, Iowa. I entered graduate school at Washington State College, where my assignment was to curate the great collections of Wilhelm N. Suksdorf, whose field notes were mostly in old German script. I collected in the Canadian Rocky Mountains and curated the herbarium in the war absence of its curator Marion Ownbey. I taught the women inmates of the Washington state penitentiary to mount the great backlog of specimens, and did fieldwork on my chosen genera of Balsamorhiza and Wyethia. My doctorate consisted of monographic studies of several genera of wild perennial sunflowers of western North America.

    During World War II I served as a conscientious objector at the Civilian Conservation Corps Camp 21 at Cascade Locks, Oregon, where my latent interest in bryophytes was rekindled, and I became a member of the American Bryological Society. I had field experience in range research in the Blue Mountains of eastern Oregon. By the time I arrived in Colorado, I already had a good knowledge of the floras of Massachusetts, New York, New Jersey, Maine, the midwestern prairies, and the Pacific Northwest, and had been a nature counselor and amateur ornithologist. I already knew much of the Colorado flora without ever having been in the Rocky Mountains.

    My first and only job as a botanist was at the University of Colorado, where in 1946 I filled Joe Ewan’s position as an instructor in the biology department, teaching general biology, genetics, evolution, plant taxonomy, and field botany.

    I had always been interested in flowering plants and bryophytes and, soon after arriving in Colorado, became deeply involved with lichens as well, pursuing them passionately for the next forty-five years. My interest in the fascinating geography of Colorado’s cryptogamic plants, combined with the freedom of travel that allowed me over the years to become acquainted with the world floras, has been invaluable. My botanical experience includes the North American Arctic, Alaska, Mexico, the Galápagos Islands, Chile, Peru, Ecuador, Australia and New Guinea, Scandinavia, Greece, the Canary Islands, Europe, Nepal, Siberia, Japan, and Antarctica.

    An Interesting Incident

    In the old days it was the plants that provided the excitement of fieldwork. The state was our garden; there were no stringent laws about trespass. With a car, cheap gasoline, good uncrowded campsites, motels where one could get by on four dollars a night, strong young legs, good eyesight, and great friends and students, we embarked on a lifetime of joyous discovery. I recall only one expedition that was frightening.

    I had a student from eastern Oklahoma, Joe McCall Anderson, whom I assigned a master’s project, the plants of Baca County, in the southeastern corner of Colorado. We planned a three-day trip to see what we could find. I enlisted Dr. E. H. Brunquist, and the three of us took off on August 30, 1949. We camped at the Dodge Ranch the first night. In the morning it began to rain heavily, and I had the stomach flu. We were unable to navigate the muddy roads, so the boys collected around the camp. Dr. Brunquist was afraid of the campstove, and Joe knew nothing more than how to boil water, so I had to feed the crew.

    The next morning it continued to rain and the campground was getting very soggy, so we broke camp and headed for the corner where Colorado, Kansas, and Oklahoma meet. We had heard that the Cimarron River cut a small corner of sand dunes on the southeastern tip of the state. As we came down from Walsh, we met a farmer named Pfeiffer, who owned the area and advised us on how to reach the sand dunes. The skies were brightening and the storm clouds and lightning retreated northward. I was feeling better. We made camp on the dunes and immediately added three snakes and a dozen plants to the state flora. I cooked up a steak dinner and we watched the sun go down. We were feeling very good. At about 11 p.m. we were about to turn in when we heard horse hooves and shouts: Turn that light on. We know God-damned well you have a light! Come out and identify yourselves!

    Three cowboys came riding up in the dark. We staggered out, Brun in his nightshirt, I in paisley pajamas, and Joe in the buff. We tried to explain, to no avail. One of the men dismounted, went to our panel truck, and yanked open the back door. At that moment the man in the middle spurred his horse and took off upstream. Keep that light on, we’ll be back! Several shots rang out and soon faded away. We just stood there. About ten minutes later, one of the three men came back and apologized. They had caught the rustler and learned that we were not the folks with the gear. Offering us the bottom of a bottle of red-eye, they said good night and rode away into the night.

    Next morning we gathered up our loot and drove north to Lamar, where we stopped at the local drugstore for a milkshake. On the rotating display of cheap books I happened to see a volume, Guns on the Cimarron, and pocketed it to give to Joe later on. But the story doesn’t end here. In the 1990s I attended the October conference at the Missouri Botanical Garden. During a break at a local café on the hill, I shared beer and talk with some friends, including Hugh Iltis, who can bear me out. I told the story straight. When I finished, one of the fellows remarked, I remember that! I was living at the Pfeiffer Ranch. Well, was this a joke or was it real? The fellow answered, No, that was no joke!

    The Field Guide: An Odyssey

    A biologist’s main contributions may all be made relatively late in life, as was true of Darwin, simply because accumulating countless small observations and fitting them into a useful and informative structure inevitably takes a long time. This tendency reflects the complexity of biological systems. The young field biologist collects specimens for taxonomic study, and records their geographic range, general ecology, habitat (with orientation and altitude where pertinent), and associations. Thus, we may pass most of our careers before we are prepared to present a complete picture of a topic. But, because we are dealing with living and genetically variable organisms, the picture is seldom really complete, and in late years we, or others, will add to and modify it. There can be no mathematical proof that the picture is correct; but we are dealing with probability rather than mathematical proof. As time passes we generally discover additional pieces to fit into the puzzle; and the probability of a correct solution finally becomes enormous and can be accepted.

    D.B.O. Savile, Evolution of a Naturalist, 2001:380

    My books on Colorado flora have been a labor of love, but more importantly they have been the way that I and my students and local colleagues have discharged our responsibility to educate the public of Colorado, and ultimately the United States, North America, and the world. Colorado has a flora that is unique in many ways, and its significance is worldwide.

    Being a taxonomic botanist in Colorado has been a rather lonely occupation. Unlike in the bustling eastern and far western states, one has few colleagues with whom to discuss problems. Few American botanists come for a visit, and it takes several days to become acclimated to the altitude. Most of our foreign visitors know about Mount Evans, Hoosier Pass, Mesa Verde, and Rocky Mountain National Park, and appreciate the Eurasian flavor of the flora, because they have received specimens from us. Here on the last of the frontier we have few serious students, the kind who were so important in helping Stebbins, Mayr, Gould, Raven, and Wilson hone their philosophies. We must rely on correspondence, travel, and our slim group of serious amateurs—bless them!

    Fieldwork in Colorado has been for me a very exciting vocation. In this edition of the book I hope to bring some of this excitement into the pages—the accidental discoveries, frustrations, sudden flashes of light when I discovered a Colorado species in the remote mountains of southern Siberia or the subalpine heights of New Guinea.

    The present edition includes the best features of the others. It has paragraphs and notes giving explanations, anecdotes, and other items that readers will find interesting enough to cause them to purchase this latest edition.

    I do not know of another flora or field guide that has been reprinted in so many different editions, ultimately reflecting the fact that a scientist’s points of view change greatly as his experience and understanding increase.

    Thus, this revision reflects my current thinking. It will not please everyone but I hope that it will not discourage amateurs, who have been my targets for so many years, from continuing to love plants and desire to know them.

    The Modus Operandi

    Local floras constructed on the state level are the most convenient field guides. In a local flora many fewer taxa are treated than in a larger regional or national flora, making identification easier. The utilization of paraphyletic genera eliminates the necessity of using subgenera and sections. On the smaller geographical scale these units have a certain reality.

    Local floras can serve better to record the precise habitats of the taxa. They need not include detailed descriptions of the species and genera, for these are available in the larger regional floras and Flora of North America (FNA). Local floras can concentrate on the most prominent distinguishing characters. Because their authors have field experience, local floras can fill in important information about the ecology of the species. In larger floras and the FNA the keys are unavoidably technical, given the size of the geographic area covered, and descriptions are very detailed and designed to be comparable throughout a genus.

    In preparing this revision, we have consulted various sources of information bearing on our justification for including or rejecting geographical reports of taxa. They consist of published accounts (included in our list of references), checklists, catalogs, databases, and examination of voucher specimens, principally in the University of Colorado Herbarium (COLO) and secondarily in the databases of the consortium of local herbaria.

    Checklists are worthwhile if only to give the user a list of names. The lists available are not all of equal quality: some give the names and authorities; some include synonyms; a few include reference to particular specimens and the herbaria in which they may be found. A bare list that lacks synonymy and references to specimens may be useless. Our Catalog of Colorado Flora (Weber & Wittmann 1992) has been cited in the FNA but we feel it is rarely used. To use a checklist as validation of occurrence requires access to the specimen for assurance that the identification is correct.

    Dot maps and their variations must be cautiously employed. The scientific method requires that one be able to trace a suspicious dot to an herbarium specimen. In the early days of monographic treatments, the maps were supported by an index to exsiccati that included brief label data and the standard abbreviation of the herbarium housing the specimen. By and large, these were thought to be wasteful of space, and were replaced by citation of a few representative specimens. Dots that are suspicious, especially those out of the known range, are the ones that are critical. Shaded areas are less reliable than dots. In a recent treatment of Colorado grasses, whole counties are shaded, giving the impression that a species is widely distributed therein, when actually there may be only a single, possibly misidentified, specimen backing up the record, and no source for the information is cited.

    In our field guides, unless we can determine where we can find a voucher specimen, and have had an opportunity to examine it, we cannot include the species. There is a pertinent anecdote about this. Wendell Holmes Camp found that John Kunkel Small’s Manual of the Flora of the Southeastern United States included species ranges that were not backed up by specimens at the New York Botanical Garden. When questioned about this, Small replied, Well, they are bound to occur there, and we might as well list them and save the cost of a revision. We have thoroughly searched the published volumes of the new FNA and traced as many doubtful reports as possible.

    Our generic concepts often differ from those of the FNA since we often recognize paraphyletic groups that are more realistic in a local or regional flora where identification is more significant than abstract phylogeny. We feel that the phylogenetic approach does not take into consideration the various kinds of barriers (cytological, seasonal, ecological, and morphological) that separate the generic divisions. In many instances we feel that the Eurasian concepts have been slighted. In our region we are dealing with a flora that is more closely related to Eurasia than has been recognized. Colorado has relatively few endemic species but a large number of vicariads, especially from Middle Asia.

    Changes in the names of plants are of two kinds, objective and subjective. Objective changes are made because of the International Rules of Nomenclature, which deal, among other things, with the accepted principles of priority. Subjective changes have to do with our opinions as scientists. At the present time, there is a great controversy as to how much weight we should give to the findings of the scientists who are mainly concerned with the evolutionary history of plants, called phylogeny. DNA studies concentrate on molecular evidence, while cladistic studies create evolutionary trees based on morphology while often ignoring other biological factors. A field guide must be useful to its readers, many of whom are dedicated amateurs.

    Our principal decisions here deal with those natural groups (genera) that are easily recognized. Dealing with a much smaller flora than that of a whole continent, these smaller genera are real for us. Technically

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