Colorado Flora: Eastern Slope, Fourth Edition a Field Guide to the Vascular Plants [4 ed.] 9781607321415, 9781607321408

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Altitude Conversion feet 3,000 3,100 3,300 3,500 3,700 3,900 4,000 4,100 4,300 4,500 4,700 4,900 5,000 5,100

meters 914 945 1,006 1,067 1,128 1,189 1,219 1,250 1,311 1,372 1,433 1,494 1,524 1,554

feet 5,300 5,500 5,700 5,900 6,000 6,100 6,300 6,500 6,700 6,900 7,000 7,100 7,300 7,500

meters 1,615 1,676 1,737 1,798 1,829 1,859 1,920 1,981 2,042 2,103 2,134 2,164 2,225 2,286

feet 7,700 7,900 8,000 8,100 8,300 8,500 8,700 8,900 9,000 9,100 9,300 9,500 9,700 9,900

meters 2,347 2,408 2,438 2,469 2530 2,591 2,652 2,713 2,743 2,774 2,835 2,896 2,957 3,018

feet 10,000 10,100 10,300 10,500 10,700 10,900 11,000 11,100 11,300 11,500 11,700 11,900 12,000 12,100

meters 3,048 3,078 3,139 3,200 3,261 3,322 3,353 3,383 3,444 3,505 3,566 3,627 3,658 3,688

feet 12,300 12,500 12,700 12,900 13,000 13,100 13,300 13,500 13,700 13,900 14,000 14,100 14,300 14,500

meters 3,749 3,810 3,871 3,932 3,962 3,993 4,054 4,115 4,176 4,237 4,267 4,298 4,359 4,420

Altitudes of Colorado Towns (meters) Alamosa

2,300

Glenwood Springs

1,751

Pagosa Springs

2,158

Aspen

2,410

Grand Junction

1,398

Pueblo

1,431

2,348

Boulder

1,635

Gunnison

Buena Vista

2,424

Hot Sulphur Springs 2,338

Rangely

1,592

Rifle

1,629

Castle Rock

1,890

Julesburg

1,060

Saguache

2,346

Cheyenne Wells

1,309

Kremmling

2,245

Salida

2,145

Colorado Springs

1,832

La Junta

1,239

Silverton

2,840

Cortez

1,890

Lamar

1,104

Springfield

1,330

Craig

1,885

Leadville

3,094

Steamboat Springs

2,041

Delta

1,512

Lake City

2,643

Trinidad

1,836

Denver

1,609

Limon

1,636

Walden

2,469

Durango

1,985

Meeker

1,905

Walsenburg

1,885

Fairplay

3,034

Montrose

1,766

Wray

1,072

Fort Collins

1,519

Ouray

2,349

Yuma

1,259

Altitudes of Colorado Passes (meters) Apishapa

3,354

Gore

2,903

Loveland

3,655

Red Mountain 3,355

Berthoud

3,449

Guanella

3,557

McClure

2,669

Slumgullion

3,463

Cameron

3,132

Hoosier

3,518

Milner

3,279

Trail Ridge

3,713

Cochetopa

3,058

Independence 3,687

Molas

3,325

Trout Creek

2,849

Cottonwood

3,696

Kebler

3,050

Monarch

3,448

Vail

3,251 3,627

Cucharas

3,030

Kenosha

3,048

Mosquito

4,020

Weston

Cumbres

3,055

La Manga

3,118

Rabbit Ears

2,873

Wilkerson

2,898

Douglas

2,519

La Veta

2,860

Raton

2,388

Wolf Creek

3,307

Fremont

3,450

Lizard Head

3,116

Colorado Flora E a s t e r n S lo p e

Bill Weber holding Ptilagrostis cf. porteri and Saussurea sp., Yabagan Pass. Altai, 1978. Photo by Ivan Krasnoborov, W. A. Weber collection

Colorado Flora E a s t e r n S lo p e F o u rt h E d i t i o n

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 proud member of the Association of American University Presses. 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

 1 0 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 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii Background of Floristic Work in Colorado . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xv Books to Inspire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxvii A Vade Mecum for the Field Botanist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxix Key to the Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Ferns and Fern Allies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Gymnosperms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Angiosperms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 491 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 497 Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 530 Illustrated Plant Structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 550

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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. xi

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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

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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. xv

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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

Background of Floristic Work in Colorado

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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.

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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

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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.

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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 com-

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pleted. 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, Hand­ book 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.

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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

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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 dis-

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mounted, 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.

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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

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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 they are offshoots of the main branches, namely paraphyletic genera. In the FNA many of our small genera are subsumed as subgenera. We feel that subgenera are not easily handled in a field flora. Because this revision will be the last one under our joint authorship, we have made an archive in the COLO herbarium of all the pertinent materials, citations, and other useful literature on which we based our decisions. All of our publications dealing with the flora are available on the Internet. We urge anyone finding novelties, range extensions of native species, or new occurrences of weeds to notify us by email and, most importantly, to deposit voucher specimens at the COLO herbarium. This is the best way we can remain up-to-date. Maintenance requires a community effort, with input from amateurs and professionals alike. Time and manpower limitations have prevented us from doing an exhaustive search of databases and for specimens of new records. The FNA is not completed, but we have searched and evaluated all the Colorado reports in the available volumes 1–5, 7, 8, and 21–26. When we say that we have not seen a specimen, it usually means that there is no specimen record within the borders of our search, which may cover databases and other herbaria. The international abbreviations for herbaria cited in the text are: COLO, University of Colorado, Boulder; CC, Colorado College, Colorado Springs; CS, Colorado State University, Fort Collins; GH, Gray Herbarium, Harvard University; RM, Rocky Mountain Herbarium, University of Wyoming.

Books to Inspire In this book I also try to lure devotees into reading some of the great books of botanical history and adventure. The historical period is being brought to life. Now you can read the journals dealing with the collecting trips of Linnaeus’ students, the struggle in England to move from the Linnean system of classification to a natural one, and how collectors were able to save their specimens in the tropics before the days of modern equipment. You must read! As the teaching of taxonomy continues to disappear from school and university curricula, our amateurs will need to become more serious about what has been a hobby. Learn the main essence of the rules of botanical nomenclature. Become associated with an herbarium or a botanic garden. Help it to survive. Make a genus your special project. Go into the field and seriously begin to make an inventory of a cliff, fen, prairie, or tundra site. And don’t confine your study to the “wild flars” (apology to James Thurber). There are good books on the mosses and the lichens. Consider looking at the fungi, algae, diatoms, and fossil plants! The world is your garden. Don’t waste your time, for life is short! This short list of books is intended to create excitement among my readers, because they are full of stories of the adventure that is field botany: taxonomists, museums, and invasive weeds. They are not merely reference books to remain on a shelf. These are books I have enjoyed. Crase, Douglas. 2004. Both: A Portrait in Two Parts [The lives of Rupert Barneby and Dwight Ripley]. 303 pp. Pantheon Books. Endersby, Jim. 2008. Imperial Nature: Joseph Hooker and the Practices of Victorian Science. 429 pp. University of Chicago Press. Fortey, R. 2008. Dry Storeroom No. 1: The Secret Life of the Natural History Museum. 335 pp. Alfred A. Knopf, New York. Furmansky, Dyana Z. 2009. Rosalie Edge, Hawk of Mercy: The Activist Who Saved Nature from the Conservationists. 352 pp. University of Georgia Press, Athens, GA. Gibbs, George. 2006. Ghosts of Gondwana: The History of Life in New Zealand. 232 pp. Craig Potton Publishing, Nelson, New Zealand. Langenheim, Jean H. 2010. The Odyssey of a Woman Field Scientist: A Story of Passion, Persistence, and Patience. 539 pp. Privately published. Shteir, Ann B. 1996. Cultivating Women, Cultivating Science: Flora’s Daughters and Botany in England, 1760–1860. 301 pp. Johns Hopkins University Press. Slotten, Ross A. 2004. The Heretic in Darwin’s Court: The Life of Alfred Russel Wallace. 602 pp. Columbia University Press, New York. Tiehm, A., & F. A. Stafleu. 1990. Per Axel Rydberg: A Biography, Bibliography, and List of His Taxa. Mem. N.Y. Bot. Gard. 58:1–75. Townsend, John Kirk. 1999. Narrative of a Journey across the Rocky Mountains to the Columbia River. 352 pp. Northwest Reprints. Oregon State University Press, Corvallis, OR. Warren, Leonard. 2004. Constantine Samuel Rafinesque: A Voice in the American Wilderness. 252 pp. University Press of Kentucky. Weber, W. A. 1997. King of Colorado Botany: Charles Christopher Parry, 1823–1890. 183 pp. University Press of Colorado. Weber, W. A. (ed.). 2000. The American Cockerell: A Naturalist’s Life, 1866–1948. 352 pp. University Press of Colorado, Boulder. Weber, W. A. (ed.). 2004. The Valley of the Second Sons: Letters of Theodore Dru Alison Cockerell: A young English naturalist writing to his sweetheart and her brother about his life in West Cliff, Wet Mountain Valley, Colorado, 1887–1890. 567 pp. Pilgrims Process Inc., Boulder. Williams, Roger L. 1984. Aven Nelson of Wyoming. 407 pp. Colorado Associated University Press, Boulder.

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A VADE MECUM FOR THE FIELD BOTANIST Scope of the Book Colorado Flora: Eastern Slope has as its subject the vascular plants—ferns, gymnosperms, and flowering plants—native and naturalized on the entire hydrologic Eastern Slope of Colorado from the Continental Divide to the Wyoming, Nebraska, Oklahoma, and New Mexico borders. It is a rich and varied territory, comprising a large segment of the Great Plains, the foothills and high peaks of the Rocky Mountains, and the interior intermountain “parks.” Edwin James, while participating in the Long Expedition of 1820, was the person to collect plants on Pikes Peak. The botanist C. C. Parry, in 1861 and 1862, explored for the first time the wilderness of South Park and the summits of the mountains surrounding the Clear Creek Valley, where he subsequently named the peaks after the famous botanists of the time: James, Gray, Torrey, Engelmann, and his favorite of all, Mount Flora. Parry Peak nestles within this cluster of magnificent botanical areas.

The Eastern Slope The Eastern Slope of Colorado includes all the land from the Continental Divide to the eastern plains. It includes the plains grasslands, the mountains, and the inter­mountain “parks.” The elevation ranges from over 14,000 ft (4250 m) down to about 3500 ft (1070 m). There are two main drainages on the plains. The South Platte drainage contains charac­teristic species of the northern Great Plains, while the Arkansas drainage is a highway of migration northward of the Chihuahuan floristic elements. Separating these drainages is a relatively low forested divide known as the Black Forest / Palmer Divide. In many respects the flora of the Black Forest resembles that of the Black Hills of South Dakota, inasmuch as the region harbors small relictual colonies of eastern American prairie and woodland species otherwise unknown in Colorado except for some protected canyons of the outer Front Range.

The High Plains, North and South The northern High Plains, or the Platte River drainage, contains several special habitats worth the attention of botanists. The Pawnee Buttes National Grassland contains outcrop­pings of rock that stand as islands in a sea of short-grass plains. Its most unusual feature is a stand of limber pine, Pinus flexilis, isolated at a much lower elevation than expected. This species evidently extended onto the High Plains during the Pleistocene and has been stranded on the Pawnee Buttes following the change in climate that produced the modern High Plains and eliminated the populations in the middle elevations. Its numerous rimrock bluffs support a number of cushion plants and dryland species unable to compete in grasslands. Among these are Leptodac­tylon caespitosum, Oreocarya celosioides, Aletes tenuifolius, and Orophaca sericea. The northern High Plains also contain areas of unstable sand dunes that have a distinctive flora. Some of the special plants to be found here are the bush morning-glory, Ipomoea leptophylla, which has a root as thick as a fencepost. A white phlox, Phlox andicola, covers the ground in season, and roadsides support stands of ring muhly grass, Muh­lenbergia torreyi, whose circular patches look like targets. In the summer and early fall, these sandy areas are glorious with displays of the sand penstemon, Leiostemon ambiguus, Palafoxia, and the common sunflower, Helianthus annuus. The southern High Plains, dominated by the Arkansas River Valley, has as its hallmark the candelabra cactus, Cylindropuntia imbricata, which marches up the valley as far north as Colorado Springs, along with the one-seed juniper, Sabina monosperma. The southeast corner of Colorado, where the Cimarron River cuts a small triangle of sand across it, supports a number of species (the so-called Chihuahuan element) that barely enter our state. The Arkansas xxix

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Valley has extensive outcrops of limestone that support a number of local, often rare, and sometimes endemic species, especially in the area between Pueblo and Cañon City. This is a dry valley, and only in exceptional years does it display its best botanical dress. Large areas of the High Plains have been modified by agriculture and no longer support native vegetation. However, wherever there are bluffs, badlands, and bottomlands, the native flora can still be found. Bonny Reservoir State Park is such an area, and the north-facing bluffs at Wray are good places to visit. Colorado botanists have tended to ignore the eastern plains because of the thunderstorms and tornadoes, the biting midges, and the heat, and have turned most of their attention to the mountains. Eastern Colorado, therefore, has been investigated more by botanists from Kansas, resulting in the 1986 publication Flora of the Great Plains. But the eastern­most tier of counties still will yield new additions to the Colorado flora of plants of the neighboring plains states.

The Mountain Front The Mountain Front includes foothills and mesas bordering the High Plains on the west and south. The southern margin of the Eastern Slope is occupied by high mesas, including the Mesa de Maya, which extends from Trinidad to Baca County and the westernmost part of the Oklahoma Panhandle. This long mesa is forested on the flanks and farmed on top; its lower slopes are covered by dense oak brush. There are deep canyons such as the Purgatoire feeding the Arkansas River system. The Mesa de Maya also has canyons running to the south into New Mexico, in which plants of the New Mexico flora barely enter Colorado. The area is very rugged and difficult to access, and except for two people, C. M. Rogers, working in the 1940s out of Wayne State University in Detroit, and Dina Clark of the University of Colorado, who has been making a continuing inventory for the past several years, it has never been thoroughly explored for plants. Here is where Colorado’s only stands of beargrass, Nolina texana, and mesquite, Prosopis glandulosa, occur. One problem with this area is that good flowering years are few and far between. The Mesa de Maya will continue to be a prime area for the discovery of new Colorado plants. The mountainous southern anchor of the Front Range, Pikes Peak, differs from the northern portion by having bedrock of a very coarse, friable granite that does not hold moisture well. Pikes Peak has several very special species. It is a very good place to see Aquilegia saximontana, two endemics—Mertensia alpina and Oreoxis humilis—and the finest stands of Telesonix jamesii, the latter best seen at Windy Point on the Cog Railroad. The Rampart Range runs north from Pikes Peak to the Platte Canyon and contains many unexplored canyons, especially in the vicinity of the Air Force Academy and Palmer Lake. The Devil’s Head area is noted for isolated stands of rare species, such as Viola biflora on the forest floor, Viola selkirkii, and the sensitive fern, Onoclea sensibilis, which is probably extinct at the only site at which it occurred, west of Sedalia. The Front Range, strictly speaking, is the great mountain mass anchored by Mount Evans on the south and Long’s Peak on the north. Here the Continental Divide reaches its easternmost point on the North American continent as well as its steepest gradient to the Great Plains. It is an area of extreme diversity, beginning, in the Boulder area, with a narrow strip we call the piedmont valleys, interspersed by low hogbacks and cliffs of shale and sandstone, followed on the west by sedimentary rocks of the Dakota Ridge and the Flatiron formations that overlie the Precambrian granite. Hard granite intrusive batholiths form spectacular “narrows” in the local canyons, providing cool and shaded north slopes and cliffs. The canyon of Clear Creek is composed in part of extensive calcareous schists, on which we have found the moss Leptodon smithii at its only known station in North America. Mount Evans is the botanical jewel of the Front Range. To enjoy a vicarious visit to the Arctic, nothing is easier or more rewarding to the botanist than a drive to Summit Lake, 13,000 feet above sea level. Summit Lake lies above timberline in a shoulder of the mountain where

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sunlight only briefly bathes the lake and its inlet and shores. Standing at the inlet, one cannot see out over the tundra to the plains or the surrounding mountains, and it is as if one were in the true Arctic. This fragile wet-tundra site has become an object of botanical pilgrimages by botanists from all over the world because of the exciting alpine species that occur here and often no other place in Colorado, recurring again only in the Arctic or in the Eurasian mountains. Summit Lake became so renowned for its botanical treasures that it was designated in 1965 as Colorado’s first Registered Natural Landmark. To ensure its preservation, a plaque was anchored to a lakeside boulder, fishing and stocking of trout were discontinued, and huge boulders have been placed across the former vehicle access to the shore. For several years, each visit to the lake turned up some new and exciting botanical find. The list includes rare and disjunct mosses, lichens, and liverworts. Among the vascular plants, one can see Phippsia algida, Kobresia sibirica, Spathularia foliolosa, Ranunculus pygmaeus, and many other rare wettundra species. Unfortunately, a large population of the introduced mountain goat threatens this botanical Eden. From Echo Lake to the summit, the drive up Mount Evans traverses subalpine forests; a renowned stand of gnarled bristlecone pine, Pinus aristata; dry, rocky tundra; massive talus block slopes; and a summit area of dwarf cushion plants including Claytonia megarhiza, Draba species, Taraxacum scopulorum, Mertensia, and Eritrichum. From the summit, one has a panoramic view of the Front Range, the Great Plains, Gray’s and Torrey’s peaks, and the expanse of South Park. The crest of the Front Range from Mount Evans to Rocky Mountain National Park is accessible and botanically very interesting. In Boulder County alone there are more than 1,500 species of flowering plants. We believe that considering all groups of plants, including the lichens and mosses, this is the richest area of Colorado, and would repay an interested amateur or profes­sional botanist an entire summer’s stay. North and south of the Front Range the Continental Divide recedes to the west. This is very apparent, especially to the north of Fort Collins, where the foothills take on a decidedly desert-steppe aspect. At Owl Canyon, north of Fort Collins, is an isolated stand of piñon pine far from its northernmost outpost at Colorado Springs. Confined to a surface outcrop of limestone that has been quarried for many years, a large part of the popula­tion was destroyed, but this stand is now a preserve. However, it is not a natural occurrence but dates back, based on archeological evidence, to an accidental planting of seeds evidently taken by squirrels or cached by Native American traders bringing seed from the south. Finding no competition on this bare outcrop, the trees thrived. The oldest trees, about 350 years old, consist of a small handful in a ravine near the north end. North of Fort Collins to the Wyoming border is a large arid highland grassland with eroded pinnacles of sedimentary rock on the east and granite domes on the west. This area is not very well explored for plants, and it is here we can expect to find some species of the Laramie Plains of Wyoming dropping over the Colorado line. One of these is Besseya wyoming­ensis. Serious study of this area is suggested.

The Intermountain “Parks” A unique feature of the Southern Rockies is the chain of intermoun­tain “parks” that divide the mountain ranges. From north to south, the principal ones are North Park, Middle Park (on the Western Slope), South Park, and the San Luis Valley. These are essentially grasslands, with dry and wet facies supporting steppe plants and fen plants, respectively. They tend to be cold-air drainage basins, hemmed by high mountains on three sides, and draining into one of the major river systems. Each park has a personality of its own.

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North Park North Park is a combination of dry sage plains, willow and sedge meadows, lakes, a few small scattered mountains, clay buttes, and sandy river benches. A shifting dune area lies on the east side against the confining Rawah Range. The Continental Divide forms the southern (Fairview Mountain) and western (Park Range) boundaries. Tributaries of the North Platte River drain the park to the north into Wyoming. While the basin of North Park looks rather uninteresting botanically, it supports a rich flora of sedges and willows, and aquatic plants of both fresh and alkaline requirements. The Park Range is easily accessible by trails along the base and up almost every valley. Here is where many plants characteristically found only on the Western Slope drop down on the east side of the Park Range. We have just begun to find them, and for this reason a number of plants thus far not reported are listed as likely to be found in North Park. The Big Creek Lake area is particularly rich in these. Trillium ovatum, Piperia unalascensis, Mimulus lewisii, M. moschatus, and Azaleastrum albiflorum occur along the flanks of the Park Range. North Park has its own endangered species, Phacelia formosula, known from a half dozen small populations near Walden. One of the most spectacular displays of wildflowers may be seen on a tour of Independence Mountain, where whole hillsides are covered by golden colonies of Helianthella uniflora and purple lupines.

South Park South Park and its surrounding unglaciated mountain ranges, some granitic, some calcareous, comprise the crown jewels of the Southern Rocky Mountains. Here are concentrated more rare and disjunct alpine, subalpine, and wetland species than anywhere else. South Park is a large grassland and wetland park bounded on the north and west by probably the richest alpine botanical area in Colorado. Hoosier Ridge, between Fairplay and Breckenridge, has many botanical treasures, but one must walk up the tundra to see them. The east side of Hoosier Pass is granitic, and the west (Mosquito Range) calcareous, so a different flora awaits in each direction. On Hoosier Ridge one can see Armeria scabra, Ipomopsis globularis, Eutrema edwardsii, Oxytropis podocarpa, and O. viscida; on the calcareous slopes of Mount Bross and Mount Lincoln, Braya humilis, Trifolium attenuatum, Parnassia kotzebuei, and Physaria alpina. And penetrating the Mosquito Range to Mount Sherman and Horseshoe Cirque, more aristocrats may be found: Askellia nana, Eriophorum altaicum, Salix calcicola, Saussurea weberi, and Trichophorum pumilum. South Park is bounded on the east by the Tarryall Range, a rugged but relatively low range most famous for its mountain sheep, and on the west by the Mosquito Range and the volcanic Buffalo Peaks. On the south, the park empties west into the Arkansas drainage through the low Trout Creek Pass, and eastward out to the main South Platte. Antero Reservoir lies in a large alkaline flat; here a rare mustard, Thellungiella salsuginea, is relatively common. Aside from its main range in eastern Siberia, this is known from only a few North American localities. The northern basin of South Park, until very recently, was a great expanse of wet meadows that were harvested for wild hay. Through the season these meadows were changing carpets of color formed by dense stands of elephantella (Pedicularis groenlan­dica) and P. crenulata, bistort (Bistorta bistortoides), marsh-marigold (Psychrophila­ leptosepala), and fringed gentians (Gentianopsis thermalis). But recent drainage and ditching of these meadows to supply water to the expanding Denver suburbs have dried them out and replaced many of them, temporarily we hope, with an uninterest­ing flora of silver sage (Artemisia frigida). In the middle of the park is an area of grassland punctuated by scattered small stands of Picea pungens. These trees grow with their feet in the water, in the midst of quaking fens fed by streams flowing out of the calcareous Mosquito Range. The water is quite sulphurous. Shallow pools occur within the extensive sedge and willow stands. Here we have a remarkably

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rich assemblage of rare boreal disjunct species: rare willows, Salix candida and S. myrtillifolia; rare sedges, Carex scirpoidea, C. viridula, and Tricho­phorum pumilum; and primroses, Primula incana and P.  egaliksen­sis. Another rich, but not calcareous, fen area in East Lost Park, in the adjacent Tarryall Range, displays extensive quaking fens dominated by the rare cotton-grass, Eriophorum gracile, up until recently thought to have become extinct in Colorado. With it occur two very rare sedges, Carex livida and C. tenuiflora, widely disjunct from their nearest localities in boreal North America. And on the fringes of the wetland, in cool, north-facing forest edges, the remarkable lichen Cladina stellaris occurs in small patches at its only station in the western contiguous United States. This beautiful little plant is a dominant understory in the subarctic muskeg, and most people recognize it as the little rubberized dyed-green “trees” that are used in miniaturizations of realtors’ three-dimensional plans.

The San Luis Valley The San Luis Valley is the largest of the parks, bounded on the east by the Sangre de Cristo Range and the Great Sand Dunes in the north and the Culebra Range in the south; across its north edge by Poncha Pass; and on the west by the La Garita and Cochetopa Hills in the north and the San Juan Mountains in the south. The Rio Grande arises in the San Juans and flows out of the valley southward into New Mexico. The valley has a rich supply of artesian water, and the basin is very sandy and alkaline, full of introduced weeds. Interestingly enough, one of the weed genera, Descurainia, produces a bushy-branched type in the valley bottom that was recently described as a completely new species, D.  ramosis­sima. A very few endemic species are known in the San Luis Valley. Astragalus ripleyi occurs at the southern end, in Conejos County. Otherwise it ranges south to Tres Piedras, New Mexico. When first discovered here, it was abundant along roadsides, but has recently been disappearing because it appears to be selectively grazed by cattle. Recent studies of the flora of the western side of the Valley at the base of the San Juans show that, floristically, those foothills are somewhat of an extension of the Front Range.

The Sawatch Range and Upper Arkansas Valley The desert-steppe vegetation of the lower Arkansas Valley follows the river almost to Leadville, and it is most surprising to see that the piñon pine climbs the south-facing slopes as high as 9000 feet along the road to Monarch Pass! The Sawatch Range includes the Collegiate Peaks, Mount Elbert, and Mount Massive. We have few or no collections from these high peaks, but recently a great deal of interest has been generated by the limestone and dolomite exposures in the tundra around Cottonwood Pass and Mineral Basin, where new discoveries of Braya glabella and B. humilis were recently made. It seems obvious that more attention needs to be paid to limestone areas at high elevations.

The Wet Mountain Valley and Spanish Peaks The Wet Mountain Valley is a small park surrounded by the Sangre de Cristo Mountains on the west and the Wet Mountains on the east. It is a beautiful, relatively unspoiled grass- and wetland, draining, by way of Texas Creek, into the canyon of the Arkansas River. This is a fine area for botanical exploration because one can headquarter at the old village of Westcliffe and reach rich areas easily. The tundra is accessible by several roads. The valley is not noted for endemics, but it is an excellent place to see Coriflora scottii in openings in the oak brush. Draba smithii, an endemic of these southern mountains, occurs on alpine ledges. The Spanish Peaks, considered to be part of the San Juan Volcanic Belt, belong to a volcanic system somewhat related to that of the San Juan Mountains of western Colorado. Several species common to both areas occur here, including Castilleja haydenii. Rydbergia brandegeei replaces R. grandiflora in the area.

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Floristic Zones Floristic zones are rough estimates of the altitudinal coverage of the easily recognizable plant communities. They are not always consistent in mountain regions because of a phenomenon called environmental compensation. There is a telescoping effect of altitude combined with slope exposure. On a south-facing slope, species of low altitudes may climb very high, and on north-facing slopes high-altitude species may reach very low altitudes because of various factors that define their ecological requirements. Statements of the floristic zone occupied by a species should always be allowed some latitude. An especially protected cool, north-facing canyon side may actually harbor typically alpine species, while an alpine tundra site with easily warmed sedimentary rock substrate and soil churned up by gophers may support typical subalpine vegetation. A well-known phenomenon of mountain floras is the transportation of high-altitude species downstream to lower-than-usual altitudes by spring floods. These waifs may survive for a year or two, but usually are selected against by the climatic norms. Mosses and lichens of high altitudes, however, are often found in cool, moist pockets on north-facing slopes of canyon sides, where they have probably survived for millennia. In this book we use a loosely construed group of floristic zones. Desert-steppe is the treeless semiarid canyon side, river bench, or talus of the lower river basins. Riparian woodlands are wetlands along the major streams. Piñon-juniper refers to the plant community dominated by these species (there seems to be nothing gained by changing the name to piñon–red cedar although technically these are not Junipers). Sagebrush refers to sites dominated by various species of Seriphidium (formerly part of Artemisia). Montane refers to the middle­altitude, relatively dry forested zone; subalpine to the spruce-fir zone below the limit of trees; and alpine to the area above the limit of trees. The American concept of subalpine and alpine zones is not the same as that used in Europe. Ruderal is a term used for sites much disturbed by the activities of humans. One should always expect to find plants growing above or below the limits given by this book. In floristic books, assigning limits to altitudinal distribution is simply an invitation for someone to prove the datum wrong.

Plant Geography The Colorado Rocky Mountain region is like a huge flattened wheel with a hub—the Rocky Mountain chain—running north to south and presenting a potential highway for plant species to move along it. Thus, it is not surprising that the majority of our high-mountain species are also found to the north and to the south at least as far as the terminus of the range near Santa Fe, New Mexico. We call this the Rocky Mountain element. Because the Rocky Mountains are less continuous to the south and because of events in geoclimatic history, we have relatively few species, if any, of a Mexican mountain element, although at one time many of our species did extend into the Mexican cordillera and are still present in small patches there. Continuing with the concept of the Rockies as a hub with radiating spokes, we see elements of other floristic regions extending into Colorado along the river valleys (the spokes) from the Great Basin, Uintah Basin, Colorado Plateau, Rio Grande Valley, Chihuahuan desert, Northern Rockies, and northern and southern Great Plains. At least one genus, Psychrophila, seems to be an Andean-Australasian group of which few other vestiges remain, including the lichen Toninia bullata. Lupines and paintbrushes also occur commonly in the South American Andes. The Northern Rocky Mountain element is especially interesting because it is restricted to the Park Range. On the west side of North Park, a number of species common in the Northern Rocky Mountains and Pacific Northwest survive in this very mesic mountain area and must have once extended north across what is now the Wyoming desert. Characteristic

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species include Azaleastrum albiflorum, Drymocallis glandulosa, Perideridia gairdneri, and Trillium ovatum. The Southern, or Colorado, Rocky Mountains have virtually no so-called amphi-Atlantic connections of species common to northern Europe and northeastern North America. Nevertheless, there are some curious plant distribu­tions involving Greenland and the Rockies, such as Sisyrinchium montanum, Draba aurea, and Festuca saximontana. This is not surprising, since Greenland was another great high-mountain mass before the Pleistocene, probably harboring a large reservoir of species common to Asia and western America. But some Rocky Mountain species occur in isolated pockets around the Great Lakes. At one time the populations were believed to have ranged across North America and were eliminated by glacial action, surviving only on ice-free nunataks. Whether this is so is still a matter of controversy. Because the Colorado Rockies are a massive, highly buffered system of moun­tain ranges whose units are isolated from each other by intermountain basins, it is to be expected that some species evolved in isolation and are restricted to a particular mountain mass. These are endemics. Endemics occur not only on mountain masses but also in isolated areas of the lowlands, particularly in connection with areas of high aridity and specialized rock strata. Species confined to the Southern Rockies, for example, but widely distributed within the area, qualify as broad endemics. Many of our common species are broad endemics. Species confined for whatever reason to a small area are called narrow endemics. Relatively few high mountain species fall in this class; Ipomopsis globularis is one. Narrow endemics are more characteristic of lower altitudes, arid climates, and special substrates. Disjunct genera and species are very important in the mountain flora. Some of these must be very ancient survivors of the Tertiary Period long before Pleistocene glaciation. Most of our disjunct species are absent between Colorado and the Canadian Rockies, Alaska, Eastern Asia or the Middle Asiatic Altai mountains; Ptilagrostis and Leucopoa are such examples. Certain alpine areas are more rewarding than others. In our experience the major northsouth-trending ranges tend to endure severe environmental stress, especially because of the drying winter foehn winds, while high east-west trending connecting ridges such as that from Mount Evans to Hoosier Pass are protect­ed from them. This, and the different exposure of the side ridges and canyons to the daily march of the sun, tends to permit these areas to remain more moist through the season as a result of slower snow-melt. In such areas the south slopes are quite as mesic as the north slopes. The greatest concentrations of rare alpine species occur in these areas.

Endangered Habitats Concern with the extremely rapid deforestation of the tropics and the concomi­tant urban development in the western United States (made possible in part by this very deforestation) has produced a nationwide movement for the preserva­tion of rare, threatened, and endangered species. The concept of rarity is discussed below. However, in Colorado our concern really should be with special or unique habitats, which in turn support populations of rare plants. If we are able to prevent destruction of their habitats, we probably need not worry about the plants. Unfortunately, many important habitats for rare plants are going by the board. For example, while environmentalists were protesting the planned im­pound­ment and filling of the spectacular canyon of the South Platte River for Two Forks Dam, no one seemed to be concerned that an Eastern Slope city had acquired water rights in South Park and, within a year or two, had destroyed, by ditching and draining, most of the wetlands of northern South Park, where a number of very rare species occur. The most severely altered area on the Eastern Slope is the urban corridor between Fort Collins and Colorado Springs. It is unfortunate that some large area of the Palmer Divide was not set aside to preserve the unique mixture of High Plains and foothills species.

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Rarity We tend to omit statements about relative abundance because the term “rare” has many connotations, and because even rare plants are often abundant where they occur. If we say something is infrequent, someone will soon find it to be abundant. We have been unable to walk over every inch of Colorado. If a species is restricted to Colorado (endemic here), we generally say so. If it is really threatened by collecting, we make a note of this. General­ly, our rare plants (except for cacti) are threatened less by collectors than by bulldozers and drainage ditches, which destroy the habitats. Rather than using the word “rare,” we prefer to use more precise words such as “infrequent,” “locally abundant,” or “local.” Rarity actually is the end result of many interacting factors. These may be characteristics of the species itself: its genetic diversity or homogeneity that may make it resistant or vulnerable to changes in the habitat or environment; its mode and recentness of origin; and its “reproductive strategy.” Others may have to do with externals: alteration of the habitat, either natural (fire, flood, climatic change) or induced by the hand of man (introduction of exotic animals, pollinators, introduced plants), and exploitation of wild plants for food, drugs, or firewood. Detailed studies of the autecology and population biology of rare plant species of Colorado have only begun. But changes in the habitat occur daily. In an important and thoughtful review of the concept of rarity, Fiedler (1986) dis­cusses in detail the concepts men­tioned here. Among other things, she shows that the rarity of Calochortus not only involves unique habitat requirements, but probably resulted in large part from the commercial bulb collection of many thousands by a single dealer and his overzealous crew. Everyone working in plant conserva­tion should study this paper. We must refine our definition of “rarity” and seek to learn its basis. Each species is a unique instance; generalizations are impossible. In Colorado there are several apparent reasons for rarity. In the high mountains, small populations have survived intact since Tertiary times (see Weber 2003) because their microhabitats have remained in place or have moved only slightly over millions of years. Our Asiatic rarities belong to this group. Their habitats include limestone outcrops, alpine wetlands, and virgin forests in areas untouched by glaciation. Since neither the hand of man, even in the mining days, nor sheep grazing has destroyed these species up to now, few alpine plants are threatened or endangered. In the desert-steppe, rare and endemic species may be restricted to certain types of rock or localized seleniferous or gypsum soils in particularly critical moisture regimes. We do not understand the specific combinations of factors that cause species to be so restricted, but can often provide educated guesses. Because these sites are localized and subject to massive injury from quarrying and recreational vehicle use, many of these plants may prove to be truly endangered. Again, each species is a unique case, and much study of the biology of the plant is required, not just fencing off a few acres. But only a few species are rare here because of past human use. The male fern was almost exterminated by pharmaceutical collectors. At one time the Colorado columbine was thought to be in great danger from people who gathered enormous bouquets on Sunday drives. Thus we have the only state law protecting a wild plant in Colorado. The habit of wholesale wildflower gathering is a thing of the past. More recently, however, some rare bog plants have been endangered by digging of peat bogs for Denver gardens. Topsoil has been stripped from foothill canyon slopes, and barrel cacti and banana yucca have been exploited for landscaping of shopping malls and residences. The preservation of crucial habitats alone is not sufficient. These sites must be adequately buffered so that the effect of change on adjacent sites will not wash over and affect a protected habitat. Unfortunately for the Eastern Slope, some of the most critical areas are being affected by human interference: the draining of South Park for the water needs of Front Range corridor cities and developments; the excessive use of the most pristine foothills around Boulder by mountain bikes, hikers and their animals, and climbers; the overpopulation of elk in Rocky

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Mountain National Park; the growing tension between homeowners and the larger wildlife; the introduction of non-native mountain goats, with no natural predators, into the most sensitive alpine areas of Mount Evans, including Summit Lake, the first site in the U.S. designated as a botanical landmark; the destruction of prairie dog towns with consequent loss of habitat for raptors, burrowing owls, and ferrets; and the covering-over of lowland agricultural areas by home developments and golf courses. In conclusion, we reiterate that it is the critical habitats that need to be preserved. Then we may have time to study over the long period the various features of the biology of plant species that make them rare, and be better able to make intelligent rather than emotional decisions for their preservation.

Revegetation Despite claims to the contrary, it is virtually impossible to revegetate an area in such a way as to reconstruct the original ecosystem. Therefore, the wisest action is to avoid disturbance in the first place. Attempts are commonly made nowadays to reclaim land denuded by development or exploitation by seeding to “native” plants. From a scientific botanical standpoint this is often regrettable, since it alters the racial mixes even of species native to this area. Because of the fact that records are seldom kept, the history of vegetational change of this kind is hard to reconstruct. Depending on the source of seed, weeds known to be potentially harmful may be introduced. Native species from other regions, if successful, may become undesirable weeds. My personal preference would be to allow the land to reclaim itself, providing the development has not produced an erosion crisis. In Australia enormous amounts of money have been spent in order to reverse the effects of one or two introductions (Opuntia cacti, Himalayan blackberry, and rabbits). This could easily happen here as a result of the complete­ly uncontrolled introduction, by well-meaning but botanically ignorant environ­mentalists, of plants for revegetation purposes.

Alien Plants A large proportion of the plants growing along roadsides, in vacant lots in towns, and on disturbed ground like road fill, mine tailings, and heavily used trails are not native to Colorado. Except for rare instances, we do not know when, how, or from what precise part of the world they came. It is a fact, however, that Eurasian plants seem to be happier here than native American plants tend to be in Eurasia. Whether or not we call them weeds is colored by how abundant they are, whether they displace native species, and how long they persist or spread. The roadside plants of Colorado must make our European tourists feel right at home! In the first edition I used the term “adventive” to denote any species that are not native in the area covered by this book. Adventive might mean deliberately introduced, escaped from a garden, or accidentally brought in by various means. Because we usually do not know the history of an introduction, it is probably better to refer to non-native plants as “aliens.” This is becoming common practice in ecological publications. We are often asked, “How do you know if a plant is native or not?” This is a question that is very difficult to answer in general, but not very difficult on a case-by-case basis. For example, we have ways of knowing whether a genus has ever been native in North America; we have case histories of weeds that first appeared on ballast dumps along the East or West Coast, and we know that they followed the railroads west­ward. Very recent introductions are fairly easy to explain. After all the easy questions are answered, several difficult ones remain, especially along the Front Range. Several species, native in eastern North America, occur sporadically near the cities of the Front Range, particularly Boulder. We know that Darwin M. Andrews had a nursery at the south edge of Boulder and that he specialized in native plants. We feel fairly sure that Sambucus canadensis, Virgulus novae-angliae, Scirpus lineatus,

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and possibly Impatiens capensis spread from his garden along the irrigation ditches. However, plants that were really native but just hanging on in a few mesic pockets may have gotten a new lease on life by the white man’s development of irrigation ditches, because now they seem to exist only thanks to these ditches. Two possibilities are Thalictrum dasycarpum and Eutrochium maculatum. The mining era saw a number of introductions. The Clear Creek Valley is now full of the yellow oriental clematis, originally grown alongside a house in Idaho Springs for its ornamental value. Butter-and-eggs, Linaria vulgaris, and bouncing bet, Saponaria vulgaris, were common garden flowers. Look at them now! We also believe that early settlers in the Boulder Valley, lacking irrigation, planted some of their ornamental plants in the creeks, such as Bluebell Gulch, near their homes, where they would get Nature’s water. How else can we explain the ornamental iris, Oriental poppies, Viburnum lantana, Viburnum lentago, and Berberis vulgaris that persist there today? More species are being introduced in the urban corridors at the present time than ever before. Significant alterations are being made in the Colorado landscape. The bulldozer and its mass movement of soil tend to transport seeds and roots far from where they originate. Revegetation of burned areas and scattering of wildflower packets and instant “meadows-ina-can” bring in new plants that are not native to the area. Many of them are undesirable; most do not survive, but some, unfortunately, compete too successfully with the native vegetation that might return naturally if given a chance. Jays and magpies carry seeds of ornamental plants from the cities out into the foothills, where some sprout and persist or actually thrive. Boulder is a case in point. Many years ago Ernest Greenman planted oaks on Green Mountain to see if they would survive north of their normal geogra­phic limit. They certainly do, and more and more of them are appearing, probably started by acorns carried by the birds. In Long Canyon we recently found a large Cotoneaster bush, a species commonly cultivated in Boulder, and a fairly large red oak. On the foot of one of the Flatirons, a scarlet oak has established itself, and the cultivated rose with purple foliage now occurs nearby. It is an interesting fact for the consideration of our own flora that alien species have their specific ecological preferences. In their homelands they are usually not weeds. Roughly speaking, the weeds of California tend to be Mediterranean in origin and environmental requirements; those of New England tend to come from Europe; those of Texas from Mexico and South America; those of Salt Lake City from some area in the Middle or Near East. The weeds of Colorado very often are Asian (southern Siberia, Caucasus, Bulgaria, etc.). A fair number of our alien species were brought to Colorado by the Spanish travelling northward from Mexico. Invasive species often begin as cultivars in municipalities. Treating escaped cultivar species in a field guide to the flora is a gray area. How do we know whether a species is naturalized? Does it have to be naturalized to be a potential noxious weed? We expect that reports of cultivars in the wild will increase alarmingly in the future, and the herbarium voucher of a first occurrence is an important record. If one encounters a cultivar that is suspect, the best thing is to get rid of it immediately. It is impossible to continue to update a field flora every time a new weed is found. Such discoveries are bound to increase as urbanization continues and vectors of transportation burgeon. Such discoveries should be reported to the agencies whose charge it is to eradicate noxious weeds. Municipalities should be aware of the potential of garden plants and advertise the need to control their spread outside town limits. Here is a list of the most aggressive aliens in the Colorado Flora, the species that have been introduced relatively recently and that need to be brought under control or eliminated. Not mentioned are some that have been pests for a long time and that we have gotten used to, like Canada thistle, cheatgrass, and bindweed; these appear to have reached a sort of balance.

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Acosta diffusa, diffuse knapweed Acroptilon repens, Russian knapweed Berteroa incana, hoary-alyssum Caragana arborescens, pea tree Cardaria latifolia, tall whitetop Dipsacus spp., teasel Hypericum perforatum, St. Johnswort Linaria genistifolia, butter-and-eggs Lythrum salicaria, purple loosestrife Onopordum tauricum, Scotch-thistle Salvia aethiopis, Ethiopian sage Tithymalus esula, leafy spurge Tithymalus myrsinites, donkeytail spurge Ulmus pumila, Siberian elm

Acosta maculosa, spotted knapweed Ailanthus altissima, stink weed Bromopsis inermis, smooth brome Cardaria draba, whiteweed Carduus acanthoides and C. nutans, thistle Elaeagnus angustifolia, Russian-olive Leucantha solstitalis, yellow starwort Linaria vulgaris, butter-and-eggs Onopordum acanthium, Scotch-thistle Potentilla recta, cinquefoil Tamarix spp., tamarisk Tithymalus uralense, leafy spurge Tripleurospermum inodorum, mayweed

Some Floristic Statistics The total number of native and alien species, ignoring varieties and subspecies, is roughly 2300. Of these, 696 are found only on the Eastern Slope. The larger families have more than 25 species. This means that, while we have 155 families to reckon with, about 75 percent of the taxa fall into the big families. If you can learn to recognize these families, you can avoid using the family key that much of the time. Success in using a field guide comes when one knows the large families without using the key.

How to Recognize the Big Families Unfortunately for beginners, the large families are also the most diverse and the least clearly defined. An understanding of them comes from learning first some species, then seeing that they are related as a genus, and gradually getting a feeling for the family by association rather than by memorization of a group of characters common to the family. A large family cannot be narrowly defined. Just as a species is a group of closely related individuals, a genus is a group of closely related species, and a family is a group of closely related genera. Of course, we like to measure relationship roughly by morpho­logical similarity, but other important factors are involved. Here are some hints for recognizing the largest families. Warning: there are exceptions to almost all of these statements! Alsinaceae (35 species): Herbaceous, leaves opposite, simple, nodes swollen, flowers 5-merous, calyx of separate sepals, petals usually white, ovary with free-central placentation. Apiaceae (41): Flowers in umbels, ovary inferior with 2 1-seeded mericarps separating at maturity. Leaves usually compound, aromatic (soap, anise, carrot). Asteraceae (378): Flowers in dense heads subtended by an involucre of bracts. Ray-flowers are good field marks but they may not be present. Heads need not be showy, but can be green and very small, as in the ragweeds and sagebrushes. Other families that mimic Asteraceae are Valerianaceae and Dipsacaceae. Boraginaceae (39): Flowers radially symmetric, short- or long-tubular, 5-merous, ovary with 4 distinct nutlets; foliage, except for Mertensia, with stiff hairs. Brassicaceae (119): Mustards have a cross-shaped corolla, with 4 sepals, 4 petals, 4 long and 2 short stamens. Ovary superior (inferior in Onagraceae, another 4-merous family) with 2 compartments separated by a thin membrane (replum) to which the seeds are attached. Most mustards lack bracts beneath the individual flowers.

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Chenopodiaceae (45): Flowers greenish, radially symmetric, petals none, ovary 1-seeded; leaves often with a mealy coating. Cyperaceae (154): Sedges usually have 3-angled stems, but some have terete ones, and 3-, hardly ever 2-ranked, leaves. The unit of inflorescence is usually a spike, each flower consisting of 3 stamens, an ovary with 2 or 3 styles, a subtending bract (never a lemma and palea) below each flower. Fabaceae (143): Possess a type of fruit common to all, a legume, exemplified by the pea or bean. Most of ours have papilionaceous flowers (like sweet-pea) with banner, wings, and keel, and 10 stamens of which 9 are united and 1 free (there are exceptions), but some have flowers lacking petals, or with many stamens. Leaves are usually, but not necessarily, compound. Juncaceae (38): Grass-like, but leaves often terete or like Iris. Flowers have all the parts of those of a lily, but they are small and brown or green. Lamiaceae (34): Leaves opposite, stem square (but see also Scrophularia), plants usually aromatic; flowers zygomorphic, ovary formed of 4 nutlets. Onagraceae (42): Flowers 4-merous, as in Brassicaceae, but stamens usually 8, and ovary inferior. Poaceae (265): Grasses have leaves 2-ranked, with a sheath, blade, and a ligule where the blade meets the sheath. The unit of inflorescence is the spikelet, which consists of a pair of glumes subtending 1 or more florets, consisting of a lemma, palea, 3 stamens, and an ovary with 2 feathery styles. Compare with Cyperaceae, Juncaceae, Juncagin­aceae. Polemoniaceae (35): Herbaceous, flowers 5-merous, styles 3. A family not easy to characterize. Think of Phlox. Polygonaceae (55): Except for Eriogonum, the presence of an ochrea (sheathing stipule) is characteristic. No differentiation between petals and sepals, fruit a 3-sided achene. Ranunculaceae (42): Herbaceous, flowers with often undifferentiated colored petals and sepals, numerous stamens and carpels (achenes). Rosaceae (73): Radially symmetrical flowers, with 5 petals and sepals, many stamens, a hypanthium, and superior ovaries (achenes or follicles). When in doubt, Rosaceae generally have stipules, Ranunculaceae do not. Salicaceae (37): Shrubs or trees, leaves alternate; catkins formed in spring. Willows have a single scale covering the bud, cottonwoods several. Saxifragaceae (28): Herbaceous, flowers 5-merous, radially symmetrical, but occasionally a few petals longer than the others, ovary with 2 united carpels, tending to be half-inferior. A difficult family to characterize, best understood by learning first one, then another, until you get a feel for it. Scrophulariaceae (92): Flowers are, with few exceptions, bilaterally symmetri­cal, leaves opposite, stem terete, ovary not lobed, with 2 internal compart­ments (compare Lam­ iaceae). Scrophularia has square stems!

Families with a Unique Character Distinctive odors or flavors: Apiaceae, Asteraceae, Brassicaceae, Lamiaceae, Oxalidaceae Milky juice: Aceraceae, Apocynaceae, Asclepiadaceae, Asteraceae (dandelion group), Campanulaceae, Euphorbiaceae, Papaveraceae Poisonous juice: Anacardiaceae (Toxicodendron), Apiaceae (Cicuta, Conium), Euphor­biaceae Spines or thorns: Asteraceae, Berberidaceae, Cactaceae, Rosaceae, Solanaceae Stinging hairs: Urticaceae, Euphorbiaceae (Tragia)

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Scientific Names Different editions of this book reflect refinements of botanical understanding and differences in points of view concerning relationships. As a result, nomenclature is not static and names do change. Here we have attempted to account for some name changes that affect Colorado by listing synonyms. For more complete coverage, see Weber & Wittmann, 1992, Catalog of the Colorado Flora. This catalog is also available and continuously updated at the web site: http://cumuseum.colorado.edu/Research/Botany/botany_databases.html For those who are interested, this site also contains complete listings of lichens and bryophytes, and bibliographic references. Further questions about the flora may be addressed to the University of Colorado Herbarium, Boulder, CO 80309.

Pronunciation In the pronunciation argument, we take the position that the most important question is whether one is being understood. Particularly, the English and American pronunciations of scientific names, whether “purist” Latin or not, seem to be utterly unintelligible to people from other parts of the world. Since the Rocky Mountain flora attracts so many Eurasians because of the similarity of their floras to ours, we should make an attempt to meet them halfway with pronunciation, because otherwise when we pronounce words like nuttallii with four syllables ending with a long I, they simply close their ears. We find that Americans and Britons are much more likely to try to understand a European’s pronunciation than vice versa. Since this is the case, we should try to reach them with something like their language, which in many ways is really more intelligible than ours. Here are our basic rules: The letter a should always be pronounced “ah.” The letter e should always be pronounced “eh.” The letter I should always be pronounced “ee.” This goes for the double ii, which may be given as either one or two syllables. The letter c can always be given the hard sound although circum­stances may dictate a soft sound; Germans often give it a “ts” sound. In pronouncing a name based on a person’s surname, try not to change its sound. Núttallii should be accented on the first, not the second syllable. In words based on two stems, such as Oxytropis, try to keep the sound of the stems intact: Oxy-trópis, rather than accentuating an unimportant vowel—Ox-ý-tro-pis; Cardio-phýlla rather than car-di-ó-phylla. Purist Latin as spoken by Americans and British is not necessarily pure Latin anyway, and it is not easily understood by others, even our own people. Try pronounc­ing a list of names and have a group of intelligent people copy them from your oral presentation and you will see what we mean. Pronouncing according to the simple rules given above will at least make our words intelligible to foreigners, and quite possibly more so to our students who were not brought up on Latin.

Common Names Common or vernacular names are of relatively little consequence in our region. Such names arise along with a culture that lives on the land and utilizes plants for food, clothing and medicine. In New England, common names were coined for this reason; in fact, many common names can be traced back to England, where many of the same plants occurred. In the American

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West, a civilization descended full-blown upon the region and common names were often transferred incorrectly to plants that were unknown in the East. This has made for continued confusion, especially now that we have visitors to Colorado who are accustomed to the traditional common names and find them misapplied here, or visitors from Eurasia who know many of our species by their botanical names, and to whom our common names mean nothing. Common names were first employed because before Linnaeus there were no scientific names available to the unlearned farmers and townspeople. Scientific names were contained in rare herbals in cloistered libraries, and they consisted of polynomial descriptions. For a delightful discussion of the common names used in Renaissance England, see Coles (1656). The purpose of our book is not to provide common names for those who are unwilling to learn the correct ones. Our translations of generic names and specific epithets are intended to help you to relate these words and their stems to your own language, and thus to make scientific names mean more. About fifty or sixty years ago a publication called Standardized Plant Names attempted to bring order to the chaos of so-called common names. Just as is happening now, people were inventing names for plants that had none, and there were lots of plants going under different names in different places, just as in America. That book tried to produce a parallel, nonscientific nomenclature to be used by­nonprofession­als. M. L. Fernald, Harvard’s Asa Gray Professor of Botany, attacked this anti-intellectual exercise. In 1942, regarding his discovery of a certain Elephantopus in Virginia, he wrote, in a footnote in Rhodora, page 367: “We thus added another to the Virginian series of elephants and their feet. We already had the Barefooted Elephant (E. nudatus), the Hairy-toed Elephant (E. tomentosus), the Stub-toed Hairytoed Elephant (E. t. forma rotundatus), and the Carolina Elephant’s-foot! We were adding the Woolly-socked Carolina Elephant’s-foot! These names, like Foul-scented Love-grass and others in Britton and Brown, and many of the crudely formed absurdities in the new Standardized Plant Names, are not colloquially used. Ours are intended as jokes; the others, unfortunately, were not. It is often said, however, that the greatest jokes are unintentional.” There is a renewed desire for common names in this otherwise advanced scientific age of biodiversity, and it is spreading like a virus through all groups of plants, including lichens and mosses. Botanical “fan clubs” and government agencies seem to feel that their workers and constituents must have a common name for every species of plant. If there is none available, one must be coined. This is an insult to the intelligence of children and adults. We should ask ourselves whether this is necessary for the progress of botany and, ultimately, for our success in saving and managing our lands. Lloyd Shinners made the most stinging remarks on common names after a lifetime of work, in his Spring Flora of the Dallas–Fort Worth Area: One of the most tiresome and irritating remarks we have to listen to over and over, runs: “Oh, don’t give me those terrible Latin names; give me something I can say and understand.” People who mouth such jawbreakers as chrysanthemum or asparagus without batting an eye are simply being childish when they say they cannot manage Latin names. That is what those two words are, without a single letter changed . . . For the Standardized Plant Names, manufac­tured according to arbitrary rules by bureaucrats, largely by translat­ ing the Latin binomials with varying degrees of accuracy, we have no use whatever. They are an insult to intelligence and a crime against good taste. There is absolutely no necessity for concocting fraudulent “common names” for plants which the “com­mon man” often cannot tell apart in the first place. Anyone with serious enough interest to want to distinguish species and varieties to the same degree that a botanist does certainly ought to be serious enough to use botanical names. Those who refuse to accept the disciplines of Science are not entitled to its benefits. The botanist has enough hard work to do without being asked to put up with a lot of artificial gobbledy-gook in the form of bogus vernacular names. We have no patience with the you-do-all-the-work, give-me-something-for-nothing attitude which lies behind demands for “common” names. Genuine

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popular names are often vivid and interesting. A study of them would be fascinating, but it belongs in the realm of folk-lore, not of science.

My experience has been that children have trouble neither with scientific names, especially if they are told the meaning, nor with technical keys. In fact, their understanding of the English language is enhanced by seeing the similarities and derivations of scientific and vernacular words, and their powers of observation are challenged by key choices. There are lots of so-called wildflower guides that require no linguistic or scientific aptitude on the part of the reader; perhaps reluctant adults should be satisfied with those. There is a tendency nowadays, stemming partly from the perceived need in the federal agencies—National Park Service, Forest Service, Bureau of Land Manage­ment, Soil Conservation Service—and local native plant societies and municipalities, to make inventories and databases of floras. These inventories are expected to discover the distributions of species of special concern (rare, threatened, and endangered). This is a laudable pursuit. However, an unfortunate consequence is that there seems to be a consen­sus of opinion that all species must have a common name. Thus, if a plant has no common name, give it one. This practice is resulting in a proliferation of different common names for the same plant, and some really ridiculous tri- and quadrinomial names. Here are some horrible examples of the lengths that this practice can go. A book on lichens (G. G. Niering’s The Lichen Book, 1947) attempted to be user-friendly by translating the scientific names. The following examples emerged: leper lichen, herd lichen, bitter knob lichen, crumbling stud lichen, pitted blind lichen, blind clay lichen, and Weigel’s leather lichen. In a book on Mosses of Singapore and Malaysia, translation of the scientific names resulted in family names: ruined tooth mosses, horizontal tooth mosses, pouched hood mosses, pott mosses, signal leaf mosses (from Sematophyllaceae—semaphore?), and species names: Sullivant’s outer net white moss, white-sheathed horizontal tooth moss, spoon-shaped two-row moss, obliquely inserted folded fruit moss, dubious bladder moss, (from Vesicularia dubyana, named for Duby, a man!), uncovered nipple moss, and hairy nipple moss. In the Colorado flora, Saussurea weberi (the genus named for a Swiss named Saussure) became Weber’s Saw-wort! Many common names repeat the scientific genus name and are really superflu­ous. Parnassia is a fine name, easy to remember. Why call it grass-of-Parnassus, when it has nothing to do with a grass? Calypso is another, with mythological interest. Why call it fairy slipper? If it looks like a slipper, it is more like a toe-dancing shoe. Unless a given species in a genus has a time-hallowed name that simply must be used, why not give all the species of the genus a “common” name, that of the genus: Eriogonum, Stipa, Geum, and Melica. Over the years, we have tried to eliminate common names embodying the name “fairy,” “baby,” and other diminutives like them. These common names were fostered during the days when botany was considered a subject for “nice ladies.” Women were, in fact, lured into university botany courses because those were less messy than zoological ones, and one did not risk soiling one’s clothes. This practice has been demeaning to the science of botany and insulting to women. It is high time we called plants by their right names, none of which are more difficult to mouth than carburetor, ibupro­fen, or lobotomy. Here are a few points to ponder: 1. Botany is a science. Despite the fact that plants are responsible for the survival of all animal life, plant products are basic to medicine, and in many parts of the world plants are crucial to generation of food and housing, our science lacks respect. Botanists are commonly considered to be lovers of wildflowers, little more. The reluctance of editors of magazines and the press to use scientific names for plants is a symptom of this. The magazine Natural History recently published a special issue on dinosaurs. All of the jaw-breaking scientific names are there, but few if any common names. Sandwiched between dinosaur articles, a short article highlights the rare vascular plants of Colorado’s High Creek Fen. Not a single scientific

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2.

3.

4.

5.

name! The editor replied to my protest to the effect that the magazine is designed for average readers. But all children quickly learn the scientific names of dinosaurs! An issue of Smithsonian Magazine featured on its cover a gorgeous color illustration of a sunflower, Scabrethia (Wyethia) scabra, growing on the sand dunes of eastern Utah. The inside cover identified this (no scientific name whatever) as sandpaper mules ears! Mules ears is a name given to the genus Wyethia, but it applies only to those species with big basal leaves; the figured species has no basal leaves. We might at least expect the scientific names to be provided in parentheses. Scientific names make it possible for everyone, anywhere in the world, to communicate with each other about organisms. Latin was chosen because it is a language that no longer is subject to change and can be used universally. On a botanical excursion to Japan, our guides often did not know the scientific names of plants, but had long-standing common names. When I asked the name of a particular plant, I pointed to it. My guide looked up the Japanese common name in the handbook, found the page, and pointed to the name and description. Lo, the scientific name was there, and everybody was happy. Scientific names are no harder to pronounce than contrived vernacular names. The public is not scientifically illiterate, nor should be encouraged to be so. Children have no trouble with scientific names. Scientific names have meaning and relevance to our own language, especially considering the great contribution of Latin and Greek to our common words. Hence they have educational value. A great proportion of scientific names have stems that are found in everyday English words. But botany is almost alone in its habit of wholesale manufacturing of vernacular names to appease what we seem to perceive as an illiterate public. Sooner or later, we will find that there is more than one vernacular name for a species. A national committee will then be set up to delete all of the alternative names and select one as a standard. An “official” list of common names has already been published for Swedish lichens!

Instead of succumbing to this new craze for common names, so dear to magazine and news editors, television, and federal agencies, let botanists make a real effort to educate the public. Give them credit for some intelligence, as other scientific disciplines do.

How to Collect and Preserve Botanical Specimens There is nothing new or strange about plant collecting. The first instructions we have seen were published about 1695 by James Petiver in a work called Gazophylacii Naturae et Artis: “Fourthly, in collecting of Plants, pray observe to get that part of either Tree or Herb as hath its Flower, Seed, or Fruit on it, but if none, then gather it as it is, and if the Leaves which grow near the root of any Herb, differ from those above, be pleased to get both to compleat the Specimen, these must be put into a Book or into a Quire of Brown Paper (which you must take with you) as soon as gathered, and once a Week shift them to a fresh place to prevent either rotting themselves or Paper.” An herbarium, or collection of dried and pressed plants, is very useful as an aid to remembering the plants you have already named, and as a reference whereby you can check your “unknowns” against the plants you already know. A collection of carefully named and preserved plants is better for these purposes than the best description or picture you might find. Collecting specimens for souvenirs is very different from collecting them professionally. In the first instance all one needs is an old book that you don’t mind ruining (please don’t use this one), a Sears-Roebuck catalog, a telephone directory, or just newspapers. You can make a

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small press, using plywood, and cut corrugated cardboard for ventilation as the plants are drying. You can put these in manila folders (but don’t use Scotch tape). Our recommendation for making a file of them is simply to place the dried specimens in a folded sheet of paper and pin the label on the right-hand corner. We strongly recommend against mounting the specimens if you ever plan to give your collection to a school or other institution. It is extremely expensive, and in time you may wind up with a white elephant on your hands. If you ever decide to give the collection to an herbarium, its staff will prefer to receive them unmounted because each institution has its own ideas of proper mounting and supplies. For most casual collectors only snippets of specimens are needed as reminders, and these can be kept in notebooks or in the way suggested above. It is important to make notes on features that would not be obvious from the specimen: height, flower color, branching mode, nature of underground parts (rhizomes, bulbs, fibrous roots, massive underground roots, etc.). Color photographs, including one of the entire plant, and a closeup of the flower, are very useful. With plants of special environmental concern, like orchids, good photographs can be used for the record, even in an herbarium. If you intend to collect specimens ultimately destined for an herbarium, or if you plan to send specimens to an herbarium for identifica­tion, you will want to operate on a higher level of sophistication. An excellent guide for this is D.B.O. Savile’s, Collection and Care of Botanical Specimens, available from Queens Printer and Controller of Stationery, Ottawa, Canada. Here are a few things that a serious plant collector should have: 1. Some sort of digging tool (a trowel, prospector’s pick, hunting knife, or weed digger) and a good sharp penknife for cutting twigs. 2. A stock of old newspapers; tabloids are best, but full-size papers may be torn along the main fold so as to make single sheets folded once into 12 × 18 inch (30 × 40 cm) folders. For the more affluent, blank newsprint is more elegant and can be written on more effectively. 3. At least 100 felt blotters, 12 × 18 inches, and twice as many corrugated cardboard ventilators, 12 × 18 inches. 4. A plant press (two flat frames of wood, either solid or a lattice of slats, 12 × 18 inches). 5. Two lengths of rope, parachute cord, or trunk straps to tie the press together. 6. A field press (12 × 18 inch satchel of lightweight waterproof fabric) for pressing along the trail. We can’t overemphasize the importance of this. Do not use plastic bags; they sweat, flowers come off, the stems bend into odd shapes, species get mixed together, and putting the specimens in the plant press at the end of the day is wearisome and disappointing in terms of quality. Specimens pressed immediately in a field press can be kept for several days before being transferred to the standard press. 7. A small field notebook for recording collection data, flower colors, plant height, and character of the underground parts if not collected. The quality of specimens varies with the good sense of the collector. The specimen is cut, pulled, or dug, and placed in the newsprint folder, with care used to clean off adhering soil and to arrange the specimen neatly. If the specimen does not have sufficient flowers or fruits, collect some extra and press them separately. Plants that are too large to fit in the folder should be folded. This especially applies to grasses. Here, simple folding is not enough (it can still produce hay; Aven Nelson once commented, on receiving a mess of folded grasses, “probably was good hay, too!”); at each fold, slip a piece of light card stock that has been slit in the middle, over the fold to hold it in place. Such specimens won’t get out of hand later on. Remember that a standard sheet of herbarium paper is 12 × 16 inches, and that a label will

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occupy about 3 × 5 inches at the lower right-hand corner. Your specimen should fit without lopping over the edges of the paper. If you anticipate having to examine flowers or fruits in detail after they are dry, be sure that you press enough extra of them that this examina­tion will not destroy the only flowers on a specimen. Take notes on flower color, height of plant, nature of the root system, and anything else that you leave in the field but might be needed later. The quality of your specimens will depend on care of preparation for the press: selecting the specimen that best exemplifies the population, cleaning off any adhering mud or soil, and collecting enough material. You can be as fastidious as you like in arranging the leaves and spreading the petals, but this must be done at the moment you first place the plant in the folder; afterwards it will be too late. Beyond this, ample pressure in the press is needed, as well as frequent changing of the blotters to carry off the accumulated moisture. When the leaves crack when manipulated, and when the stem feels dry, the specimen is ready to remove from the press. A field notebook is essential. Number your specimens. We suggest that a simple consecutive number system be used. (We abhor systems that combine year, month, day, and specimen number!) Numbers are cheap; they can be used or deleted, and your ego will swell after your numbered list gets to a thousand or more. Place the number in big, bold black characters in the middle of the newspaper sheet (edges of sheets tend to get gnawed by use or rodents). For each collecting foray, start a new heading: state, county, locality, altitude (rounded out to the nearest 500 feet or 100 meters is good enough), and date. Follow this with your numbers; ecological situation and supplemen­tal information can be added after the number. Sticking to a standard routine will make it easier to produce labels, either by hand or on a computer program. It seems that there are as many formats of specimen labels as there are curators. Some labels seem to trumpet ownership or advertise an herbarium (HERBARIUM OF Y. Y. FLERTCH, PEORIA, ILLINOIS, for instance, spread in block letters across the top), rather than give adequate specimen information. We have tried, by precept and example, to minimize the advertising on specimen labels. The identification of the herbarium can be relegated to an unobtrusive line at the bottom. One important feature we would like to see on more labels is the family name or its abbreviation. Labels can be printed in two columns on standard 8½ × 11 inch, acid-free 100 percent rag paper (other stocks are unstable over time). We prefer to use unperforated stock because some labels will be more detailed than others; a paper cutter does not add too much time. A sample label is given here: COLORADO, U.S.A. Penstemon versicolor Pennell

SCR

Las Animas Co.: 4 mi N and 1/8 mi E of Andrix. T30S R51W S24, ca. 1300 msm. Restricted to caliche outcropping. Flowers bright blue-purple, basal leaves present at anthesis, stems ascending. 22 May 1993

Weber & Wittmann 18546 Herbarium COLO (Boulder)

The Importance of Voucher Specimens People come to the herbarium and expect to find the complete record. This is an impossibility. Many years ago I attempted to determine just how good our coverage was. I selected the species of one family, the Liliaceae, which contains mostly easily seen species whose distributions, from experience, I could fairly well predict. I prepared a table of the counties of

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Colorado and the species that would be expected to occur in each county. I found that the herbarium contained only 19 percent of those to be expected. Of course the herbarium at the time contained only about 30,000 specimens. However, fifty years later the percentage is not much higher, for collectors have not been able to mount expeditions into each county on a scientific basis, and we tend to revisit those few areas that were especially fruitful for our efforts. Therefore, amateurs can make an invaluable contribution by collecting in the counties that are not well covered: make good specimens, and deposit them (vouchers) in an institutional herbarium. Our herbarium has available hand checklists for each county that list those species that have been deposited in the past, so that only those specimens really needed might be watched for and collected.

Collecting Ethics Because of the current emphasis on conservation, many potential collectors of plant specimens feel that any collecting should be discouraged. This is certainly not the case. Woody plants, for example, can have representative branches in leaf, flower, or fruit collected without any serious damage to the plant. Annual species are usually abundant enough to permit judicious collecting; they also usually produce an abun­dance of seeds. Perennials will not suffer to have one or two plants taken. Of course, there are rare instances when there are not enough specimens to allow the taking of many, but it is a rare species indeed that cannot suffer the loss of one or two speci­mens. If the proper habitat is available, there are probably many other stands that were not seen. If the habitat is extremely localized, there may be good reason to be especial­ly careful; in fact, some of our best publicized rare plant sites are pillaged by large numbers of botanists and gardeners who still have the acquisitive habits of the vanity collector. However, in general, deer, elk, and other animal foragers, fungus infestation, bulldozers,and clearing for home-building and lawns is much more destructive to wild popula­ tions than the collections of a few botanists. In the last analysis, it is extremely important to have the record of where the plants grew, how many there were, and the nature of the habitat preserved as speci­mens in the herbarium. Inevitably, some of the populations are going to be destroyed no matter how conscientious we ourselves may be in our fieldwork, but if vouchers are preserved at least we will have them as long as our civilization preserves the herbaria and museums on which our cultures are based.

Plant Identification No single book is enough to help you learn the art of analysis and identifica­tion of plants. For beginners, some elementary book in plant analysis is essential. There are many and they go back over a hundred years. Asa Gray’s Lessons in Botany, written in the 1850s, is still extremely useful; editions of it still can be found in used book stores. For general vocabulary and descriptions and illustrations of families, V. H. Heywood (ed.), Flowering Plants of the World (1978) is ideal. H. D. Harrington & L. W. Durrell’s How to Identify Plants (1957), P. H. Davis & J. Cullen’s The Iden­tification of Flowering Plant Families (1979), J. P. Baumgardt’s How to Identify Flowering Plant Families (1982), J. P. Smith’s Vascular Plant Fami­lies (1977), and S. B. Jones & A. E. Luchsinger’s Plant Systematics (1986) are all useful. The two biggest hurdles in becoming adept at plant iden­tification are mastering the terminology and learning to recognize the bigger plant families so as to avoid having to use the family key. Family keys are extremely difficult to write because there are so many exceptions to the stereotype. Small families are usually much easier to learn because they are more cleanly circumscribed.

How to Use the Keys The dichotomous key has become the standard method for the identifica­tion of organisms and should need little introduction. It involves the presentation of successive pairs of choices,

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from which the reader arrives at a decision and then proceeds to the next choice (as indicated by the paren­thetical number on the right) and the next, until arriving at the name of the organism. Keys are simply means by which one arrives at a tentative decision, which should be followed up by a careful reading of a detailed description and examination of authentically named specimens and illustrations. This is why we have herbaria for reference and great multivolume editions of regional and state floras. In a field guide (the so-called Excursion-flora of the European), the need for conciseness makes an extensive treatment impossible. As a compromise, our keys are somewhat more detailed, and notes are added concerning the habitat, altitudinal range, and special characteristics that help to confirm the final decision. There are several kinds of keys. The first is a key that is relatively infallible because it uses all the critical features of the plants regardless of whether they are easily visible or require elaborate techniques of dissection. It might require observation of features, such as flower buds, seeds, or winter buds that are not always present. It also tends to place related species or genera close to each other. This kind of key is written for the person who makes it rather than for the user in the field. The second kind of key is designed for the user, and its intention is to make possible iden­tification of plants under almost any field condi­tions. This is an impossible task, at best, but is a goal that writers of field handbooks aim for. Such a key may also deliberately make mistakes, that is, it will assume the reader’s observation may be incorrect but will bring him to the correct con­clusion anyway. Key-writing is a fine art, and the use of keys has to be developed in somewhat the same spirit. One key-writer of note has pointed out that “if the presence of various small features is useful as a means of identifying some kinds of plants, then the absence of the same features must be equally useful in distinguishing other kinds. Persons using a key for identification seldom have any difficulty in recognizing the presence of a structural feature but often find it difficult to convince themselves of its absence. This is purely a matter of mental attitude and has nothing to do with the size and conspicuousness of the feature in question. Those who use this or any similar work should guard against this tendency” (Gleason, New Britton and Brown Illustrated Flora, 1952). The limitations of the author’s experience and the fact that ecological compensation places plants of higher altitudes lower down on a north-facing slope than plants of lower altitudes should be a warning to readers that alpine species might be expected occasionally to occur at lower altitudes and vice versa. In such matters the reader is requested to bear with the author, for a definite statement subject to modification is often prefer­able to silence. Plants with blue flowers will throw white-flowered mutants. Low plants growing on a well-manured site may assume giant proportions. And the beginner will always find a flower with six petals when there should be only five. One of my old bird-watching mentors always used to mumble under his breath, “Birds don’t read books.” Neither do plants.

Eponymy: Botanists Honored in Colorado Plant Names To identify all the people who have been honored in the names of Rocky Mountain plants would fill this book. Most of them are mentioned briefly in the text. The ones listed here are those who, again and again, appear in the specific epithets or in generic names. Most of these were collectors in Colorado or worked intensively on Rocky Mountain plants. For additional names and details, see Joseph and Nesta Dunn Ewan (1981) Biographical Dictionary of Rocky Mountain Naturalists. The best index to biographical and bibliographic information on botanists will be found in the three-volume reference work Bibliographic Notes on Botanists, produced from a card file accumulated over many years by the late George Hendley Barn­hart, of the New York Botanical Garden. This man entered briefly into my career, for in the first half of the first decade of my life Mr. Barn­hart lived in the apartment above ours on Valentine Avenue in the North Bronx, and my mother often invited him down for tea by knocking on the ceiling with her broomstick.

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Baker, Charles F., 1872–1927, collected extensively in Montrose and Gunnison counties. His collec­tions were described by Greene in a series of papers called Plantae Bakerianae. Barneby, Rupert C., 1911–2000, indefatigable collector and monographer. Preeminent specialist in Fabaceae, notably for Colorado, Astragalus and Oxytropis. Bigelow, John Milton, 1804–1878, botanist, surgeon on the Mexican Boundary Survey, 1848– 1854. Brandegee, Townshend Stith, 1843–1925, botanist with the Hayden Surveys, also surveyor of Fremont County, made important collections in the Mesa Verde area. Clements, Frederick E., 1874–1945, ecologist, originator of the plant succession concept, had experimental gardens on Pikes Peak. Constance, Lincoln, 1909–2001, American specialist in Apiaceae, University of California at Berkeley. Coulter, John Merle, 1851–1928, botanist with the Hayden Surveys and author of Manual of the Botany of the Rocky Mountain Region (1885). Crandall, Charles Spencer, 1852–1929, horticul­turist, professor at Colorado Agricultural College, collected extensively in Colorado. Douglas, David, 1799–1834, Scottish botanist and explorer of the West, never visited Colorado but described and collected many plants native to the state. Eastwood, Alice, 1859–1953, high school teacher in Denver, collected around Grand Junction in 1890, later became curator at the California Academy of Sciences where she gained lasting fame by segregating the type specimens and eventually saving them during the great earth­quake. Engelmann, George, 1809–1884, St. Louis physician-botanist and a founder of the Missouri Botani­cal Garden. Major contributions were made in the cacti and conifers. Fendler, Augustus, 1813–1883, collector for Engel­mann and Gray. Frémont, John Charles, 1813–1890, explored widely in the West and collected plants and animals. Unfortunately, many of his specimens were lost in accidents along the way. Gray, Asa, 1810–1888, student of Torrey, develo­per of the Harvard Herbarium, the dominant taxonomist of his era. Visited Colorado twice, in 1872 for the dedication of Gray’s and Torrey’s Peaks, and for a few days in 1877, with Sir Joseph Dalton Hooker. Greene, Edward Lee, 1843–1915, field botanist and clergyman in Colorado and New Mexico, later botanist at the University of California, Berkeley, one of the most knowledgeable persons of his time about the Colorado flora. Hall, Elihu, 1820 (or 1822?)–1882, Illinois botanist, collected in South Park with J. P. Harbour and C. C. Parry in 1862. Harbour, J. P., dates unknown, collected in South Park in 1862 with Elihu Hall and C. C. Parry. Harrington, Harold D., 1903–1981, professor at Colorado State University, Fort Collins, and author of Manual of the Plants of Colorado (1954), the only complete flora of the state with keys and descriptions, indispensable to present and future floristic work. Hayden, Ferdinand Vandeveer, 1829–1887, leader of the U.S. Geological and Geographical Survey of the Territories, 1867–1879, in which many botanists participated. Holm, Herman Theodor, 1854–1932, Danish botanist, collected in Colorado in 1896 and 1899, and pub­lished a classic paper on the plant geography of the Rocky Mountains.

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Hooker, Sir Joseph Dalton, 1817–1911, director of the Royal Botanic Gardens, Kew, an early leader in plant geography, visited Colorado with Asa Gray in 1877 and was the first to note the strong Asiatic element in our flora. Hooker, Sir William Jackson, 1785–1865, British botanist, described many western North American plants from the historic voyages of exploration and from Geyer’s trip across Wyoming; director of the Royal Botanic Gardens, Kew. Most species commemorating Hooker in our flora refer to W. J. rather than to his son, Sir Joseph Dalton Hooker. James, Edwin, 1797–1861, surgeon-naturalist with the Long Expedition, collected plants in 1820 from Pikes Peak to the Platte River. See G. J. Goodman and C. A. Lawson (1995), Retracing Major Stephen H. Long’s 1820 Expedition. Jones, Marcus Eugene, 1852–1934, exceptional field botanist, probably the greatest collector the West has known, mining engineer, contemporary and great competitor of Greene. Letterman, George Washington, 1841–1913, a reclusive Missouri school­teacher, collected on Long’s Peak. Löve, Áskell, 1916–1994, Icelandic cytotaxonomist, professor at the University of Colorado, and prolific botanical writer: Flora of Iceland, Cytotaxono­mical Atlas of the Arctic Flora. Nelson, Aven, 1859–1952, Wyoming botanist and founder of the Rocky Mountain Herbarium at Laramie, first lifelong resident botanist of this region. Great teacher and developer of botanists, such as L. N. Goodding, Elias Nelson, J. F. Macbride, Marion Ownbey, and George J. Goodman. Nuttall, Thomas, 1786–1859, English botanist, explorer of the West with the second Wyeth Expedition, Harvard professor, and a perceptive taxonomist, especially in Asteraceae. Osterhout, George E., 1858–1937, amateur naturalist and resident collector in Colorado and southern Wyoming. Parry, Charles Christopher, 1823–1890, physician-botanist of Davenport, Iowa, collected exten­sively on the Eastern Slope. See W. A. Weber (1997), King of Colorado Botany: Charles Christopher Parry, 1823–1890. Patterson, Harry Norton, 1853–1919, botanist-printer of Oquawka, Illinois, collected in the Gray’s Peak area. Payson, Edwin Blake, 1893–1927, promising student of Cockerell and Nelson, collected extensively from his father’s cattle ranch at Naturita. A brilliant botanist, specialized in Brassicaceae and Boraginaceae. Penland, C. William T., 1899–1982, professor at Colorado College, avid alpine botanist, dis­ covered many rare alpine species on Hoosier Pass, specialized in Penstemon. Pennell, Francis W., 1886–1952, Pennsylvania botan­ist, specialized in the Scrophu­lariaceae of the Rocky Mountain region. Porter, Thomas C., 1822–1901, professor at Lafayette College, Pennsylvania, collected with the Hayden Survey and published the first Colorado flora in 1874, Synopsis of the Flora of Colorado. Pursh, Frederick, 1774–1820, German botanist, author of Flora Americae Septentrio­nalis, which described the collections of the Lewis and Clark Expedition. Rothrock, J. T., 1839–1922, student of Asa Gray, botanist with the Wheeler Expeditions, 1869–1879. Rydberg, Per Axel, 1860–1931, Swedish immigrant and curator at the New York Botanical Garden; with Greene and Nelson, one of the most important figures in Rocky Mountain botany, published Flora of the Rocky Mountains and Adjacent Plains, 1917 [1923], and collected extensively in Colorado.

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Schmoll, Hazel M., 1891–1990, University of Chicago student, wrote thesis on Chimney Rock area, Pagosa-Piedra region, curator of botany at the Colorado State Museum, 1919–1923, and collected at Mesa Verde National Park; mayor and “majordomo” of the town of Ward for many years, after giving up botany completely. Vasey, George, 1822–1893, curator in the U.S. National Herbarium, collected extensively in eastern Colorado. Watson, Sereno, 1826–1892, botanist on U.S. exploring expeditions, later assistant to Asa Gray at Harvard and then curator.

Notes Generic names: We have presented generic names together with their authors and dates of publication. We have also tried to present a derivation of each name for its educational value. Specific names: A specific name consists of a generic name and a specific epithet, followed by the name of the author of the name. Botanical nomenclature differs from zoological nomenclature in the way authors of the names are cited. The name of an author in parentheses means that this author first published the species. The second author is the person responsible for the name in its present form, usually under another genus. Counties: Colorado is divided into sixty-four counties as shwon on the map on the inside cover. County abbreviations are given on the facing page. Family abbreviations: Collectors should indicate on their labels the family to which a species belongs. This makes for ease in filing and retrieval from collections. Because family names are usually long, we have developed a standard three-letter abbreviation for each family (Weber 1982a). These abbreviations are given at the head of each family treatment and are used in the running heads. Family arrangement: Families are listed alphabetically within the groups: Ferns and Fern Allies, Gymnosperms, and Angiosperms. Synonymy: We have listed synonyms italicized and in parentheses. Meanings: We show the meaning of the name in square brackets. Occurrence: In the keys, genus and species are bolded in roman typeface, indicating occurrence in Colorado’s eastern slope. Genus and species in italics indicates that it may be expected to occur here, but has not yet been recorded. Please, let us know if you find these! The county abbreviations are shown for known distribution. Figures: Figures are referred in bold, e.g. 27F. Anecdotes: Explanations and accounts in first person singular refer to Bill Weber’s experiences; first person plural to Bill Weber and Ron Wittmann.

Colorado Flora E a s t e r n S lo p e

KEY TO THE FAMILIES Note: In the longer keys reference is made, in square brackets, to the number of the couplet from which you last came; page references are not given, since the families are in alphabetic order. Starting pages for major groups are as follows: Ferns and Fern Allies, p. 17 Gymnosperms, p. 38 Angiosperms, p. 42 1a. Plants not producing seeds or true flowers, but reproducing by spores; fern-like, moss-like, rush-like plants. Ferns and Fern Allies 1b. Plants producing seeds, either by means of flowers or cones; plants of various aspects (seed plants) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. Leaves needle-like or scale-like; evergreen trees and shrubs, never with flowers; ovules and seeds on the open face of a scale or bract (rarely the cone becomes a fleshy “berry” in Juniperus and Sabina). Gym­nosperms 2b. Leaves various, seldom needle-like or scale-like (if so, flowers are present), rarely evergreen; ovules and seeds borne in a closed cavity (carpel; ovary). Angiosperms, Flowering Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 3a. Parasitic or saprophytic, often highly colored but not green (mistletoe, in this category, has some chlorophyll but is yellowish and epiphy­tic). Key A 3b. Not parasitic, or at least having green leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 4a. Stems thick and succulent, spiny; true leaves absent or greatly reduced and early deciduous. Cactaceae, Cactus Family 4b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) 5a. Leaves all basal, with circular blades covered with stalked, glistening red glands; flowers in a raceme. Droseraceae, Sundew Family 5b. Not as above, not insectivorous plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6) 6a. Submerged plants, with or without floating leaves. Key B 6b. Terrestrial or semiaquatic, not submerged nor with floating leaves . . . . . . . . . . . (7) 7a. Vines, climbing or twining among other plants, often possessing suckers or tendrils, not merely creeping on the ground. Key C 7b. Herbaceous or woody plants, not vines (the Smilacaceae have tendrils) . . . . . . . . (8) 8a. Leaves usually parallel-veined; flower parts in 3s; stem hollow or with scattered vascular bundles; herbaceous (except Agavaceae and Nolinaceae); seeds with 1 cotyledon (but see also Limnanthaceae, which has floral parts in 3s, otherwise a typical dicot). Key D 8b. Leaves usually netted-veined; flower parts in 5s, 4s, or 2s; stems with vascular bundles arranged in a ring around the pith; herbaceous or woody; seeds usually with 2 cotyledons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (9) 9a. Trees or shrubs. Key E 9b. Herbaceous, sometimes woody at the very base. Key F

KEY A (Parasites/saprophytes) 1a. 1b. 2a. 2b.

Attached to the bark of trees, or by suckers to the aerial stems of herbs . . . . . . . (2) Without obvious attachments to the aerial parts of their hosts . . . . . . . . . . . . . . . (3) Attached to the trunks of branches of evergreen trees. Viscaceae, Mistletoe Family Thread-like orange or yellow plants attached by suckers to aerial parts of herbs. Cuscu­taceae, Dodder Family 1

2

Key to the Families: Parasites/saprophytes

3a. Flowers actinomorphic, in a spike-like erect or nodding raceme. Monotrop­aceae, Pinesap Family 3b. Flowers zygomorphic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 4a. Flowers tubular, the petals united; ovary superior. Orobanchaceae, Broomrape Family 4b. Flowers with separate petals; ovary inferior. Orchidaceae, Orchid Family

KEY B (Aquatics) 1a. Plants usually not more than 1 cm long, free-floating or submerged, with no attachment whatever . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 1b. Plants not free-floating, or much larger if appearing so . . . . . . . . . . . . . . . . . . . . . (3) 2a. [1] Plants disk-like or thallus-like, without true stems and leaves, free-floating or submerged. Lemnaceae, Duckweed Family 2b. Plants with obvious leaves arranged in 2 ranks along a short stem, free-floating. Salviniaceae, Waterfern Family, in Ferns and Fern Allies 3a. [1] Stems short and lacking, the leaves attached to the bottom, linear-elongate, the tips floating on the surface. Sparganiaceae, Burreed Family 3b. Plants with definite stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 4a. [3] Leaves simple, entire or slightly toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) 4b. Leaves distinctly lobed, compound, or finely dissected . . . . . . . . . . . . . . . . . . . . (17) 5a. [4] Leaves linear or oblong, arranged in whorls . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6) 5b. Leaves variously shaped, not whorled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (7) 6a. [5] Leaves translucent, lax, 2 cell-layers thick; flowers, if present, sessile (carpellate) or long-pedicelled (staminate). Hydrocharitaceae, Frogbit Family 6b. Leaves opaque, rather rigid unless submerged, more than 2 cell-layers thick; flowers sessile in the leaf axils. Hippuridaceae, Mare’s Tail Family 7a. [5] Leaves almost orbicular, deeply cordate, very thick and leathery; flowers large, yellow, solitary. Nymphaeaceae, Pondlily Family 7b. Leaves narrower, not cordate; flowers not as above . . . . . . . . . . . . . . . . . . . . . . . . (8) 8a. [7] Flowers blue or yellow, with 6 tepals and 3 stamens. Pontederi­aceae, Pickerelweed Family 8b. Flowers greenish or colored, but not blue or yellow . . . . . . . . . . . . . . . . . . . . . . . . (9) 9a. [8] Leaves linear or filiform . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (10) 9b. Leaves with distinctly broadened blades . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (14) 10a. [9] Flowers in spikes. Potamogetonaceae, Pondweed Family 10b. Flowers sessile in the leaf axils or on slender, often coiled pedun­cles . . . . . . . . . (11) 11a. [10] Fruit minute, blackish, on an elongate, often coiled peduncle; leaves filiform, over 3 cm long. Ruppiaceae, Ditchgrass Family 11b. Flowers and fruits sessile in the leaf axils; leaves shorter . . . . . . . . . . . . . . . . . . . (12) 12a. [11] Fruit rounded or emarginate, oblong or wider, not beaked. Callitrichaceae, Water Starwort Family 12b. Fruit narrowly cylindric, tapered to a beak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (13) 13a. [12] Fruit flattened, slightly curved, with a stout beak; leaves filiform. Zannichelli­ aceae, Horned Pondweed Family 13b. Fruit terete, straight, the beak whitish, not rigid; leaves linear, flat, the margins very finely toothed (under high magnification). Najad­aceae, Waternymph Family

Key to the Families: Aquatics

3

14a. [9] Leaves alternate, at least 1 cm long; flowers not sessile in the leaf axils . . . . . (15) 14b. Leaves opposite, less than 1 cm long; flowers inconspicuous, sessile in the leaf axils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (16) 15a. [14] Floating leaves pinnately veined; flowers pink, with showy pink perianth parts. Persicaria in Polygonaceae, Buckwheat Family 15b. Floating leaves with parallel veins, or floating leaves absent; flowers greenish, not showy. Potamogetonaceae, Pondweed Family 16a. [14] Stipules lacking; calyx and corolla absent; ovary 4-locular; floating and submerged leaves often strikingly different. Callitrichaceae, Water Starwort Family 16b. Stipules present; calyx and corolla often present; ovary 3- or 5-locular; leaves not dimorphic. Elatinaceae, Waterwort Family 17a. [4] Leaves 3-foliolate; flowers white, petals fringed. Menyanthaceae, Buckbean Family 17b. Leaves not 3-foliolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (18) 18a. [17] Leaves bearing small balloon-like traps; flowers showy, yellow, spurred, on racemes projecting above water level. Lentibulariaceae, Bladderwort Family 18b. Leaves not bearing bladders; flowers not spurred . . . . . . . . . . . . . . . . . . . . . . . . . (19) 19a. [18] Leaves alternate; flowers with white or yellow petals. Ranun­culaceae, Buttercup Family 19b. Leaves whorled; flowers greenish, inconspicuous . . . . . . . . . . . . . . . . . . . . . . . . . (20) 20a. [19] Leaf divisions dichotomous, finely serrate; flowers sessile in the axils of normal leaves. Ceratophyllaceae, Hornwort Family 20b. Leaf divisions pinnate, entire; flowers in an interrupted spike resembling a knotted cord. Haloragaceae, Water Milfoil Family

KEY C (Vines) 1a. 1b. 2a. 2b. 3a. 3b. 4a. 4b. 5a. 5b. 6a. 6b. 7a. 7b. 8a. 8b. 9a. 9b.

Leaves simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) Leaves compound (Caution! Poison-ivy is in this category) . . . . . . . . . . . . . . . . . . (7) Leaves palmately lobed, sometimes only slightly so . . . . . . . . . . . . . . . . . . . . . . . . (3) Leaves not lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) Plants with tendrils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) Tendrils absent. Humulus in Cannabaceae, Hops Family Herbaceous; fruit a papery, spiny balloon or a gourd. Cucurbitaceae, Cucumber Family Woody; fruit a fleshy “grape.” Vitaceae, Grape Family Venation parallel; flowers and fruits in umbels. Smilacaceae, Smilax Family Venation netted; flowers and fruits not umbellate . . . . . . . . . . . . . . . . . . . . . . . . . (6) Flowers 1 cm long or more; petals united, pleated. Convolvulaceae, Morning­-glory Family Flowers smaller; perianth parts (tepals) separate. Fallopia in Polygonaceae, Buck­ wheat Family Leaves pinnately compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (8) Leaves trifoliolate, palmately 5–7-foliolate, or ternately compound . . . . . . . . . . . (9) Leaflets entire; flowers with banner, wings, and keel (sweetpea type). Fabaceae, Pea Family Leaflets serrate; flowers tubular. Bignoniaceae, Catalpa Family Leaves palmately 5–7-foliolate. Vitaceae, Grape Family Leaves not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (10)

4

Key to the Families: Vines

10a. Leaves with 3 shiny leaflets; flowers greenish; plant short (scarcely viny in this area), commonly bearing clusters of greenish-white berries. Toxicoden­dron, poison-ivy, in the Anacardiaceae, Sumac Family 10b. Leaves twice ternately compound or, if 3-foliolate, the flowers blue or yellow, with long feathery styles in fruit. Ranunculaceae, Buttercup Family

KEY D (Monocots) 1a. Woody plants with narrow evergreen leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 1b. Herbs; leaves otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 2a. [1] Leaves stiff and erect; flowers perfect, more than 3 cm long, pendent, white. Agavaceae, Agave Family 2b. Leaves supple, long, and arching; flowers unisexual, less than 1 cm long, not pendent, cream-colored. Nolinaceae, Sotol Family 3a. [1] Tall, fern-like plants, the true leaves minute, triangular, papery, subtending clusters of filiform green cladodes; flowers small, yellowish; fruit a red berry. Asparagaceae, Asparagus Family 3b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 4a. [3] Leaves cordate-ovate; stem with stipular tendrils. Smilacaceae, Smilax Family 4b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) 5a. [4] Cattail-like plants with a tight spike of minute flowers protruding from the side of the stem; foliage very fragrant when fresh. Acor­aceae, Sweetflag Family 5b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6) 6a. [5] Flowers minute, enclosed in chaffy bracts; 3- or 6-parted perianth lacking; flowers arranged in spikes or spikelets (grasses and sedges) . . . . . . . . . . . . . . . . . . . . . . . . (7) 6b. Flowers not enclosed in chaffy bracts or scales; perianth usually present, with 3 or 6 parts that may themselves appear papery or chaffy . . . . . . . . . . . . . . . . . . . . . . . . (8) 7a. [6] Leaves in 2 rows on the stem, the sheaths usually open, the margins not fused (few exceptions); stems cylindric or flattened and almost always hollow; anthers attached to filaments at their middles. Poaceae, Grass Family 7b. Leaves 3-ranked, sometimes absent; sheaths usually closed, the margins fused; stems almost always triangular and solid (a few cylindric and hollow); anthers attached at one end. Cyperaceae, Sedge Family 8a. [6] Flowers with a rudimentary perianth of bristles or scales, or none . . . . . . . . . (9) 8b. Flowers with sepals and petals (sometimes the two are similar in shape and texture: tepals) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (10) 9a. [8] Flowers in elongate terminal spikes, the looser staminate flowers in a separate group above the dense brown carpellate ones. Typhaceae, Cattail Family 9b. Flowers in spherical clusters, staminate ones above the carpellate ones. Spargani­aceae, Burreed Family 10a. [8] Carpels numerous (over 6), separate and distinct, in a whorl or ball. Alismata­ceae, Water-Plantain Family 10b. Carpels 3 or 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (11) 11a. [10] Ovary wholly inferior, the floral parts attached to the top of the ovary . . . (12) 11b. Ovary superior or only partly inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (15) 12a. [11] Flowers radially symmetrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (13) 12b. Flowers bilaterally symmetrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (14) 13a. [12] Flowers blue; leaves gladiate. Iridaceae, Iris Family 13b. Flowers yellow; leaves grass-like. Hypoxidaceae, Yellow Stargrass Family

Key to the Families: Monocots

5

14a. [12] Flowers slipper-shaped with a rounded toe; leaves more than 1; functional stamens 2. Cypripediaceae, Lady’s slipper Family 14b. Flowers, if slipper-shaped, the toe pointed and leaf solitary; functional stamen 1. Orchidaceae, Orchid Family 15a. [11] Perianth of 6 chaffy or scale-like similar segments, hardly petal-like, but arranged in 2 alternating groups of 3; grass-like plants. Juncaceae, Rush Family 15b. Perianth segments petal- or sepal-like, not chaffy or scale-like . . . . . . . . . . . . . . . (16) 16a. [15] Tepals minute, greenish; stamens sessile; carpels separating as units at maturity; annual, or perennial from a rhizome; grass-like plants of alkaline flats and mountain fens. Juncaginaceae, Arrowgrass Family 16b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (17) 17a. [16] Outer and inner perianth segments strongly differentiated in color or size . . . (18) 17b. Outer and inner perianth segments similar (tepals) . . . . . . . . . . . . . . . . . . . . . . . (20) 18a. [17] Petals less than 2 cm long, all purple or 2 blue, 1 white. Com­melinaceae, Spiderwort Family 18b. Petals over 2 cm long, white or rose colored . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (19) 19a. [18] Leaves broadly ovate, in a whorl of 3; flowers white, with green sepals. Trilliaceae, Trillium Family 19b. Leaves linear, alternate; flowers white or rose, with a prominent gland of colored hairs. Calochortaceae, Mariposa Family 20a. [17] Flowers in umbels subtended by a group of papery bracts; stem arising from a bulb. Alliaceae, Onion Family 20b. Flowers not in umbels; stems from fibrous roots, rhizomes, or bulbs . . . . . . . . . (21) 21a. [20] Low, weak plants rooted in mud; flowers blue or yellow; stamens 3; leaves linear or narrowly oval. Pontederiaceae, Pickerelweed Family 21b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (22) 22a. [21] Inner tepals with a prominent gland at the base; ovary with prominent styles; carpels separate partway down. Melanthiaceae, False-Hellebore Family 22b. Tepals without glands, or the gland small and indistinct; ovary usually without styles (don’t confuse these with separate stigmas); carpels united . . . . . . . . . . . . . . . . . (23) 23a. [22] Leaves basal; if not, then the flowers large and showy. Liliaceae, Lily Family 23b. Leaves alternate, cauline; flowers small, 2 cm or less . . . . . . . . . . . . . . . . . . . . . . (24) 24a. [23] Inflorescence a terminal raceme or panicle; tepals white, wide spreading. Con­vallariaceae, Mayflower Family 24b. Inflorescence axillary or terminal, flowers solitary or a few; flowers bell-shaped, tepals yellowish. Uvulariaceae, Bellwort Family

KEY E (Woody dicots) (See Key C for woody vines) 1a. Leaves minute (less than 5 mm long), scale-like, overlapping, and appressed to the stem. Tamaricaceae, Tamarisk Family 1b. Leaves larger and otherwise not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. [1] Leaves covered by silvery or brownish peltate scales. Elaeagnaceae, Oleaster Family 2b. Leaves not covered by peltate scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 3a. [2] Leaves and branches opposite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 3b. Leaves and branches alternate or scattered . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (17)

6

Key to the Families: Woody Dicots

4a. [3] Fruit a samara . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) 4b. Fruit otherwise (or fruits not present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6) 5a. [4] Leaves pinnately veined and pinnately compound, with a leathery texture. Fraxinus in Oleaceae, Olive Family 5b. Leaves palmately veined, simple or compound (Negundo sometimes has 5 leaflets, thus pinnately compound); leaves not leathery. Aceraceae, Maple Family 6a. [4] Leaves palmately lobed or compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (7) 6b. Leaves neither palmately lobed nor palmately compound . . . . . . . . . . . . . . . . . . . (9) 7a. [6] Terminal bud long-pointed, not protected by overlapping scales; fruit a berry. Viburnum in Caprifoliaceae, Honeysuckle Family 7b. Terminal bud blunt or merely acute, covered by overlapping scales . . . . . . . . . . . (8) 8a. [7] Leaves pinnately veined and pinnately compound, with a leathery texture. Fraxinus in Oleaceae, Olive Family 8b. Leaves palmately veined, simple or compound (Negundo sometimes has 5 leaflets, thus pinnately compound); leaves not leathery. Aceraceae, Maple Family 9a. [6] Leaves evergreen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (10) 9b. Leaves deciduous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (12) 10a. [9] Leaves entire, paler beneath; plants of subalpine fens and pond margins. Kalmia in Ericaceae, Heath Family 10b. Leaves serrulate or crenate, not pale beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . (11) 11a. [10] Leaves elliptic; low, spreading shrubs, leaves spreading in one plane; flowers small, axillary, reddish. Celastraceae, Stafftree Family 11b. Leaves broadly oval; creeping plant with only slightly woody stems; flowers in pairs, pendent from an erect stalk. Linnaea in Caprifoli­aceae, Honeys­ uckle Family 12a. [9] Leaves linear, with smaller leaves fascicled in the axils; leaves glabrous, appearing terete, the margins tightly revolute; flowers white; restricted to gypsum soils. Fran­keniaceae, Frankenia Family 12b. Leaves broader, not in axillary fascicles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (13) 13a. [12] Leaves pinnately compound. Sambucus in Caprifoliaceae, Honeys­ uckle Family 13b. Leaves simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (14) 14a [13] Lenticels conspicuous; fruit a drupe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (15) 14b. Lenticels inconspicuous; fruit a capsule or several-seeded berry . . . . . . . . . . . . . (16) 15a. [14] Twigs red; leaves broadly ovate with raised, curved parallel venation; flowers in compound cymes. Cornaceae, Dogwood Family 15b. Twigs gray-brown; leaves narrowly to broadly oblong; venation inconspicuous; flowers greenish, in axillary clusters. Forestiera in Oleaceae, Olive Family 16a. [14] Bark exfoliating; leaves oblong or oval, commonly distinctly hairy; flowers white, waxy; fruit a dry capsule. Hydrangeaceae, Hydrangea Family 16b. Bark not exfoliating; leaves ovate; if oblong, then glabrous or very minutely hairy, sometimes slightly lobed. Caprifoliaceae, Honeys­ uckle Family 17a. [3] Leaves compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (18) 17b. Leaves simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (26) 18a. [16] Leaves spine-margined, evergreen (resembling holly); inner bark yellow. Ber­b eridaceae, Barberry Family 18b. Leaves not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (19) 19a. [18] Leaves with 3 leaflets; stems never prickly (Caution! Poison-ivy is in this category) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (20)

Key to the Families: Woody Dicots

7

19b. At least some leaves not trifoliolate, usually pinnately compound (stems may be prickly) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (21) 20a. [19] Shrub; fruit red and fuzzy or an ivory-white, hard berry. Anacar­diaceae, Sumac Family 20b. Small tree; fruit a disk-shaped samara. Ptelea, in Rutaceae, Rue Family 21a. [19] Leaves even-pinnately compound (lacking a terminal leaflet) . . . . . . . . . . . . (22) 21b. Leaves odd-pinnately compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (23) 22a. [21] Trees, never with thorns. Sapindaceae, Soapberry Family 22b. Shrubs, sometimes with thorns. Fabaceae, Pea Family 23a. [21] Fruit a legume; leaflets more than 9, entire. Fabaceae, Pea Family 23b. Fruit not a legume; leaflets various, but if numerous, then serrate or with shallow lobes or auricles at the base of the leaflets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (24) 24a. [23] Leaflets 11 or fewer; if more, then the pith not occupying a major portion of the stem section. Rosaceae, Rose Family 24b. Leaflets more than 11; branches stout, the pith occupying a major portion of the cross-section . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (25) 25a. [24] Leaflets serrate; fruits red, round, with a velvety surface. Anacar­diaceae, Sumac Family 25b. Leaflets entire except for basal auricles; fruit an elongate samara with a central seed. Simaroubaceae, Quassia Family 26a. [17] Stems with thorns or spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (27) 26b. Stems spineless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (35) 27a. [26] Thorns often more than 1 cm long, formed by modification of whole branchlets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (28) 27b. Thorns shorter, formed at the nodes (modified leaves or stipules) . . . . . . . . . . . (33) 28a. [27] Leaves linear, often somewhat thick and succulent . . . . . . . . . . . . . . . . . . . . (29) 28b. Leaves not succulent, broader . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (30) 29a. [28] Rigid, short-branched, dense, low shrub less than 1 m tall, with narrowly oblong pale green leaves, the older branches rigid, thorn-like; flowers small, white, with petals and sepals. Crossosomataceae, Crossosoma Family 29b. Shrubs either over 1 m tall or not rigidly branched, with succulent or gray-farinose leaves; flowers lacking petals, subtended by characteristic bracts. Chenopodiaceae, Goosefoot Family 30a. [28] Thorns smooth, sharp, reddish, over 2 cm long, derived from branchlets but along the normal branches. Crataegus in Rosaceae, Rose Family 30b. Thorns either terminating short branchlets or representing stipules, at the leaf bases, less than 2 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (31) 31a. [30] Fruit a berry; petals united; leaves usually with indistinct lateral veins. Lycium in Solanaceae, Nightshade Family 31b. Fruit not a berry; petals separate; leaves usually with distinct lateral veins . . . . . (32) 32a. [30] Leaves with 3 prominent veins; thorn a modified branchlet; low shrub. Ceano­thus in Rhamnaceae, Buckthorn Family 32b. Leaves with pinnate venation; thorn replacing a stipule; introduced tree. Maclura in Moraceae, Mulberry Family 33a. [27] Leaves linear; young twigs woolly tomentose. Tetradymia in Aster­aceae, Sunflower Family 33b. Leaves broader; young twigs not tomentose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (34)

8

Key to the Families: Woody Dicots

34a. [33] Leaves elliptic, entire, toothed, or spine-toothed; fruit elliptical. Berberid­aceae, Barberry Family 34b. Leaves ovate, lobed, and toothed; fruit globose. Grossulariaceae, Gooseb­ erry Family 35a. [26] Leaves pinnately lobed or spine-toothed, leathery; fruit an acorn. Fagaceae, Oak Family 35b. Leaves not as above; fruit not an acorn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (35) 36a. [35] Leaf blades unequal at the base (one side attached lower than the other). Ulmaceae, Elm Family 36b. Leaf blades not unequal at the base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (37) 37a. [36] Leaves broadly ovate-cordate or deeply 5–7-lobed, crenate; fruits juicy, black­ berry-like (a fleshy catkin); introduced. Moraceae, Mulberry Family 37b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (38) 38a. [37] Leaves palmately lobed (sometimes shallowly) . . . . . . . . . . . . . . . . . . . . . . . (39) 38b. Leaves not palmately lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (41) 39a. [38] Foliage aromatic (sagebrush odor). Asteraceae, Sunflower Family 39b. Foliage not aromatic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (40) 40a. [39] Stamens numerous; carpels few to numerous, separate; fruit dry; flowers never tubular. Rosaceae, Rose Family 40b. Stamens 5 or fewer; carpels 2, united, forming a fleshy berry; flowers often tubular. Grossulariaceae, Gooseberry Family 41a. [38] Leaf buds (in the leaf axils) with a single covering scale; leaves from narrowly linear to broadly lanceolate or elliptic. Salix in Salicaceae, Willow Family 41b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (42) 42a. [40] Petiole flattened perpendicular to the leaf face; leaves deltoid-serrate, evenly serrate, or crenulate. Populus, in Salicaceae, Willow Family 42b. Petiole not flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (43) 43a. [41] Either staminate or carpellate catkins present through part or most of the year (in Alnus the carpellate catkin is persistent and woody, and in Corylus it bears a solitary flower and fruit); leaves irregularly serrate. Betu­laceae, Birch Family 43b. Catkins not present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (44) 44a. [43] Flowers in heads, each flower cluster surrounded by an involucre; mostly low desert shrubs. Asteraceae, Sunflower Family 44b. Flowers not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (45) 45a. [43] Desert shrubs with farinose pubescence. Chenopodiaceae, Goosefoot Family 45b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (46) 46a. [45] Leaves never with 3 prominent veins; flowers usually vase-shaped (but wide open in Azaleastrum and campanulate in Gaultheria); petals united, waxy. Erica­ceae, Heath Family 46b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (47) 47a. [46] Leaves with 3 prominent veins (or, in Rhamnus and Frangula, with many parallel lateral veins), elliptic or ovate; stamens opposite the petals. Rham­naceae, Buck­ thorn Family 47b. Leaves variously toothed or lobed; stamens numerous or not opposite the petals (when in doubt, it is usually this choice; it is just difficult to write a key separating these families without flowers). Rosaceae, Rose Family

Key to the Families: Herbaceous Dicots

9

KEY F (Herbaceous dicots) 1a. Cauline leaves in whorls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 1b. Cauline leaves not in whorls . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6) 2a. Leaves in a single whorl of broad leaf blades at the top of the stem. Corn­aceae, Dogwood Family 2b. Leaves in several whorls along the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 3a. Stems square, usually with recurved hooks. Rubiaceae, Madder Family 3b. Stems not square, not armed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 4a. Plant prostrate; petals lacking. Molluginaceae, Carpetweed Family 4b. Plant erect; petals present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) 5a. Plants tall, stout; leaves oblanceolate; flowers 4-merous, greenish. Frasera, in Gen­tianaceae, Gentian Family 5b. Plants low, slender; leaves linear; flowers 5-merous, white. Spergula in Alsina­ceae, Chickweed Family 6a. Flowers several to many, sessile in heads, each flower cluster surrounded or sub­ tended by an involucre. Key F1 6b. Flowers not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (7) 7a. Perianth none or of a single set of parts (tepals or sepals), these all much alike in color and texture. Key F2 7b. Perianth present, evidently double, the outer segments (sepals) and inner segments (petals) usually conspicuously different in texture, color, or both . . . . . . . . . . . . . (8) 8a. Petals separate. Key F3 8b. Petals united (at least at the base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (9) 9a. Corolla radially symmetrical. Key F4 9b. Corolla bilaterally symmetrical. Key F5

KEY F1 (Flowers in heads with involucre) 1a. Leaves linear, all basal; head single, scapose; flowers pink, the head papery-textured; ovary superior. Limoniaceae, Thrift Family 1b. Not as above; ovary inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. [1] Involucre papery or umbrella-like, consisting of a single, undivided cup; flowers obviously separate and not tightly confined by the involucre. Nyctagin­aceae, Fouro’clock Family 2b. Involucre not papery or umbrella-like, the flower cluster a dense head . . . . . . . . (3) 3a. [2] Inflorescence a spike of inconspicuous flowers, below which is a circle of large white or pink petal-like bracts, producing the effect of a single flower; leaves basal, fleshy; plant stoloniferous, pepper-scented. Saururaceae, Lizardtail Family 3b. Plants not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 4a. [3] Corolla 4-lobed; stamens separate. Dipsacaceae, Teasel Family 4b. Corolla 5-lobed or strap-shaped or of both types; stamens with united anthers. Asteraceae, Sunflower Family

KEY F2 (Perianth absent or of tepals) 1a. 1b. 2a. 2b.

Plants dioecious . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) Plants not dioecious . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) [1] Leaves simple, not lobed or compound. Amaranthaceae, Amaranth Family Leaves lobed or compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3)

10

Key to the Families: Herbaceous Dicots

3a. [2] Staminate flowers in racemes, carpellate in clusters; fruit nut-like; leaves digitate­ly compound with narrow serrate leaflets. Cannabis in Can­nabaceae, Hops Family 3b. All flowers in open panicles; achenes ribbed, in clusters; leaves ternately compound with small palmately lobed leaflets. Thalictraceae, Meadowrue Family 4a. [1] Ovary inferior or half inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) 4b. Ovary superior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (9) 5a. [4] Ovary with 2 locules, 1 ovule in each; fruit 2-seeded . . . . . . . . . . . . . . . . . . . . . (6) 5b. Ovary with 1 locule, this with 1–2 ovules (or ovary with 1–3 locules but only 1 locule containing an ovule); fruit 1-seeded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (7) 6a. [5] Petals united at the base; leaves opposite or whorled; flowers in cymes, never umbels. Rubiaceae, Madder Family 6b. Petals separate; leaves alternate or basal; flowers in umbels. Apiaceae, Parsley Family 7a. [5] Leaves alternate, glaucous; flowers greenish white; fruit a drupe. Comandraceae, Comandra Family 7b. Leaves opposite; flowers white or pink; fruit an achene . . . . . . . . . . . . . . . . . . . . . (8) 8a. [7] Leaves simple, entire; flowers pinkish or flesh-colored; fruits hard and bony or with papery wings. Nyctaginaceae, Four-o’clock Family 8b. Leaves pinnately lobed or divided; flowers white; fruits provided with a delicate parachute of feathery bristles. Valerianaceae, Valerian Family 9a. [4] Carpels separate, 2–many in each flower (if 1, the fruit a fleshy several-seeded berry); stamens usually numerous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (10) 9b. Carpels solitary or several united; stamens 1–many (usually not over 10 in most families) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (13) 10a. [9] Tepals greenish, shorter than the stamens; achenes ribbed; herbaceous plants with compound leaves, the leaflets ovate, lobed. Thalictraceae, Meadowrue Family 10b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (11) 11a. [10] Carpels enclosed in a 4-angled calyx cup; leaves with stipules; flowers in a dense head. Sanguisorba in Rosaceae, Rose Family 11b. Carpels not enclosed in the calyx; leaves lacking stipules; flowers not in a dense head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (12) 12a. [11] Carpels consisting of dehiscent, several-seeded follicles (in Actaea a berry). Helleboraceae, Hellebore Family 12b. Carpels consisting of 1-seeded indehiscent achenes. Ranunculaceae, Butterc­ up Family 13a. [9] Ovary with 2 or more locules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (14) 13b. Ovary with 1 locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (19) 14a. [13] Plants with milky juice. Euphorbiaceae, Spurge Family 14b. Plants without milky juice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (15) 15a. [14] Flowers unisexual; ovary on a stalk (in this group the flowers are reduced to single stamens on single gynoecia, but the stamens and gynoecium are surrounded by a cup-like involucre that resembles a perianth). Euphor­biaceae, Spurge Family 15b. Flowers perfect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (16) 16a. [15] Leaves opposite or whorled, entire; stamens 1–many (rarely 2); flowers axillary, solitary, or in small clusters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (17) 16b. Leaves alternate or crowded at the base of the stem, usually toothed; stamens 2; flowers in terminal spikes or racemes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (18)

Key to the Families: Herbaceous Dicots

11

17a. [16] Calyx lobes hooded, with a subapical, prolonged appendage; capsule circumscis­ sile; plants perennial, prostrate, succulent. Aizoaceae, Fig-Marigold Family 17b. Calyx lobes not as above; capsule not circumscissile. Alsinaceae, Chickw ­ eed Family 18a. [16] Perennials; flowers in spikes; fruit several-seeded. Besseya in Scrophulari­aceae, Figwort Family 18b. Annuals; flowers in racemes; fruit 2-seeded (1 seed in each locule). Lepidium in Brassicaceae, Mustard Family 19a. [13] Ovary with several–many ovules; fruit a capsule . . . . . . . . . . . . . . . . . . . . . . (20) 19b. Ovary with 1 ovule; fruit a 1-seeded achene or utricle . . . . . . . . . . . . . . . . . . . . . (21) 20a. [19] Perianth of united tepals, pink; leaves oblong, glaucous. Glaux in Primula­ceae, Primrose Family 20b. Perianth of separate tepals; leaves otherwise. Alsinaceae, Chickweed Family 21a. [19] Leaves with stipules either papery or sheathing the stem . . . . . . . . . . . . . . . (22) 21b. Stipules none or not sheathing the stem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (23) 22a. [21] Leaves opposite; stipules papery. Paronychia in Alsinaceae, Chickw ­ eed Family 22b. Leaves alternate; stipules united around the stem in a sheath just above the nodes. Polygonaceae, Buckwheat Family 23a. [21] Conspicuous, persistent stipules present; leaves opposite . . . . . . . . . . . . . . . (24) 23b. Stipules lacking; leaves alternate (except in a few Amaran­thaceae) . . . . . . . . . . . (25) 24a. [23] Plants small, with spreading, prostrate, or densely caespitose stems rarely over 30 cm tall; stinging hairs not present. Alsinaceae, Chickweed Family 24b. Plants with erect stems usually over 30 cm tall; stinging hairs present. Urticaceae, Nettle Family 25a. [23] Flowers perfect, the flower clusters subtended by a cup-like involucre; stamens 6–9; fruit an achene. Polygonaceae, Buckwheat Family 25b. Flowers perfect or unisexual but not subtended by a cup-like involucre; stamens 1–5; fruit an achene or utricle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (26) 26a. [25] Bracts and perianth ± papery or membranaceous. Amaranthaceae, Amaranth Family 26b. Bracts and perianth herbaceous to fleshy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (27) 27a. [26] Style and stigma 1; leaves alternate and entire; fruit an achene; annuals. Parietaria in Urticaceae, Nettle Family 27b. Styles and stigmas 1–3 (but, if 1, the leaves toothed); fruit a utricle; annuals or peren­­nials; weedy species, often coarse and scurfy-pubescent. Chenopodi­aceae, Goosefoot Family

KEY F3 (Petals present, separate) 1a. Stamens alternating with branched staminodia with terminal yellow anther-like glands. Parnassiaceae, Grass-of-Parnassus Family 1b. Stamens lacking alternating branched staminodia . . . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. [1] Sepals and petals 3, petals shorter than sepals; ovary superior, fruit a schizocarp of 3 nutlets; delicate wetland plants with pinnatifid leaves. Limnanthaceae, Meadow­ foam Family 2b. Floral parts not in 3s; otherwise not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 3a. [2] Ovary inferior, at least part of the lower half fused to the hypanthium or calyx tube . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 3b. Ovary superior (if hypanthium present, the ovary may seem inferior, but dissection shows it is not imbedded in the hypanthium tissues; cf. rose hip) . . . . . . . . . . . . . (9)

12

Key to the Families: Herbaceous Dicots

4a. [3] Prostrate succulent herbs with rounded oblanceolate, thick leaves; stamen number variable; petals yellow. Portulaca in Portulacaceae, Purslane Family 4b. Not prostrate, succulent herbs; otherwise not as above . . . . . . . . . . . . . . . . . . . . . (5) 5a. [4] 2 or more styles present. Saxifragaceae, Saxifrage Family 5b. Only 1 style present (stigmas may be lobed) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6) 6a. [5] Foliage sandpapery, with minutely barbed hairs. Loasaceae, Loasa Family 6b. Plants lacking barbed hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (7) 7a. [6] Stamens 2, 4, or 8; petals 2 or 4; style 1, locules usually 4 (1 in Circaea). Ona­graceae, Evening-Primrose Family 7b. Stamens 5, rarely 4; petals usually 5; styles 2 or more; locules 2–6 . . . . . . . . . . . . (8) 8a. [7] Locules 4–6; fruit a several-seeded berry; leaves basal, ternately compound. Araliaceae, Ginseng Family 8b. Locules 2; fruit dry, separating into 2, 1-seeded mericarps. Apiaceae, Parsley Family 9a. [3] Flowers asymmetrical; petals 4–7, toothed or cleft; stamens numerous, inserted on one side of the flower; leaves pinnatifid. Resedaceae, Mignone­ tte Family 9b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (10) 10a. [9] Corolla bilaterally symmetrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (11) 10b. Corolla radially symmetrical . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (17) 11a. [10] Leaves pinnately or palmately compound . . . . . . . . . . . . . . . . . . . . . . . . . . . (12) 11b. Leaves simple, entire to deeply lobed or pinnatifid, but never com­pound . . . . . (14) 12a. [11] Sepals 2, very minute and scale-like; corolla spurred; leaves greatly dissected. Fumariaceae, Fumitory Family 12b. Sepals 4 or 5; corolla not or very inconspicuously spurred; leaves once or twice compound . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (13) 13a. [12] Ovary with 1 placenta; petals 5 (a banner, 2 wings, and a keel that consists of 2 partly united petals enclosing the stamens and style); flowers usually shaped like those of sweet-pea. Fabaceae, Pea Family 13b. Ovary with 2 placentae on opposite sides of the ovary; petals 4; stamens exserted. Cleomaceae, Cleome Family 14a. [11] Stamens many; carpels more than 1; capsule dehiscing along 1 suture (thus a follicle). Helleboraceae, Hellebore Family 14b. Stamens 10 or fewer; ovary of a single or several united carpels; fruit a dehiscent capsule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (15) 15a. [14] Flowers not spurred, but with a larger upper petal (the banner). Polygalaceae, Milkw ­ ort Family 15b. Flowers spurred . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (16) 16a. [15] 1 sepal spurred, petaloid (only 3 present); ovary with 5 locules; flower shaped like a gnome’s slipper. Balsaminaceae, Jewelweed Family 16b. Sepals not spurred (5 sepals present), but a petal prominently spurred; ovary with 1 locule and 3 parietal placentae. Violaceae, Violet Family 17a. [10] Stamens of the same number as the petals and opposite them . . . . . . . . . . . (18) 17b. Stamens fewer or more numerous than the petals, or, if the same number, then alternate with them . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (21) 18a. [17] Sepals, petals, and stamens each 6 in number, 3 of the sepals petal-like; leaf margins spiny. Berberidaceae, Barberry Family 18b. Sepals, petals, and stamens 2–5 (sepals rarely 6); spineless . . . . . . . . . . . . . . . . . . (19) 19a. Hypanthium present, stamens 5. Rosaceae 19b. Hypanthium absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (20)

Key to the Families: Herbaceous Dicots

13

20a. [19] Styles, stigmas 1; sepals usually 5. Primulaceae, Primrose Family 20b. Styles, stigmas 2 or more; sepals usually 2. Portulacaceae, Purslane Family 21a. [17] Ovary deeply lobed; plants with glandular dots, very strongly scented; floral parts in 4s. Thamnosma, in Rutaceae, Rue Family 21b. Plants not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (22) 22a. [21] Ovary 1 (a single unit) with 1 locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (23) 22b. Ovaries more than 1, or, if 1, then with 2 or more locules . . . . . . . . . . . . . . . . . . (37) 23a. [21] Stamens 13 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (24) 23b. Stamens 12 or fewer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (30) 24a. [23] Ovary simple (of a single carpel having 1 placenta, 1 style, 1 stigma; many such ovaries may be present in a single flower) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (25) 24b. Ovary compound (2 or more placentae, styles, or stigmas) . . . . . . . . . . . . . . . . . (26) 25a. [24] Fruit a 1-seeded achene. Ranunculaceae, Buttercup Family 25b. Fruit a several-seeded, dehiscent follicle or a fleshy berry. Hel­leboraceae, Hellebore Family 26a. [24] Placenta free-central or basal. Portulacaceae, Purslane Family 26b. Placentae parietal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (27) 27a. [26] Ovary with 2 parietal placentae; plants usually viscid and ill-smelling. Cleomaceae, Cleome Family 27b. Ovary with 3 or more placentae; plant not viscid nor ill-smelling . . . . . . . . . . . . (28) 28a. [27] Leaves simple, alternate, entire; flowers of 2 kinds, conspicuous ones with large yellow petals and many stamens, and inconspicuous, cleistogamous, axillary ones lacking petals. Cistaceae, Rockrose Family 28b. Plants not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (29) 29a. [28] Leaves opposite, entire, with minute translucent dots (hold up to the light); juice not milky; leaves and petals often with black marginal dots. Hypericaceae, St. Johnswort Family 29b. Leaves alternate, toothed or lobed, without translucent dots; juice milky; calyx forced off intact as a cone by the swelling petals; flowers white or cream-colored, rarely yellow or orange. Papaveraceae, Poppy Family 30a. [23] Gynoecium a single carpel with 1 placenta, style, and stigma . . . . . . . . . . . (31) 30b. Gynoecium compound (more than 1 placenta, style, or stigma) . . . . . . . . . . . . . (32) 31a. [30] Stamens and petals attached to the rim of the calyx tube (hypanthium). Rosa­ceae, Rose Family 31b. Stamens and petals not attached to the calyx tube. Fabaceae, Pea Family 32a. [30] Petals inserted on the throat of a bell-shaped or tubular calyx. Lyth­raceae, Loosestrife Family 32b. Petals inserted on the receptacle; calyx of separate or united sepals . . . . . . . . . . (33) 33a. [32] Ovules attached to base of ovary or to a free-central placenta . . . . . . . . . . . (34) 33b. Ovules attached to 2 or more parietal placentae . . . . . . . . . . . . . . . . . . . . . . . . . . (36) 34a. [33] Calyx of united sepals; petals with claws. Caryophyllaceae, Pink Family 34b. Calyx of separate sepals; petals not stalked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (35) 35a. [34] Sepals 2 or numerous; stamens commonly opposite the petals, sometimes fewer than the petals, sometimes numerous; leaves commonly basal and succulent. Portulacaceae, Purslane Family 35b. Sepals usually 5; stamens not opposite the petals, usually 5 or 10 (rarely 3); leaves usually opposite, not very succulent. Alsinaceae, Chickweed Family

14

Key to the Families: Herbaceous Dicots

36a. [33] Ovary with 2 parietal placentae; sepals and petals 4 each. Cleomaceae, Cleome Family 36b. Ovary with 3–5 parietal placentae; sepals and petals 5. Hypericaceae, St. Johnsw ­ ort Family 37a. [22] Plants with milky juice or stinging hairs; ovary stipitate, exserted from the cyathium. Euphorbiaceae, Spurge Family 37b. Plants without milky juice or stinging hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (38) 38a. [37] Flower parts in 2s or 4s. Brassicaceae, Mustard Family 38b. Flower parts in 5s or numerous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (39) 39a. [38] Leaves trifoliolate, acrid tasting. Oxalidaceae, Woodsorrel Family 39b. Leaves not trifoliolate nor acrid tasting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (40) 40a. [39] Stamens united in a column around the styles. Malvaceae, Mallow Family 40b. Stamens not united in a column around the styles . . . . . . . . . . . . . . . . . . . . . . . . (41) 41a. [40] Leaves simple, linear or oblong, succulent. Crassulaceae, Stonecrop Family 41b. Leaves not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (42) 42a. [41] Stamens numerous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (43) 42b. Stamens not more than 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (45) 43a. [42] Leaves elliptic, with translucent dots and often minute, black, marginal dots on leaves and petals; stamens tending to be in 5 groups; fruit a capsule. Hyperica­ceae, St. Johnswort Family 43b. Leaves and stamens not as above; fruit an achene . . . . . . . . . . . . . . . . . . . . . . . . . (44) 44a. [43] Stipules lacking; hypanthium not developed. Ranunculaceae, Buttercup Family 44b. Stipules present; hypanthium always developed. Rosaceae, Rose Family 45a. [42] Petals waxy; anthers opening by terminal pores; leaves often leathery. Pyrola­ceae, Wintergreen Family 45b. Petals not waxy; anthers opening by slits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (46) 46a. [45] Fruit 5-carpellate, separating at maturity into 5, 1-seeded segments (meri­carps) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (47) 46b. Fruit not 5-carpellate, or, if so, not separating into mericarps . . . . . . . . . . . . . . . (48) 47a. [46] Flowers pink or white; mericarps not spiny. Geraniaceae, Geranium Family 47b. Flowers yellow to orange; mericarps stoutly spiny. Zygophyllaceae, Caltrop Family 48a. [46] Petals yellow, copper, or blue, falling within a few hours; capsule 10-locular (5 locules each with an additional septum). Linaceae, Flax Family 48b. Petals white, yellow, or pink-purple, not fugacious; capsule not as above . . . . . . (49) 49a. [48] All leaves opposite, with scarious stipules; stem glandular-hairy; low, prostratespreading plants of mudflats in the piedmont valleys. Bergia, in Elatina­ceae, Water­ wort Family 49b. Plants not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (50) 50a. [48] Calyx completely free from the ovary but tightly surrounding it; petals mount­ed on the calyx tube. Lythraceae, Loosestrife Family 50b. Calyx not closely surrounding the ovary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (51) 51a. Calyx fused to the lower part of the ovary only; petals on the margin of a hypan­ thium (flowers perigynous). Saxifragaceae, Saxifrage Family 51b. Calyx free; flowers hypogynous. Zygophyllaceae

Key to the Families: Herbaceous Dicots

15

KEY F4 (Petals united, radially symmetrical) 1a. 1b. 2a. 2b.

Plants with milky juice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) Plants without milky juice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) [1] Corolla bell-shaped, without special horn-like structures . . . . . . . . . . . . . . . . . (3) Corolla rotate, with a central body consisting of the fused stigmas and stamens; corona present, enclosing horn-like structures. Asclepiadaceae, Milkweed Family 3a. Ovary superior. Apocyn­aceae, Dogbane Family 3b. Ovary inferior. Campanulaceae, Bellflower Family 4a. [1] Ovary superior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) 4b. Ovary inferior or half inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (18) 5a. [4] Ovary deeply 2-lobed, circumscissile; corolla yellow, 7–12 mm long, the lobes longer than the tube. Menodora in Oleaceae, Olive Family 5b. Ovary not 2-lobed nor circumscissile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6) 6a. [5] Stamens more numerous than the corolla lobes, 6–many . . . . . . . . . . . . . . . . . (7) 6b. Stamens as many as the corolla lobes or fewer . . . . . . . . . . . . . . . . . . . . . . . . . . . . (8) 7a. [6] Stamens many, united into a tube around the style. Malvaceae, Mallow Family 7b. Stamens 6–10, separate and distinct; anthers opening by pores at the basal end; petals waxy. Pyrolaceae, Wintergreen Family 8a. [6] Stamens 5, opposite the petals; ovary with 1 locule; placenta basal or free-central. Primulaceae, Primrose Family 8b. Stamens alternate to the petals or fewer; ovary more than 1-loculed, or, if 1-loculed, then the placenta rarely basal or free-central . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (9) 9a. [8] Ovary 4-lobed, developing into 4 (or by abortion fewer), 1-seeded nutlets . . (10) 9b. Ovary not 4-lobed; fruit a capsule or berry, usually several-seeded . . . . . . . . . . . (11) 10a. [9] Leaves alternate; stem not square in cross-section; not aromatic. Boragin­aceae, Borage Family 10b. Leaves opposite; stems square in cross-section. Lamiaceae, Mint Family 11a. [9] Ovary with 1 locule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (12) 11b. Ovary with 2 or more locules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (14) 12a. [11] Leaves basal; flowers solitary, scapose; stoloniferous plants rooted in mud. Limosella in Scrophulariaceae, Figwort Family 12b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (13) 13a. [12] Leaves opposite or whorled, entire; style 1 or none; plants mostly glabrous; inflorescence not curled in the bud. Gentianaceae, Gentian Family 13b. Leaves usually alternate (if opposite, then not entire); styles 2, or single and 2-cleft above; plants mostly hairy; inflorescence commonly curled in the bud. Hydrophyl­laceae, Waterleaf Family 14a. [11] Stigma 3-lobed or style 3-branched; ovary with 3 locules. Polemoni­aceae, Phlox Family 14b. Stigma entire or 2-lobed, or style 2-cleft; ovary usually with 2 locules . . . . . . . . (15) 15a. [14] Flowers yellow, in dense terminal spikes or racemes over 20 cm long; filaments hairy. Verbascum in Scrophulariaceae, Figwort Family 15b. Flowers variously colored, never in spikes or elongate racemes . . . . . . . . . . . . . (16)

16

Key to the Families: Herbaceous Dicots

16a. [15] Styles 2, distinct, each one again 2-cleft; ovules 2 in each locule; flowers axillary, the corolla lavender, with darker pleats; foliage silky-hairy. Convolvu­laceae, Morning­-glory Family 16b. Style 1, or if 2, rarely separate to the base, never again 2-cleft; ovules usually more than 2 per locule; inflorescence various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (17) 17a. [16] Style 1, the stigma entire or 2-lobed; fruit a capsule or berry. Solanaceae, Nightshade Family 17b. Styles 2 or definitely 2-branched below the stigmas; fruit a capsule. Hydrophyl­laceae, Waterleaf Family 18a. [4] Leaves ternately compound, basal; flowers in a few-flowered, tight umbel-like cyme; delicate herbs. Adoxaceae, Adoxa Family 18b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (19) 19a. [18] Leaves alternate or basal. Campanulaceae, Bellflower Family 19b. Leaves opposite or whorled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (20) 20a. [19] Stem creeping, slightly woody; leaves opposite, crenate; flowers 2, pink, pendent from an erect stalk. Linnaea in Caprifoliaceae, Honeysuckle Family 20b. Stem erect or sprawling, herbaceous; leaves opposite or whorled, entire; flowers white, minute, in cymes. Rubiaceae, Madder Family

KEY F5 (Petals united, bilaterally symmetrical) 1a. Corolla with 3 petals united at the base by their narrowed claws, the other 2 minute, gland-like; fruit globose, spiny. Krameriaceae, Ratany Family 1b. Not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. [1] Ovary superior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 2b. Ovary inferior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (7) 3a. [2] Stem 4-angled; leaves opposite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 3b. Stem not 4-angled; leaves opposite, alternate, or basal . . . . . . . . . . . . . . . . . . . . . . (5) 4a. [3] Corolla usually strongly 2-lipped; style arising from the base of the gynoecium between the nutlets; foliage usually with a minty odor. Lami­aceae, Mint Family 4b. Corolla open, flat, from a narrow tube; style terminal; foliage never with a minty odor. Verbenaceae, Vervain Family 5a. [3] Corolla papery; leaves basal; inflorescence a spike; fruit a circumscissile capsule (dehiscent as one would open a soft-boiled egg). Plantaginaceae, Plantain Family 5b. Corolla not papery; otherwise not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (6) 6a. [5] Calyx split to the base along the lower side; corolla 3–5 cm long and nearly as wide; fruit woody, with 2 long curved claws. Martyniaceae, Unicorn plant Family 6b. Calyx not split to the base on the lower side; corolla various; fruit not as above. Scrophulariaceae, Figwort Family 7a. [2] Flowers not spurred or gibbous-based but highly colored, red, or blue-and-white striped. Lobelia, in Campanulaceae, Bellflower Family 7b. Flowers gibbous-based, white or cream-colored. Valerianaceae, Valerian Family

FERNS AND FERN ALLIES 1a. Plants floating on the water, about 1 cm long, with minute, sessile, 2-lobed leaves arranged on 2 sides of the stems, giving a braided appearance. Salviniaceae, Waterfern Family 1b. Plants not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. Stems jointed, hollow, green (except the fertile stems of Equisetum arvense, which are yellowish brown), the nodes circled by sheaths. Equisetaceae, Horsetail Family 2b. Stems not jointed, seldom green; sheaths absent . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 3a. Aquatic, inhabiting lakeshores or actually submerged in ponds and lakes . . . . . . (4) 3b. Terrestrial, growing on soil or rocks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (5) 4a. Leaves grass-like, their bases swollen, each bearing a pair of sporangia, the whole forming an onion-like bulb; plants submerged in shallow water of mountain lakes and ponds for the greater part of the growing season. Isoëtaceae, Quillwort Family 4b. Leaves with distinct petioles and blades, the blades 4-parted, resembling a four-leaf clover; spores borne at the base of the plant in round, nut-like “sporocarps”; borders of ponds and sandy streamsides at lower altitudes. Marsileaceae, Pepperwort Family 5a. Leaves very numerous, lanceolate or linear, often bract-like, sessile, spirally or oppositely arranged in 4 to many ranks upon branched perennial stems . . . . . . . . . . . (6) 5b. Leaves relatively few, broad or ± dissected (except in Asplenium sep­tentrionale, the Grassfern, which has linear leaves), arising from an underground stem . . . . . . . . (7) 6a. Leaves minute (less than 3 mm long). Selaginellaceae, Little Club-moss Family 6b. Leaves larger (5 mm–1 cm or more long). Lycopodiaceae, Club-moss Family 7a. Fronds (the “leaves” of ferns) narrowly linear, divided towards the tip into a few narrow forks. Asplenium septentrionale, in Aspleniaceae, Spleenwort Family 7b. Fronds broader, ± dissected (fern-like) [entire in Ophioglossum] . . . . . . . . . . . . . . . (8) 8a. Not all fronds alike; entire fronds or branches modified for spore produc­tion . . . (9) 8b. All fronds essentially alike, whether fertile or sterile . . . . . . . . . . . . . . . . . . . . . . . (11) 9a. One entire branch of each frond completely altered in appearance, modified for spore production, the remainder of the frond green and not producing spores. Ophioglossaceae, Moonwort Family 9b. Fronds of 2 kinds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (10) 10a. Tall plants of wet forests in the foothills; sterile fronds tall with broad lobes; fertile fronds with pinnules forming globular, berry-like divisions containing sporangia; rare (or extinct in Colorado). Onocleaceae, Sensitive Fern Family 10b. Small plants of rocky sites or calcareous cliffs; sterile fronds short with much divided pinnae, their ultimate divisions blunt-tipped; fertile fronds taller, with podshaped pinnules. Common. Cryp­togrammaceae, Rock Brake Family 11a. Fronds golden underneath, palmately branched, curling up when dry. Notholaena standleyi in Cheilanthaceae, Lip Fern Family 11b. Fronds not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (12) 12a. Fronds with the lower pair of branches ± equaling the central branch or facing forward, creating the illusion of a 3-branched frond . . . . . . . . . . . . . . . . . . . . . . . . . (13) 12b. Fronds distinctly pinnate, with 1 main axis from which the pinnae arise along the sides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (14)

17

18

Ferns and Fern Allies

13a. Fronds tall and coarse, forming thicket-like stands; sporangia, when present, borne on infolded edges of the pinnules. Hypolepidaceae, Bracken Family 13b. Fronds small, delicate; sporangia, when present, borne on the flat undersides of the pinnules. Gymnocarpium in Athyriaceae, Ladyfern Family 14a. Fronds with submarginal sori, the indusium formed by the rolled or folded edge of the pinnule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (15) 14b. Fronds with the sori on the face of the pinnule . . . . . . . . . . . . . . . . . . . . . . . . . . . (16) 15a. Pinnules large, thin, smooth, green, fan-shaped or reniform. Adiantaceae, Maiden­ hair Family 15b. Pinnules small, often thickish, hairy or scaly or wax-coated below. Cheilanthaceae, Lipfern family 16a. Frond merely deeply pinnatifid, the bases of the pinnae fused to the stipe. Polypodi­aceae, Polypody Family 16b. Frond distinctly pinnate, the pinnae not fused to the sides of the stipe . . . . . . . . (17) 17a. Sori linear-elliptic; indusium curved, crescentic, or absent. Athyri­aceae, Ladyfern Family 17b. Sori round or greatly elongate; indusia not crescent-shaped . . . . . . . . . . . . . . . . (18) 18a. Plants large, stout; indusium forming a somewhat circular shield, attached at the sinus. Dryopteridaceae, Shieldfern Family 18b. Plants small, delicate; indusia not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (19) 19a. Sori greatly elongate, with a flap-like indusium attached along a vein. Aspleniaceae, Spleenwort Family 19b. Sori round or, if slightly elongate, then without indusia . . . . . . . . . . . . . . . . . . . (20) 20a. Indusium cup-like, attached centrally under sporangia, deeply cleft at matur­ity; pinnae thickish, widest above the middle. Woodsi­aceae, Woodsia Family 20b. Indusium hood-like, attached by its side at the base, not cleft at maturity; pinnae thin, broadest below the middle or at the base. Cystopteris in Athyri­aceae, Ladyfern Family

ADIANTACEAE Newman 1840. MAIDENHAIR FAMILY (ADI) Maidenhairs are the most delicate and rarest of our ferns. The fronds are sometimes branched once dichotomously, with the pinnae radiating from one side of each branch like the comb of a Spanish mantilla. In our species the frond has 1 main branch, from which the pinnae arise on each side in a pyramidal form. The sori are arranged in a row underneath the folded margin of the pinnule. Most species are found in the tropical highlands. ADIANTUM L. 1753 [the ancient name, meaning un­wetted, shedding raindrops] One species, A. capillus-veneris L., Venus’s Hair Fern. Rare or infrequent, on dripping sandstone cliffs, Mesa de Maya.

ASPLENIACEAE Newman 1840. SPLEENWORT FAMILY (ASL) One of the crown jewels of the Colorado flora belongs to this family. D. M. Andrews, a Boulder nurseryman, discovered a fern at the turn of the last century, in horizontal crevices on south-facing cliffs east of Boulder. Aven Nelson described it in 1904 as a new species Asplenium andrewsii. We now include this taxon in the widely world-disjunctive A. adiantumnigrum. Though common in other parts of the world, this fern is very rare in North America and deserves federal protection.

Ferns and Fern Allies

19

ASPLENIUM L. 1753 [Greek, asplenon, a name used by Dioscorides for some fern supposed to cure spleen diseases]. Spleenwort 1a. Fronds linear, simple or only forked at the tips, in dense clumps in rock crevices, resembling dark green grass. A. septentrionale (L.) Hoffmann [northern], Grass ­Fern, 6A. Fairly common on arkosic rocks in the outer foothills. A species widely scattered in the mountains of the Northern Hemisphere, including the Altai, Alps, Pyrenees, the Caucasus, and in Scandinavia. 1b. Fronds with distinct pinnae, not grass-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. Fronds broadly triangular in outline, with branches bearing pinnae. A. adiantumnigrum L., 2A. Restricted to sandstone cliffs, White Rocks, BL. Protected. (A. andrewsii.) 2b. Fronds unbranched, with lateral oblong pinnae . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 3a. Pinnae opposite. A. resiliens Kunze [resilient]. Sandstone cliffs, Mesa de Maya region. 3b. Pinnae alternate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 4a. Pinnae with a basal auricle (shallow lobe), at least on the leading edge. A.  platy­neuron Britton, Sterns & Poggenburg [flat vein]. Ebony Spleenwort. Sandstone cliffs and rimrock, BA, LA. ­ ort, 1C. 4b. Pinnae lacking any auricles. A. trichomanes L., Maidenhair Spleenw Infrequent, cliff crevices, canyons of eastern foothills.

ATHYRIACEAE Alston 1956. LADY FERN FAMILY (ATY) The earmark of the lady ferns is in the shape of the indusium, which, instead of being round as in the shield ­ferns, is crescent- or comma-shaped. But the indusium varies from genus to genus in the family. In the As­­plen­iaceae the indusium is linear. These are indications of how important it is for students to learn the fine details of the sporangia, sori, indusium, and, in fact, the venation of ferns, since the sori are positioned at different points on the fronds vis-àvis the veins of the pinnules. 1a. Fronds large, over 3 dm long, up to 15 cm or more wide. Athyrium, Lady Fern 1b. Fronds less than 3 dm long, up to about 10 cm wide . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. Fronds broadly triangular, with 3 main branches . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 2b. Fronds pinnate. Cystopteris, Brittle Fern 3a. Pinnules oblong, not deeply toothed. Gymnocarpium, Oak Fern 3b. Pinnules tapered, deeply toothed or lobed. Cysto­pteris, Brittle Fern ATHYRIUM Roth 1799 [Greek, athyros, doorless, the sporangia only tardily forcing back the margin of the indusium]. Lady Fern Subspecific taxa are poorly understood in this genus. See the FNA for details. 1a. Indusium crescentic, usually plainly visible; frond well expanded, the pinnae not appearing crowded or directed sharply toward the apex of the frond; fronds few to a clump. A. filix-femina (L.) Roth 1b. Indusium rarely seen, minute, withering early; frond narrow, the pinnae rather crowded or directed sharply toward the frond apex. A. distentifolium Tausch ex Opiz. Subalpine screes. American plants are sometimes called A. americanum (Butters) Maxon. CYSTOPTERIS Bernhardi 1805 [Greek, cystis, bladder, + pteris, fern]. Brittle Fern 1a. Fronds broadly triangular, with three main branches. C. montana (Lamarck) Bernhardi. Rare and local in moist, rich spruce forests, on the Eastern Slope known from a collection near Mount Antero. 1b. Fronds pinnate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2)

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2a. Stipe with a few long brownish hairs near the base of the lower pinnae; lower pinnae pinnate-pinnatifid to twice pinnate; rhizome usually long-creeping; spores averaging 33–41 microns. C. reevesiana Lellinger [for Timothy Reeves, contempo­rary American pteridologist]. Restricted mostly to the southernmost counties. Known to form triploid hybrids with C. fragilis. 2b. Stipe lacking hairs; lower pinnae pinnatifid to pinnate-pinnatifid; rhizome shortcreeping; spores averaging 39–60 microns. C. fragilis (L.) Bernhardi, 2E. Our most common fern, from low to high altitudes. The stipes are produced in clusters and are very brittle. GYMNOCARPIUM Newman 1851 [Greek, gymno, naked, + carpos, fruit]. Oak Fern One species, G. dryopteris (L.) Newman, 2B. Shaded woods and thickets, mon­tane, subalpine.

CHEILANTHACEAE Nayar 1970. LIP FERN FAMILY (CHL) The name “lip fern” alludes to the fact that the margins of the pinnules curve over the sori, not in itself a really diagnostic character, since it is shared by other families. The family is essentially one of the dry tropics, only a few of them reaching Colorado in rocky steppe-desert and foothill sites. A variety of growth forms occur, some pinnate, others dichotomous or even triangular. The species often are distinguished by deposits of white or yellow wax, or various combinations of hairs and scales, on the stipes and pinnules. 1a. Fronds green and glabrous. Pellaea, Cliff Brake 1b. Fronds either hairy or scaly, or with a white or yellow wax on the under­side . . . (2) 2a. Fronds with hairs or scales, not waxy beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 2b. Fronds waxy beneath . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (4) 3a. Fronds once pinnate; lower surface densely covered with fringed or stellate scales. Astrolepis, Cloak Fern 3b. Fronds more than once pinnate. Cheilanthes, Lip Fern 4a. Fronds triangular, with a yellow wax beneath. Noth­olaena, Lip Fern 4b. Fronds widely branched, with very small ternate pinnules, white waxy beneath. Argyrochosma, Lip­Fern ARGYROCHOSMA Windham 1987 [Greek, argyros, silver, + chosma, flow, alluding to the white waxy coat on the pinnules]. Lip Fern One species, A. fendleri (Kunze) Windham, 2C. Talus and cliff crevices of arid canyonsides. (Cheilanthes cancellata, Notholaena fendleri.) ASTROLEPIS Benham & Windham 1992 [Greek. astro, star, + lepis, scale]. Cloak Fern One species, A. integerrima (Hooker) Benham & Windham. Two records from sandstone cliffs, BA, LA. (Cheilanthes.) CHEILANTHES Swartz 1806 [Greek, cheilos, margin, + anthos, flower]. Lip Fern 1a. Fronds densely tomentose, at least below, the pinnules hairy above; rhizomes erect or decumbent but not long-creeping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 1b. Fronds glabrous except for scales; pinnules glabrous above; rhizomes slender, creeping . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 2a. Scales present on rachises of pinnae; stipes usually scaly. C. eatonii Baker [for D. C. Eaton, 1834–1895, American fern specialist]. Cliffs of the Mesa de Maya, and Front Range canyons to CC. 2b. Fronds reddish-hairy beneath, lacking scales. C. feei Moore [for Antoine L. A. Fée, French pteridologist, 1789–1874]. Sandstone cliffs and overhangs, the most common species.

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3a. Scales of frond rachises ciliate; rhizome scales persisting. C. wootonii Maxon [for E. O. Wooton, 1865–1945, New Mexican botanist, author, with Standley, of Flora of New Mexico]. Cliffs of Mesa de Maya. 3b. Scales not ciliate; rhizome scales deciduous. C. fendleri Hooker. On granitic rocks, foothills of Front Range, and Great Sand Dunes. NOTHOLAENA R. Brown 1810 [Greek, nothos, spurious, + chlaena, cloak, alluding to the nature of the “indusium” of the original species]. Lip Fern One species. N. standleyi Maxon [for P. C. Standley, 1884–1963, botanist of Chicago Field Museum, author of Flora of New Mexico]. Sandstone cliffs of the Mesa de Maya. (Cheilanthes.) PELLAEA Link 1841 [Greek, pellos, dusky, from the dark stipes]. Cliff Brake 1a. Stem scales bicolored, with a dark central region and lighter, brown margins . . . (2) 1b. Stem scales uniformly reddish brown to tan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 2a. Fronds 2–3-pinnate at base, the basal pinnae with more than 3 pairs of pinnules. P. wrightiana Hooker [for Charles Wright, 1811–1855, plant collector]. Cliffs and rimrock of the Mesa de Maya. An isolated population has been found recently in Boulder Mountain Park. 2b. Fronds once pinnate at base, the basal pinnate usually with a trio of closely grouped pinnules. P. truncata Goodding. Cliffs of canyons, known from Royal Gorge and Phantom Canyon near Cañon City. 3a Ultimate segments with the vein glabrous on the upper side, or if not, then the rachises nearly glabrous. P. glabella Mettenius. Cliffs of the Mesa de Maya. 3b. Ultimate segments with the vein sparsely villous on the upper side. P.  atropur­purea (L.) Link [black purple]. In rock crevices of sedimentary formations from LR to BA and LA. Recently P. gastonyi Windham has been collected at one locality in LR by Scott F. Smith. This taxon reputedly originated through hybrids involving P. glabella and P. atropurpurea. It most closely resembles the latter. See the FNA for details.

CRYPTOGRAMMACEAE Pichi-Sermolli 1963. ROCK BRAKE FAMILY (CRG) Our single genus, Cryptogramma, contains two species, one of them extremely common and the other extremely rare. These plants are known by their develop­ment of a special fertile frond in which the pinnules are converted to pod-like sporangial units, with the pinnule margins rolled over the sori. The rare species C. stelleri, instead of being densely tufted, is slender and rhizomatous, forming very few fronds and growing in the crevices of limestone cliffs. A prime locality to discover this might be the limestone areas of the Collegiate and Sangre de Cristo ranges. CRYPTOGRAMMA R. Brown 1823 [Greek, cryptos, hidden, + gramme, a line, alluding to the lines of sporangia]. Rock Brake One species, fronds robust, crowded on a short rhizome; lower parts of the stipes persistent. C.  acro­stichoides R. Brown [like the genus Acro­stichum]. Rocky places, montane to alpine. (C. crispa subsp.)

DRYOPTERIDACEAE Herter 1949. SHIELD FERN FAMILY (DRY) The term “shield fern” alludes to the shape of the indusium, which covers the cluster of sori like a circular shield, attached by the middle. This family is very well represented in eastern North America but only a few species occur in the Rocky Mountain states. Polystichum mu­nitum, a common species in the forests of the Pacific Northwest, is imported to our area for use in floral arrangements. One’s first examination of the sporangia of ferns under the microscope is an exciting experience. Each sporangium is shaped like a swollen coin, with easily ruptured faces and a rim of cells with differentially thickened walls, with the outer wall

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Ferns and Fern Allies

thin and the side and bottom ones thick. The interaction of wetting and drying causes the rim to fly open and reverse itself, scattering the spores far and wide. 1a. Frond simply pinnate, the pinnae spine-toothed, with an auricle at the upper base. Polystichum, Christm ­ as Fern 1b. Frond more than once-pinnate. Dryopteris, Shield Fern DRYOPTERIS Adanson 1763 [Greek, drys, oak, + pteris, fern]. Shield Fern 1a. Frond bipinnatifid, the ultimate divisions blunt; indusium very prominent. D. filix-mas (L.) Schott, Male Fern, 1A. Infrequent, shaded canyons, foothills of Front Range. Formerly gathered for the oil in the rhizomes, from which a worm medicine was prepared; no longer gathered commer­cially, but could be exterminated easily and should be protected. 1b. Frond bipinnate, the ultimate divisions sharp-pointed; indusium not so prominent. D. expansa (C. Presl) Fraser-Jenkins & Jermy. Rare, in rich subalpine forests, known only from Rocky Mountain National Park. Not listed for Colorado in the FNA. (D. assimilis.) POLYSTICHUM Roth 1799 [Greek, polys, many, + stichos, row, referring to the sori]. Christmas Fern One species, P. lonchitis (L.) Roth [a name used by Pliny for some plant with a tongueshaped leaf], Holly Fern, 1B. Infrequent among rocks, alpine and subalpine.

EQUISETACEAE Michaux ex de Candolle 1804. HORSETAIL FAMILY (EQU) The horsetails belong to one of the most ancient lineages of land plants, abundantly found fossilized in the Coal Measures of the Paleozoic era. Their basic structures have changed relatively little over time. The deposition of mineral silica on the surface of the stems gives them the name scouring-rushes, and they served very nicely in scouring out pots and pans in colonial times, as they do now at modern-day campgrounds. What we see is the sporophyte only. The gameto­phytes are almost never encountered, since they are small and rarely produced. 1a. Green stems bearing numerous branches in whorls at the nodes (fertile, simple brown stems also belong here). Equisetum, Horsetail 1b. Green stems stout, simple or occasionally with a few short branches scattered irregularly on the main stem. Hippochaete, Scouring-Rush EQUISETUM L. 1753 [Greek, equus, horse, + Latin, seta, bristle]. Horsetail One species, E. arvense L., 4C. Sterile stems with ascending branches, 4-angled; central cavity one-fourth the diameter of the stem; rhizome with scattered, blackish tubers; fertile stems brown, abruptly wilting; stem smoothly tubercu­late, the teeth in whorls of 3–5(6). Wet ditches and floodplains. The stems are of 2 kinds, branched and sterile, or brown, unbranched, and bearing a sporangiate cone at the apex. The latter are produced in early spring and soon wither. (E. pratense was reported in error.) HIPPOCHAETE Milde 1865 [Greek, hippos, horse, + chaite, mane]. Scouring-Rush Formerly included in Equisetum. Hybrids occur involving all species. 1a. Stems slender, 5–12-angled and -grooved; sheaths loose, with fine-pointed persistent teeth; central cavity of stem usually half its diameter. H. variegata (Schleicher) Bruhin. Sandbars of streams. 1b. Stems stout, 16–48-angled and -grooved; sheaths loose or tight, the teeth persistent or deciduous; central cavity of the stem more than half its diame­ter . . . . . . . . . . (2) 2a. Stem dying after one season; sheaths lacking a dark band at base; cone rounded at apex. H. laevigata (A. Braun) Farwell [smooth], 4A. Wet ground of ditches and streamsides.

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2b. Stem enduring several years; sheaths commonly with a dark band at the base; cone pointed at the apex. H. hyemalis (L.) Bruhin subsp. affinis (A. Braun) W. A. Weber [of winter; related], 4B. Similar habitats.

HYPOLEPIDACEAE Pichi-Sermolli 1970. BRACKEN FAMILY (HPL) Bracken is a fern found almost everywhere in the world, in the tropics as well as the northern and southern temperate zones. Bracken forms an almost impen­etrable brushland on the highlands of the Galápagos Islands. It is easily recognized as a large fern with 3 obvious main branches, and the sori are marginal on the pinnules and protected by their folded edges. Fiddleheads of bracken are said to be edible, but the enzyme thiamin­ase in bracken is very poisonous to livestock if much is ingested, either green or dry. PTERIDIUM Gleditsch 1760 [diminutive of Pteris, another genus]. Bracken One species, P. aquilinum (L.) Kuhn subsp. lan­uginosum (Bongard) Hultén [of an eagle, from the wing-shaped fronds; woolly]. Dry, open woodlands. Our largest native fern.

ISOËTACEAE Reichenbach f. 1828. QUILLWORT FAMILY (ISO) Identification of the quillworts is a technical task requiring examina­tion of the mega­spores with a high-powered dissecting microscope and comparison material of correctly named specimens. Mature plants are needed, which means collecting in late summer or fall, when the spores in the leaf bases are ripe, and the plants then tend to fall apart. Young culms of grasses, sedges, and burreeds may easily be mistaken for quillworts if they are submerged by seasonal high water at the margins of ponds. ISOËTES L. 1753 [name used by Pliny for a species of Sedum]. Quillwort 1a. Megaspores with scattered low tubercles becoming joined to form low ridges. I. bolanderi Engelmann [for H. N. Bolander, 1831–1897, California botanist], 5D. Small lakes and ponds, upper montane and subalpine. 1b. Megaspores with high, sharp spines and jagged ridges . . . . . . . . . . . . . . . . . . . . . . (2) 2a. Leaves strictly erect, rather stiff; megaspores 0.5–0.6 mm diameter, with long low ridges and micro-ornamentation of short spines joined at bases. I. occidentalis L. F. Henderson. Deep water of Bierstadt Lake, Rocky Mountain National Park. The plants are often washed ashore by wave action. (I. lacustris of previous editions.) 2b. Leaves somewhat recurved, rather flexible; megaspores 0.2–0.5 mm diameter, spiny with spines of 2 distinct sizes. I. echinospora Durieu. Known from an old collection from Pikes Peak, and more recently from Bierstadt Lake in Rocky Mountain National Park. (I. setacea subsp. muricata of previous editions.)

LYCOPODIACEAE Palisot de Beauvois ex Mirbel in Lamarck & Mirbel 1802. CLUB-MOSS FAMILY (LYC)

The spores of Lycopodium were once used as a fine baby powder and an inflammable powder for flash photography. In Scandinavia, where they are abundant ground cover in forests, lycopods are gathered in enormous quantities for ornamental Christmas greens. In Colorado they are so infrequent as to be considered in need of protection. The FNA reports the circumpolar Diphasiastrum alpinum for Colorado. We have been unable to find any records, and its occurrence here is suspect. 1a. Stem not creeping, the erect branches tightly bunched; sporangia in the axils of unmodified leaves, not in discrete cones. Huperzia, Fir Club-moss 1b. Stem creeping extensively, the erect branches not tightly bunched; spores produced in an elongate cone. Lycopodium, Club-moss

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Ferns and Fern Allies

HUPERZIA Bernhardi 1801 [for Johann Peter Huperz, died 1816]. Fir Club-moss One species, H. haleakalae (Brackenridge) Holub [from Haleakala, Maui], 5B. Local in rocky cirque-basins and cliffs near or above timberline. (Lycopo­dium, Huperzia selago.) A separate family, Huperziaceae, may be used for this species and its relatives. LYCOPODIUM L. 1753 [Greek, lycos, wolf, + pous, paw]. Club-moss 1a. Leaves widely spreading or somewhat reflexed, rich green, 5–10 mm long, acute, regularly toothed; branches 10–15 mm wide. L. annotinum L. [a year old, from the marked separation of annual branches], 5A. Infrequent in subalpine spruce forests and under willow thickets. 1b. Leaves ascending, yellowish-green, 3–7 mm long, sparingly toothed or entire; branches 3–7 mm wide. L. dubium Zoëga. Rare, upper subalpine and alpine. This differs from L. annotinum in having very short, appressed leaves on short erect branches. There is no intergradation of these taxa in Colorado. The FNA calls our plant a trivial variant, var. alpestre C. Hartman, of L. annotinum. This is a northern circumpolar taxon and is not likely to be a mere environmental ecad. We follow Böcher, Holmen, & Jacobsen (1968) in their use of L. dubium. Johan Zoëga (1742–1788) was a Danish disciple of Linnaeus who wrote the Flora of Iceland (1775).

MARSILEACEAE Mirbel in Lamarck & Mirbel 1802. PEPPERWORT FAMILY (MSL) Plants of Marsilea are not very fern-like and might be passed off as four-leaf clovers, but the venation is dichotomous-parallel. In late summer the sporocarps, hard, nut-like organs at the base of the plants, are conspicuous. The sporocarps of Nardoo, M. drummondii, of Australia, are edible and gathered by the aborigines, who taught the early explorers to use this wild food as a last resort against starvation in the outback. MARSILEA L. 1753 [for Luigi Marsigli, 1658–1730, Italian naturalist]. Pepperwort One species, M. vestita Hooker & Greville, Hairy Pep­perwort, 5C. Edges of muddy streams and ponds at low altitudes on the plains and in piedmont valleys. The plants become conspicuous in August and September when ponds dry up. (M. mucronata of previous editions.)

ONOCLEACEAE Pichi Sermolli 1970. ONOCLEA FAMILY (ONC) ONOCLEA L. 1753 [name used by Dioscorides for some plant, certainly not this one] One species, O. sensibilis L. [sensitive (to early frosts)]. Sensitive Fern. Known in Colorado from one collection in the foothills of the Front Range near Sedalia. The population has disappeared. An Arcto-Tertiary relict, otherwise common in the eastern US and Asia.

OPHIOGLOSSACEAE C. Agardh 1822. MOONWORT FAMILY (OPH)

Contributed by Donald R. Farrar (Iowa State University) and Steve J. Popovich (United States Forest Service) The Ophioglossaceae is an ancient family of plants distantly related to modern ferns. Plants produce only a single leaf per year that is divided dichotomously into a sterile, foliaceous part (trophophore) and a fertile, spore-bearing part (sporophore). (For this treatment, the term “leaf ” is preferred to “frond,” which is used elsewhere in this book.) Worldwide, the Ophioglossaceae contains 10 genera. Three occur in Colorado, adjacent states, and southwestern South Dakota: Botrychium, the moonworts; Botrypus, the rattlesnakeferns; and Sceptridium, the grape-ferns. Traditionally the latter two groups have been included in Botrychium, but morphology, and to a greater extent genetics, provide convincing evidence for their separation into distinct genera (Hauk et al. 2003). The Ophioglossaceae has long held special fascination for those who have encountered it. The name “moonwort” reflects medieval beliefs of the plants’ mystical powers to render

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doors unlocked, horses unshod, and invisibility to anyone dusted with their spores. These diminutive plants continue to receive attention disproportionate to their size and diversity. Alice Eastwood was one of the first to collect Ophioglossaceae in Colorado, in 1887 from Pikes Peak and in 1891 near Steamboat Springs. Subsequent collections over the next century were sparse until the 1980s, when intensive search and discovery by Warren (Herb) and Florence Wagner resulted in numerous collections. Colorado’s first “moonwort aficionado,” Peter Root, inspired by the Wagners, continued to increase collection records. Further discoveries by Warren Hauk, Toby Spribille, Annette Kolb, Jim Montgomery, Scott (Scotty) Smith, David Steinmann, Erica Smith Sokoloski, Kevin Kovacs, and Ben Legler added many new records, and greatly expanded knowledge of the species and their distributions in Colorado and surrounding areas. Dedicated U.S. Forest Service botanists have made similar discoveries in the Black Hills of South Dakota and Wyoming. A revision of the Botrychium lunaria complex by Mary Stensvold (2008) has affected the taxonomy of several of our species. New taxa continue to be described, and important discoveries undoubtedly lie ahead as fieldwork and genetic analysis promote increased understanding of Ophioglossaceae species and their ecology. The coverage area for this treatment emphasizes Colorado, but includes all adjacent states (AZ, KS, NE, NM, OK, UT, and WY) and the southwestern quadrant of South Dakota. It also accommodates all of the Rocky Mountain Region (Region 2) of the U.S. Forest Service. The treatment excludes Ophioglossum and Sceptridium species that occur in our area only in the eastern deciduous woodlands of Nebraska, Kansas, and Oklahoma. Distribution and habitat descriptions are for Colorado unless noted otherwise, but in many cases apply reliably to adjacent states. Couplet drawings illustrate underlined text. The authors thank Karen Pille for couplet illustrations, Chanda Skelton for Diagram 1, and Cindy Johnson for critical review. 1a. Trophophore (foliaceous leaf part) simple, entire; veins reticulate; sporophore (spore-bearing leaf part) with sporangia sunken in axis. Ophioglossum, Adder’s tongue

1b. Trophophore compound (rarely simple, entire in Botrychium simplex); veins forked; sporophore with sporangia not sunken in axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . (2) 2a. Trophophore ternate or pinnate (rarely simple), less than 4.5 cm wide, no more than twice divided. Botrychium, Moonwort 2b. Trophophore ternate, generally more than 6 cm wide, 3 to 4 times divided (2 times divided in small plants) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 3a. Trophophore withering in the fall; trophophore and sporophore joined several cm or more above ground level; trophophore sessile; pinnae thin and delicate. Botrypus, Rattlesnake-Fern 3b. Trophophore green (or bronze, but still photosynthetic) through the summer of its second year; trophophore and sporophore joined ± at ground level; trophophore stalked; pinnae thick and leathery. Sceptridium, Grape-Fern BOTRYCHIUM Swartz 1800 [Greek, botry, bunch of grapes, from resemblance of the spor­ angia to clusters of grapes]. Moonwort Moonworts are the most diverse and abundant Ophioglossaceae in the Rocky Mountains, and 16 taxa occur in Colorado. The common name is derived from Latin luna, moon, in reference to the crescent-moon shape of the pinnae of the type species, B. lunaria, and Old English wyrt, herb, for alleged curative properties. Although a few moonwort species in our area have specialized habitat preferences, most occur on well-drained, seasonally mesic soils in sites dominated by perennial herbaceous vegetation. Moonworts have a close association with mycorrhizal fungi on which they depend for water, minerals, and carbohydrates. Botrychium distribution and abundance are linked to the

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Ferns and Fern Allies

co-occurrence of these fungi. Moonwort habitats are diverse, ranging from natural meadows and riparian areas to previously disturbed (now stabilized) areas, including burns, avalanche chutes, roadsides, mining districts, earthen dams, power lines, timber harvest clearings, old town sites, and ski slopes! Populations tend to appear about 20 years following disturbance, and subsequently decline with canopy closure by woody plants. Moonworts generally occur at elevations above 9000 ft in Colorado, predominantly in the subalpine, but also in montane and alpine areas. Although they occur on various soil types, Botrychium seem to be especially frequent on soils developed over limestone or other sources of calcium. A good, but not always reliable, indicator of moonwort habitat in Colorado’s subalpine is the presence of two or more of the following associates: Androsace, Antennaria, Fragaria, Senecio, Solidago, Taraxacum, and juvenile (knee- to head-high) conifers. While moonworts are difficult to find, recent fieldwork has revealed populations more abundant and widespread than previously believed. Some moonworts are markedly more prevalent than others, and 95 percent of the time, those encountered in Colorado are Botrychium echo, B. hesperium, B. lanceolatum (red phenotype), B. minganense, and B. “neolunaria.” The key is intended to identify typical specimens. Determining what is “typical” requires examination of a number of plants within a population. Very small and very large individuals are often atypical, and multiple species may co-occur in productive habitats. Moreover, some populations exhibit sterile hybrids not included in the key. Identification of Botrychium species requires careful attention to details of leaf structure (Diagram 1). The most important characters for identification are the shape and cutting (dissection) of the pinnae and the stalk lengths of the trophophore and sporophore. Trophophore stalk length ranges from none (sessile) to longer than the average of the distances between the attachment points of the first (basal) and second, and second and third, pinnae pairs. Sporophore stalk length ranges from one-fourth of the total length of the trophophore to longer than the total trophophore length. The sporophore stalk elongates last, and lengths provided in the key apply after sporangia have matured (yellow and plump prior to spore release, or brown and shriveled following spore release). Throughout the key, “sporophore stalk” refers to the length of the sporophore stalk only, whereas “total trophophore length” refers to the total length of the trophophore including its stalk. Pinna span is important in once-pinnate species and is measured by superimposing an arc of a circle with the point of pinna attachment as the central angle’s vertex (Diagram 1). Pinna span is measured in degrees, and unless stated otherwise, it and other pinna characters refer to the basal pair of pinnae. As with all technical keys, most descriptive characters are understood to be modified by the word “usually” or “generally,” and this is especially true for Ophioglossaceae. These terms have been retained in the leads only when particularly applicable. Underground structures are not useful for identification. If collection is necessary, carefully cut (do not pinch) the leaf at ground level. This preserves the underground stem and roots. Specimens must be carefully pressed to ensure that the sporophore and trophophore do not overlap and that pinnae margins are fully spread. Please refrain from over-collecting. Photographs that clearly show the features described above are helpful and may substitute for collections from small populations. In the geographic area covered by the key, Colorado has received the greatest survey effort, but all states remain incompletely surveyed. See the table following the key for species occurrences listed by state. Enjoy and don’t get discouraged! With practice, you will acquire a feel for keying moonworts and it will become easier as you gain familiarity with this interesting genus. 1a. Leaf divided into two sporophores, more or less equal in size and shape; trophophore absent. B. paradoxum W. H. Wagner [referring to the peculiar absence of a trophophore], 3-III. One small site, a subalpine grassy slope near Crested Butte,

Ferns and Fern Allies

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GN. In Colorado, individuals of other species have been observed to produce two sporophores and no trophophore, but this is rare and the sporophores usually differ in size and shape. 1b. Leaf divided into a sporophore and a trophophore . . . . . . . . . . . . . . . . . . . . . . . . . (2)

2a. Trophophore blade width (at the base) greater than or approximately equal to the blade length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (3) 2b. Trophophore blade longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (9)

3a. Sporophore absent or tiny (mostly