Steyermark's Flora of Missouri, Volume 1 [1]

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STEYERMARK’S

Flora of Missouri

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STEYERMARK’S

Flora of Missouri Revised edition

George Yatskievych

The Missouri Department of Conservation Jefferson City, Missouri

in cooperation with

The Missouri Botanical Garden Press St. Louis, Missouri

PRINTING HISTORY

First edition, published as “Flora of Missouri” by the Iowa State University Press, Ames, Iowa Published November 1963 Second printing (with errata), 1968 Third printing, 1972 Fourth printing, 1975 Fifth printing, 1977 Sixth printing, 1981 Seventh printing, 1996

Revised edition, published by the Missouri Department of Conservation, Jefferson City, Missouri, in cooperation with the Missouri Botanical Garden Press, St. Louis, Missouri.

Published December 1998

Copyright © 1998 by the Conservation Commission of the State of Missouri

ISBN: 1-887247-19-X

[Insert Cons. Dept. and Mo. Bot. Garden Logos?]

Missouri Department of Conservation P.O. Box 180 Jefferson City, MO 65102-0180

Missouri Botanical Garden P.O. Box 299 St. Louis, MO 63166-0299

Equal opportunity to participate in and benefit from programs of the Missouri Department of Conservation is available to all individuals without regard to their race, color, national origin, sex, age or disability. Complaints of discrimination should be sent to the Department of Conservation, P.O. Box 180, Jefferson City, MO 65102, or U.S. Fish & Wildlife Service, 18th and C streets NW, Washington D.C. 20240, Missouri Relay Center 1-800-735-2966 (TDD).

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DEDICATION

The first edition of this book was dedicated to the famous student of the Missouri flora, Ernest J. Palmer, who, unfortunately, did not live to see its completion. Similarly, the present edition is dedicated to the memory of Julian A. Steyermark, in the hope that he would have approved of its contents.

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CONTENTS

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix Illustration Credits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xii

Part I: Introduction How to Use This Book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 A History of Floristic Botany in Missouri. . . . . . . . . . . . . . . . . 7 An Overview of Missouri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 1. Geography, Climate, and Geology. . . . . . . . . . . . . . . . . . 43 2. Vegetation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Natural Divisions of Missouri . . . . . . . . . . . . . . . . . . . 52 Plant Communities. . . . . . . . . . . . . . . . . . . . . . . . . . . . 60 Analysis of the Flora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

Part II: The Flora Summary of the Major Groups . . . . . . . . . . . . . . . . . . . . . . . . 94 Pteridophytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 Gymnosperms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162 Monocots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174

Glossary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 915 Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 929 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 953

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ACKNOWLEDGMENTS A long-term project such as this would not be possible without the assistance of countless individuals whose efforts, large and small, helped lead to its completion. As compiler of the information in these pages, however, responsibility for the content rests solely with the author, and any weaknesses in the treatments may be attributed to me. Because the work is ongoing, this list necessarily is incomplete, and I apologize for any names that my failing memory may have caused to be omitted. In the beginning, it was Peter Raven whose vision provided the impetus for a revision of Missouri’s flora that developed with his continuing support into the present project. Others at the Missouri Botanical Garden directly involved in the project’s administration have included Marshall Crosby, Enrique Forero, Bob Magill, Nancy Morin, Mike Olson, and Doug Stevens. John Wylie was the project’s first champion at the Missouri Department of Conservation, and his efforts have been followed by several able administrators there, including James H. Wilson, Rick Thom, Don Kurz, David Urich, Al Brohn, and Ken Babcock. The Missouri Conservation Commission is to be commended for its support of floristic programs in the state. My project assistants, interns, and volunteers have worked thousands of hours at jobs ranging from routine to very unusual and challenging, and all have performed above and beyond the call of duty. They have included: Alan Brant, Mary Broida, Karen Bynum, Germán Carnevali, Charles Caverly, Linda Ellis, Barbara Fry, Brad Guhr, Claire Hemingway, Paige McClure, Bill Summers, Ba¯a¯di Tadych, Charlotte Taylor, Joanna Turner, Matthew Wagner, Pat Walker, Jackie Wallach, Andoni Westerhaus, Kathleen Wood, and Jane Whitehill. In particular, the work could not have been completed without the tireless efforts of Bill Summers and Joanna Turner, who have been helping me since nearly the beginning of the work in 1987. Information in the introductory chapters was compiled from numerous sources. In particular, I am indebted to the following individuals, who provided information or graphics: Nicki Aberle, Natural History Biologist, Missouri Department of Conservation; Chris Barnett and Chris Fulcher, University of Missouri Center for Agricultural, Resource, and Environmental Systems; Charles Brown, Manager of Information Services, St. Louis Mercantile Library; Chris Dietrich, Natural History Biologist, Missouri Department of Conservation; Susan Fraser, Archivist, New York Botanical Garden; Betty (Bauer) Hedges, Kirkwood; Albert and Dorothy Heinze, Naturalists, Imperial; Thomas P. Hertel, Data Processing Specialist, Division of Geology and Land Survey, Missouri Department of Natural Resources; Blane Heumann, Head of the Lower Ozarks Field Office, The Nature Conservancy, Missouri Chapter; Anita L. Karg, Archivist, Hunt Institute for Botanical Documentation, Carnegie Mellon University; Robin Kennedy, Curator, University of Missouri Herbarium; Pamela Knobeloch, Ethical Society of St. Louis; Martin Knorr, Librarian, Harris Stowe State College; Doug Ladd, Director of Science and Stewardship, The Nature Conservancy, Missouri Chapter; Marshall Magner, Naturalist, Webster Groves; Betty Nellums, Webster Groves Nature Study Society Historian; Frank Mercer, Professor and Dean, Emeritus, St. Louis College of Pharmacy; Martha Riley, former Archivist, Missouri Botanical Garden; Tim Smith, Botanist, Missouri Department of Conservation; Rick Thom, Natural History Section Chief, Missouri Department of Conservation; and Cora J. Wilson, Archivist, Jackson County Historical Society. Information on the artists and funding sources for the plates of species illustrations are listed separately. A number of specialists have helped with the completion of various plant families for Volume 1. It is a pleasure to acknowledge their assistance. The following are among the botanists who provided data and/or reviewed treatments: Susan Aiken, Canadian Museum ix

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of Nature (Festuca); Charles Bryson, U.S. Department of Agriculture (Carex); Julian Campbell, The Nature Conservancy, Kentucky Chapter (Elymus); Richard Carter, Valdosta State University (Cyperus, Eleocharis); David Castaner, Central Missouri State University (Carex); Gerrit Davidse, Missouri Botanical Garden (Cyperaceae, Poaceae); Jerrold Davis, Cornell University (Puccinellia, Torreyochloa); Robert Faden, U.S. National Herbarium (Commelinaceae); Roy Gereau, Missouri Botanical Garden (Poaceae); Charles Horn, Newberry College (Pontederiaceae); Terry Jacobsen, Hunt Institute (Allium); Robert Kral, Vanderbilt University (Fuirena, Xyris); Clair Kucera, University of Missouri (Poaceae); Ronald L. McGregor, University of Kansas (Sporobolus); Paul McKenzie, U.S. Fish and Wildlife Service (Cyperaceae, Poaceae); Robert Naczi, Western Kentucky State University (Carex); John and Charlotte Reeder, University of Arizona (Poaceae); Jeff Rettig, College of the Ozarks (Carex); Anton Reznicek, University of Michigan (Carex); John Ricketson, Missouri Botanical Garden (aquatics); Terry Robison, Missouri Department of Conservation (Pinus); S. Galen Smith, University of Wisconsin at Whitewater (Eleocharis, Scirpus and allies, Typhaceae); W. Carl Taylor, Milwaukee Public Museum (Pteridophyta); Sue Thompson, Carnegie Museum (Acoraceae, Araceae); and Gordon C. Tucker, Eastern Illinois University, Charleston (Cyperaceae). These days, computers are the lifeblood (and bane) of any scholarly undertaking. Fortunately, I have had lots of help with the cybernetic aspects of the project. Rick Wolters and Todd Larivee (Missouri Department of Conservation) and Jennifer Reis (Missouri Botanical Garden) have been my principal advisors in matters of hardware and software. Jeffrey Kunce of the Conservation Department designed a very useful program for mapping countylevel distributions of plants and Deborah Briedwell produced the final maps in ArcInfo from my database. Chris McMahon and Miriam Fica of the Garden also continue to provide excellent support, especially with network, Internet, and World Wide Web–related problems. The raw materials upon which this revision is based originated in the libraries, the herbaria, and the landscape of Missouri. I am greatly indebted to the staff of the Missouri Botanical Garden’s library, whose knowledge of their substantial accessions is truly impressive. Dale Johnson and Linda Oestry were especially helpful in helping to locate obscure references. I also owe a great debt to the curators of the herbaria consulted both within Missouri and elsewhere in the United States for their courtesy in allowing me to examine and borrow specimens that were studied during the preparation of Volume 1. At the Missouri Botanical Garden Herbarium, David Brunner, Jim Solomon, Sue Taylor, and especially Lois Ganss were a big help in making my research possible. The process of turning a manuscript into a published volume is lengthy and involved. I am indebted to my editor, Julie Schroeder, for her attention to detail, and to Lisa Skokowski, who completed the layout and design, for rising to a difficult challenge. At the Garden, Amy McPherson and Diana Gunter provided invaluable technical assistance on matters of production, as did Carol Davit and Bernadette Dryden at the Conservation Department. Since the start of the project in 1987, collectors have added about 5,000 new Missouri specimens to the Missouri Botanical Garden herbarium annually. When one adds to this the new accessions at other herbaria around the state, it becomes apparent that a large number of amateur and professional botanists have generated valuable new information on the state’s flora, including numerous distributional records. This lively interest, along with my interactions with these botanists, provided a strong impetus to see the work through to completion. In particular, the questions, discussions, and continual prodding by members of the Missouri Native Plant Society and the Botany Group of the Webster Groves Nature Study Society, provided a kind of positive support that kept me going at times when the project seemed a bit overwhelming. Amateur botanists of special note have included Carl and Dolly Darigo, Patrick Delozier, Karen Haller, Jack and Pat Harris, Stanton Hudson, John

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Molyneaux, Ann Wakeman, and the late Arthur Christ and Edgar Denison. A number of present and former biologists with the Missouri Department of Conservation have been particularly helpful with new records of uncommon species, including Craig Anderson, Nina Bicknese, Rick Clawson, Greg Gremaud, Sherry Holmes, Brad Jacobs, Karen Kramer, Hillary Loring, David Moore, Tom Nagel, Doug Newman, Tim Nigh, Deborah Rowan, Joe Ryan, Mike Skinner, and especially Tim Smith. Similarly, a number of naturalists and biologists at the Missouri Department of Natural Resources have provided new records, including Michael Currier, Dan Drees, Ken McCarty, Ron Mullikin, Paul Nelson, and Bruce Schuette. Paul McKenzie of the U.S. Fish and Wildlife Service’s Columbia Field Office has been a tireless student of the Missouri flora. Similarly, Doug Ladd and his associates at the Missouri Chapter of The Nature Conservancy have been very helpful both in providing new records and in discussions concerning vegetation and flora. No list of thank-yous could be complete without an acknowledgment of the late Julian Steyermark’s role in the project. It was with his permission that this revision became possible, and his support at the beginning was invaluable in getting me started. Julian Steyermark is a model of what a botanist should be, and his high standards of scholarship are something I continue to aspire to but fear I will never reach. Would that I had known while he was still living the questions that I would have liked to ask him now. First and last, my wife, Kay, is responsible for this volume reaching completion. In addition to her prodigious knowledge of midwestern flora, her love and support carried me through all phases of the work.

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ILLUSTRATION CREDITS The revised edition features entirely new plates and includes considerably more species than were in Steyermark’s first edition. These illustrations are a major contribution to the manual and are an invaluable resource for both learning about various species and verifying specimen determinations. The 586 new plates of line drawings that appear in the Flora of Missouri would not have been possible without the generous contributions from the following funding sources: The Academy of Science of St. Louis Timon Primm Memorial Fund (at the Missouri Botanical Garden) Mrs. John S. Lehmann The John Allan Love Foundation The Missouri Department of Conservation The Wednesday Club of St. Louis

The following artists applied their considerable talents to the completion of the 194 plates that appear in Volume 1. Their dedication and patience with the author’s often detailed instructions are greatly appreciated. Phyllis Bick

Plates 1–3, 6–19, 22–25, 67–87, 110–18, 131–91

Linda Ellis

Plates 97, 98

Kate Johnson

Plates 193, 194

Lewis Johnson

Plate 89

Ellen Lissant

Plates 20, 21, 119–30

John Manning

Plates 27, 28, 88, 90–92, 99–109, 192

Jean Meske

Plates 4, 5

Paul Nelson

Plates 26, 29–66

Paula Nicholas

Plates 93–96

Many materials on Plates 29–66 were cofunded and prepared in conjunction with a forthcoming volume on the genus Carex in Robert H. Mohlenbrock’s Illustrated Flora of Illinois series. They are reproduced here courtesy of Southern Illinois University Press. The plates in the Missouri and Illinois manuals contain somewhat different materials and are reproduced in different formats with different magnifications. A number of sources generously provided materials used or adapted in the figures included in the introductory chapters. They are credited individually in the figure captions. Other photographs and figures, unless noted, are the author’s.

PART I

Introduction

HOW TO USE THIS BOOK This floristic manual is intended to serve several purposes, depending upon the needs of the user. At its heart, it is an encyclopedia of the plants to be found growing outside of cultivation in the state. To this end, it includes information on the appearance, flowering time, distribution, and habitats of each species, along with discussions of plant uses and other pertinent topics. The introductory chapters include background information on the history of botany, climate, geography, geology, and vegetation of Missouri. Perhaps of greatest importance to serious students of the flora is the lengthy bibliography of books and papers cited in the text. However, a second group of users is interested primarily in a manual for the determination of unknown plants. To this end the descriptions and other text are integrated with a series of illustrations, distribution maps, and diagnostic keys, which are intended to facilitate identification of all of the vascular plants growing in Missouri. Other user groups, particularly in the professional community, may see the book as a tool for comparing Missouri’s flora to that of surrounding states; a set of baseline, organismal data as an aid to developing land management strategies; a summary of the native, introduced, weedy, endangered, or other classes of plants to correlate with other studies, or simply as a starting point for further research on topics in need of closer study. Whatever the background of the user, the intent of Julian Steyermark (as well as the present author) has been to provide scientifically accurate information with sufficient detail to be useful without too technical a style of presentation. Technical terminology has been kept to a minimum (at the slight expense of verbal precision in some cases), and most of the specialized terms found necessary to communicate various characteristics of plants are defined in the text and/or the glossary at the end of each volume. An effort has been made to standardize terms as much as possible, both for plant parts and for ecological information. In this way, continued use of the volume (hopefully) will result in increased user proficiency. When asked, nearly all botanists will answer that the way to become familiar with the flora of an area is to practice determining unknowns using a floristic manual. The prerequisites for using this volume are few, but important. First and foremost, the user must be able to read carefully, as utilizing both the diagnostic keys and the descriptions requires attention to detail. The user also must be familiar with at least some basic botanical terms, such as can be found in even the nontechnical literature, as well as the metric system for linear measurements (a ruler is included on the back endpaper of the volume). For many groups of plants, details necessary for plant determinations can be seen only with magnification. For this purpose, users are encouraged to purchase a good-quality hand lens, preferably of 10¥ magnification or better. A good hand lens is both one with clear optics and one with which the user feels comfortable, and readers are urged to try out various models before making a choice. A dissecting microscope conveys a wonderful advantage for plant identification as it leaves the hands free for manipulation and dissection of specimens, but is not a requirement to use this book. Other tools that may prove useful for plant determinations are a pair of fine-point forceps, a razor blade or sharp knife, and a dissecting probe or needle.

TAXONOMIC FRAMEWORK The desire to want names for objects is both human nature and a necessity for communication. People also like to “pigeonhole” objects by relating them to other, similar objects, which perhaps grew from a need to feel secure about the potential properties of unknowns through analogy with known objects of similar appearance. These drives to name things and 1

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to classify them were the psychological roots of the earliest efforts at taxonomy. A taxon (plural, taxa) is an entity in a system of classification, regardless of which level (family, genus, etc.) is being discussed, and taxonomy is the organized study of taxa. Systematic botany, the study of plant classification, has come to mean about the same thing as plant taxonomy in most of the botanical literature. In biology there is a nested hierarchy of levels at which objects are classified. This hierarchy is utilized in the present volume. Within the plant kingdom (one of several kingdoms, including animals, fungi, bacteria, etc.), there are a number of major groups known as phyla or divisions. Division names end in -phyta; for example, the division of flowering plants is known scientifically as the Magnoliophyta in the modern literature. Within each division, there may be one or more classes, whose scientific names end in -opsida. For example, the monocots are the Liliopsida, whereas the dicots are the Magnoliopsida. Each class contains one or more families of organisms, whose names usually end in aceae. Families are treated alphabetically within each of the major groups in the present volume. They are an important level in the classification, as many botanists find it useful to learn to recognize important plant families, such as the Poaceae (grasses), Cyperaceae (sedges), Fabaceae (legumes), Asteraceae (composites), and so forth. Edgar Denison (1994), in the fourth edition of his popular field guide, Missouri Wildflowers, noted that nearly 80 percent of Missouri wildflowers are contained in only 24 of the more than 100 families in the flora, and that familiarity with these 24 common families can be very helpful in facilitating plant determinations. Many field botany classes at colleges and universities focus on teaching characters to recognize plant families. Families contain one or more genera (singular, genus), also treated alphabetically in this work (as are the lower levels of classification). There is no consistent ending for generic spellings as there are for higher taxonomic levels. Each genus contains one or more species. Species are the basic unit of all taxonomic classifications, and most people, regardless of background, have some intuitive grasp of species in nature. However, the way that taxonomists decide how clusters of related species should be grouped into genera is one of the most perplexing problems for nonbotanists to understand. In addition, telling apart species in some difficult groups can cause headaches for even experienced botanists. Within a species, not all individuals look exactly the same. This variation between individuals, or between populations of individuals, is due to a combination of environmental and genetic causes. Environmental causes are generally ignored for taxonomic purposes, because the particular appearance of a plant under environmental control cannot be passed on to that plant’s offspring (seeds from a stunted plant growing in the dry, nutrient-poor crack in a sidewalk will grow into plants of very different appearance if sown in the rich, moist soil of a garden plot). However, genetically based variation can be important for classification purposes. Minor variations in a single character, such as flower color mutants or nearly glabrous individuals in a population of otherwise hairy plants, have been named as forms by some botanists. Steyermark (1963) was a great proponent of recognizing forms, which can have horticultural importance, but in general such minor variants are not considered important by most botanists. Thus, not much space is devoted to discussion of forms in the present volume. In some cases, however, more complex variation allows a species to be divided into more or less coherent groups, which often differ somewhat in geographic range, habitat preferences, as well as appearance. These variants are often classified as varieties or subspecies, depending upon the preferences of the particular taxonomist. They are components of a species that have differentiated somewhat, but not enough to be separated as different species, and can nearly always hybridize freely with fertile offspring. On the other hand, species theoretically are stable biological entities that are separated from other related species by some sort of boundary or boundaries (several other definitions

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of the word species exist, which will not be discussed here). Usually these boundaries are genetic; that is, two related species may not be able to interbreed successfully for one or more reasons, producing only sterile offspring. In other cases, species may be isolated reproductively by different flowering times or specialized pollinators that prevent hybridization from occurring in nature. In some cases, nonoverlapping distributions or different habitat requirements may prevent interbreeding. However, even in cases where the potential for rampant hybridization is present (as in some wind-pollinated plants, such as oaks and willows), some mechanism exists to insure that the species maintain their individuality over successive generations. In cases where hybridization with fertile offspring becomes excessive, related taxa may be “swamped” genetically and unable to maintain their distinctions. This apparently has occurred in the sugar maple alliance of the genus Acer, where several groups (often treated as separate species) became separated geographically at some point in the past (perhaps during Pleistocene glaciation) and diverged morphologically but were forced back together by subsequent climate changes and recent alterations in land management by humans, to the point where separate species cannot be recognized over most of the cumulative range of the complex. To determine specimens, a belief in evolution is not necessary. However, the biologists who name and classify organisms use evolution as a basic framework for classifying species (or other levels in the hierarchy). The explanation that two species are related to one another not only because they share similar morphological features, but because they share a common ancestry (phylogeny), is essential to an understanding of taxonomy. Most modern systematists seek to understand the phylogeny of groups of taxa, that is to say the evolutionary and genealogical relationships among the organisms under study. Species differentiate over time by fragmentation of populations into different geographic areas; spread of genetic mutations through some populations, but not others; the effects of environmental or other selection for survival of some individuals and not others (“survival of the fittest”); and random chance affecting the matrix of genetic variation within and between populations. An explanation of evolutionary theory is beyond the scope of the present volume (several excellent texts are available at most libraries), but the idea that taxonomists name species and adopt classifications for other reasons in addition to morphological differences helps to explain why botanists sometimes persist in maintaining species that may be difficult to differentiate in the field. Ideally, a taxonomic classification should both express phylogenetic relationships and be of practical use for plant determinations, but organisms exist and evolve without foreknowledge that humans are trying to identify them. On a practical level, the classification of plant families used in the present volume is adapted with only minor alterations from those of Cronquist (1981) and Gleason and Cronquist (1991). The genera and species accepted are derived from the author’s understanding of the taxonomic literature and the wealth of specimens of Missouri plants. Where a given taxon has been called by a different name or recognized at a different taxonomic level in earlier literature on the Missouri flora, these synonyms are listed under the accepted name. Each species name consists of two parts, the genus name and the species epithet, known collectively as a binomial. The citation of authorities for various names is a bibliographic tool primarily of interest to taxonomists, but it also gives recognition to the authorities who have studied various taxa. The abbreviations for authors of plant names generally have been standardized following Brummitt and Powell (1992). The convention for plant names is that the one or more authors whose names appear in parentheses were the ones who first coined the particular epithet in question, whereas the one or more names immediately outside the parentheses denote the authors who adopted the accepted classification for that taxon. For example, the Ozark wake robin is presently known scientifically as Trillium pusillum Michx. var. ozarkanum (E.J.

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Palmer & Steyerm.) Steyerm. André Michaux, the great French botanist who explored portions of North America in the early 1800s, named this species based on specimens collected to the east of Missouri. The varietal epithet “ozarkanum” was coined by Ernest J. Palmer and Julian Steyermark, two of Missouri’s most famous botanists (see the history of botany chapter later in this volume), but they originally described it as a separate species, Trillium ozarkanum E.J. Palmer & Steyerm. Later, Steyermark completed more detailed studies on the taxonomy of this group and decided that the plants of the Ozarks were not a separate species from those growing in the Appalachians, and he transferred Trillium ozarkanum to a variety of Trillium pusillum, which was the older (first published) of the two names at the species level and thus had nomenclatural priority.

DETERMINING SPECIMENS When confronted with a specimen of an unknown species, there may be a temptation to flip through the illustrations in the hope of “getting lucky” and finding a match. Although this can be useful in some situations where the genus or species group is known, in general the practice should be avoided. One reason for this is that not all species are illustrated (even though most are), so the plates provide an incomplete record of the flora. The plates also are incomplete in that not all parts of every species are shown—for many species only diagnostic details have been illustrated, rather than the entire plant or even the whole inflorescence. Finally, a single plate cannot convey the range of variation present in any given species, so a specimen may not match the corresponding illustration exactly. The first step in determination of an unknown plant is the use of the diagnostic keys. Keys are a method of breaking down large amounts of complex information on plants into smaller, manageable units known as couplets. In the present volume there are keys at several hierarchical levels, including families, which contain genera, which are composed of species, some of which are further divided into varieties or subspecies. Users with some botanical experience may be able to skip the key to families and start with the key to genera for a known family or the key to species if the genus has been recognized. Whatever the level in the taxonomic hierarchy, the key consists of one or more couplets, which are organized in an indented form similar to that of a written outline. A couplet is a pair of contrasting statements (leads) that require the user to choose which set of character states (variations for that character) most closely approximates those of the specimen at hand. After deciding on the most correct lead, the user moves on to the next couplet, indented under that lead. Successive couplets eventually lead to the plant name. In some cases, when the plant under consideration seems intermediate between the two leads of a given couplet or the couplet asks about structures not present on the specimen, the user will have to go both ways at that juncture and decide which of the two paths leads to a better end. Often, in such cases, the incorrect initial lead will be followed by couplets in which neither lead makes sense or fits in the context of the specimen. Keying plants can be a challenging activity, particularly for large genera or morphologically similar groups. For this reason, users should verify their determinations by comparing the specimen to the written description of the taxon, as well as the illustrations. Comparison of unknowns with authenticated specimens at one of the state’s many herbaria (see the history section later in this volume) is especially useful in confirming identifications. In some cases, the range maps and habitat data also may provide information useful in making the decision on whether the determination is correct.

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TYPES OF INFORMATION INCLUDED The descriptions vary in length from group to group. Within a group, the descriptions are organized roughly in parallel fashion; that is, the same characters are discussed in the same order for each taxon, to facilitate comparisons. An important point is that the descriptions are hierarchical. For example, a character that is constant for all species in a given genus is included in the generic description, but not repeated under each species, to save space. Thus users should carefully read the generic descriptions in addition to those for individual species. Also, for genera in which only a single species occurs in Missouri, only the species description is given, without a separate one for the genus, again to save space. The descriptions contain characters in the same order: whole plant, underground structures, aboveground stems, leaves, inflorescences, flowers, perianth (sepals, petals), stamens, carpels, and fruits. They end with a listing of any chromosome numbers reported for the species, taken without verification or citation from the literature. There is also a range of months of flowering given, based on the specimen record and data from Steyermark (1963). Other information under each species includes: common name(s); a short summary of Missouri and overall ranges; a list of habitats in which the species occurs; discussions of taxonomy, biology, and/or other pertinent topics; citations of plate numbers and county dot map numbers; and a treatment of any subspecies or varieties in the flora. For information on habitats of Missouri plants, see the discussion of this topic in a later introductory chapter. Illustrations were prepared from a combination of fresh plants and dried specimens. Although nearly all of the species are illustrated, not all parts of every species are shown. Instead, for many species groups, whole plants or inflorescences are shown for only some species, with details important for determinations, such as fruits or flowers, illustrated for the remainder. Exotic species are indicated with the word introduced at the beginning of the paragraph on distributions and ecology. Native species, which form the bulk of the flora, are not indicated as such. In general, an introduced species is one that probably was not present in Missouri prior to European colonization of the state a little more than 200 years ago. However, there are species, such as Lycopodium digitatum, Woodwardia areolata, and some species of Eupatorium, that apparently have expanded their natural ranges into Missouri during the past two centuries and must be considered native to the flora. Nonnative species may have been introduced in a variety of ways (escapes from cultivation, accidental introductions from overseas, or accidental spread of species native elsewhere in the United States) and range from widespread, noxious weeds to single records that may not have persisted in the state for very many years. Classifying a plant as introduced or native can be a controversial topic for some species, and each decision on this represents the author’s best guess, based on personal experience and the literature. Perceptions of native vs. nonnative also change; for example, many earlier authors, including Steyermark (1963), thought that watercress (Nasturtium officinale) was native in the midwestern states, but it is now generally accepted to have been introduced from Europe. Once noted and collected in Missouri by some individual, the general criteria for including any new species in the flora are the ability of that species to persist in the wild and its ability to reproduce and potentially to spread. The coverage in the manual probably is slightly overinclusive, as the decision was made to provide treatments of species for which information on reproductive success presently is unknown, including some of the introduced species collected along railroad tracks once or only a few times. The paragraph on distribution and ecology also includes a qualitative assessment of how abundant a species is in the state. This has been simplified to three states: uncommon, scattered, and common. Common species may be expected to be encountered regularly within the portions of the state indicated and to occur in two-thirds of the counties or more. Uncommon species are those mostly present in five counties or fewer or with very few occurrences with-

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in any of the Missouri counties indicated. Scattered species are those intermediate in abundance between common and uncommon. No attempt has been made to indicate those species considered rare or endangered at the state level by the Missouri Department of Conservation’s Natural Heritage Program (the few species listed officially under the Federal Endangered Species Act are, however, indicated as such), because the information on rare Missouri plants is too incomplete for such a listing to remain up-to-date for more than a few months, and because the definitions and contents of the state list recently have undergone detailed revision. Users interested in the state-level status of plants should order the latest edition of the official list of species of conservation concern from the Missouri Natural Heritage Program (1998). The distribution and ecology paragraph contains a brief summary of the general distribution of each species in the state. This may include a simple summary of parts of the state in which a species occurs or may include the names of specific Natural Divisions to which it is restricted. For some species the range stated refers to the Missouri River, which is a convenient geographic marker for the southern limits of glaciation in the state. The introduction of this book contains discussions of the state’s Natural Divisions and the effects of glaciation, as well as the range of habitat types. The information on each plant’s ecology was taken mostly from the specimen record, supplemented in some cases with data from the literature. An attempt has been made to standardize the terms for various habitats in order to facilitate comparisons between species. The county dot maps were prepared from a computerized database of label data from herbarium specimens. Thus, every dot represents one or more voucher specimens that verify the occurrence of the species in that county. Information from the database presently is available from the author, and eventually will become available on the World Wide Web following publication of both volumes of the new edition. For purposes of convenience, the maps do not show a separation in distributions between St. Louis County and the City of St. Louis, which are separate political entities, but difficult to separate geographically at the small size of the present maps. It should be noted that the dot maps cannot convey how common a species is within a given county, or whether it is still extant there. A dot on a map only signifies that a species was collected in that county at least once at some point in the past. For most counties, no complete listing is available of the present-day, extant flora, although this information would be very useful to land managers, botanists, and conservationists. Users with access to botanical libraries are encouraged to take advantage of the abundant citation of references throughout the text. Although space limitations do not permit an exhaustive listing of the literature pertinent to every species, the references cited include the recent taxonomic literature as well as books and papers on various historical, biological, or other topics. These will serve as a starting point for more in-depth studies on groups of plants. By the same token, groups for which there are no references cited frequently are those groups in greatest need of taxonomic research. The present volume represents the first attempt at a comprehensive study of the Missouri flora in more than 35 years. However, as noted repeatedly throughout the volume, knowledge of the state’s flora remains woefully incomplete. It is hoped that publication of this revised edition will stimulate both amateur and professional botanists to seek out new distributional records in the field. Portions of the northeastern and southeastern quarters, as well as all of the counties bordering other states, are in the greatest need of further collecting and fieldwork. For those interested, but inexperienced, in preparing plant specimens, instructions may be obtained from any textbook on plant taxonomy or by contacting the author. I would also be happy to receive any corrections or suggestions of improvements for the future.

A HISTORY OF FLORISTIC BOTANY IN MISSOURI Prior to colonization of North America by Europeans, Native Americans already had developed a rich storehouse of information on plants. Without a written language to record data, native plant uses were preserved through an oral tradition that was to be impacted greatly during the upheavals of European conquest of America. Some information was transmitted to fur traders, missionaries, and other early settlers, but much of what has been written about Native American plant lore has been reconstructed by ethnobotanists, ethnologists, and archaeologists (for example, see Moerman [1977, 1986], for information on medical ethnobotany). The European presence in what is now the state of Missouri can be traced back to the sixteenth-century expedition of the Spanish conquistador, Hernando DeSoto (1499?–1542), who discovered the Mississippi River and traveled through portions of southern Missouri in 1541. Several French missionaries and explorers made casual observations of the landscape during travels in the seventeenth and eighteenth centuries (Beilmann and Brenner, 1951). However, the botanical exploration of the state did not begin until early in the nineteenth century. Some authors (Spaulding, 1908–1909) have cited the famous French botanist, André Michaux (1746–1802; commemorated by Croton michauxii G.L. Webster and Quercus michauxii Nutt., among others), as being the first botanist in Missouri, but although Michaux’s extensive travels took him in 1795 to Kaskaskia and Cahokia, Illinois, there is no evidence that he botanized west of the Mississippi River (Klem, 1914; McKelvey, 1955).

THE AGE OF EXPEDITIONS The earliest phase of botanical studies in Missouri involved expeditions that set out from the St. Louis area and primarily were intended to explore the newly opened American West, following the acquisition of lands by the United States through the Louisiana Purchase of 1803. The first such expedition to make botanical observations and collections was that of Meriwether Lewis (1774–1809) and William Clark (1770–1838) from May 1804 to September 1806, which involved a search for a route to the Pacific Ocean by boat. Then-president Thomas Jefferson, who had a keen interest in natural history, commissioned the expedition to make scientific observations and to collect seeds and botanical specimens. André Michaux apparently was under consideration as botanist on the trip, but political tensions between France and the United States prevented his going. Thus, the expedition had no trained biologist. Nevertheless, the journals of the trip, which have been published in various forms (see Moulton [1983–1997] for the most complete, annotated edition), contain the first observations of the vegetation and botany of the Missouri River valley and adjacent areas. The group traveled from St. Charles up the Missouri River before passing on to Iowa and eventually the Pacific Northwest. Of the about 150 specimens that have survived from the journey, which are housed in the herbarium of the Academy of Natural Sciences of Philadelphia, apparently none was collected in Missouri (Meehan, 1898). Likewise, it does not seem probable that seeds from the trip, which eventually made their way into the hands of several prominent nurserymen of the time (and perhaps also to Jefferson), originated from the Missouri portion of the trip. The first two botanists to collect specimens in Missouri were John Bradbury and Thomas Nuttall, both British by birth. These two traveled and botanized together on a remarkable journey into the western United States, but their subsequent careers took dramatically different turns. Bradbury died in relative obscurity, whereas Nuttall became regarded as one of the preeminent botanists of his time.

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John Bradbury (1768–1823; commemorated by Monarda bradburiana L.C. Beck and the genus Bradburia Torr. & A. Gray, among others) was commissioned by the Liverpool Philosophical Society to collect specimens and seeds of American plants for propagation and study at the Liverpool Botanic Garden in England, and arrived in St. Louis at the end of December 1809 (McKelvey, 1955). He botanized around St. Louis during the following year, before making arrangements to accompany the fur-trading expedition of William Price Hunt (1783–1842), who was attempting to retrace the route of Lewis and Clark to the Pacific Northwest for John Jacob Astor’s Pacific Fur Company. Bradbury’s travels lasted from March to July Fig. 1. Thomas Nuttall, from an oil painting by R. 1811 and took him as far west as North Dakota. A. Clifford, dated 1859. Photograph courtesy of the He was accompanied by Thomas Nuttall, and Missouri Botanical Garden Library Archives, St. he also often communicated with Henry Marie Louis. Brackenridge (1786–1871), a friend who was traveling along somewhat the same route with a competing expedition under the command of Manuel Lisa (1772–1820) of the Missouri Fur Company. Brackenridge made no collections, but the observations of Missouri vegetation in his accounts of his trip (Brackenridge, 1814, 1816) add to the meager storehouse of information from that early period. Bradbury’s journals originally were published in 1817 and have been reprinted and annotated several times since then. After returning to St. Louis, Bradbury made his way to New Orleans, witnessing the destruction of New Madrid during the great earthquake of December 1811, but he was prevented from completing his return to England until 1817 by the outbreak of the War of 1812 and subsequently by lack of funds. In the interim, his plant specimens, which had preceded him to Europe, were studied and described without his permission by the unscrupulous botanist, Frederic Pursh (1774–1820; commemorated by Phacelia purshii Buckl., among others). These factors prevented the development of Bradbury’s scientific career, and he made no further contributions to botany. After publication of his journals in 1817, he returned to the United States and lived first in St. Louis and then Middleton, Kentucky, until his death in 1823 (Rickett, 1934). Bradbury’s specimens, which apparently were dispersed by his son (McKelvey, 1955), were subjected to a convoluted series of changes of ownership, with the primary sets eventually coming to reside at the Academy of Natural Sciences of Philadelphia and the Public Museum of Liverpool. Thomas Nuttall (1786–1859; commemorated by Sedum nuttallianum Raf., among others) (Fig. 1) traveled extensively in the United States and completed three major expeditions west of the Mississippi River: the 1811 trip with Bradbury; an 1818–1820 southern trip through present-day Arkansas and Oklahoma, which only incidentally included stops near New Madrid, Missouri (Lottinville, 1980); and an 1834–1836 journey that began as another retracing of Lewis and Clark’s route, but wound up including lengthy side trips to Hawaii and the California coast. On his first trip west, he arrived in St. Louis during the autumn of 1810 (Pennell, 1936), with financial support from the eminent professor of botany at the University of Pennsylvania, Benjamin Smith Barton (1766–1815; commemorated by the genus Bartonia Muhl. ex Willd., among others). He had been instructed to conduct botanical explorations from Philadelphia to the Great Lakes and westward into present-day Saskatchewan, returning the following year via a southern course along the Missouri River (Pennell, 1936; McKelvey, 1955), but somehow he came to deviate significantly from these orders.

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Nuttall became acquainted with Bradbury and together the two botanized in the environs of St. Louis during the winter and spring of 1810–1811. Nuttall was able to join Bradbury on the 1811 Astorian expedition to retrace Lewis and Clark’s route to the Pacific Northwest, making his own collections. After returning to St. Louis, he made his way to New Orleans on a faster boat than Bradbury, thus missing the New Madrid earthquake (Lottinville, 1980). Unlike Bradbury, he was able to travel to England prior to the start of the War of 1812, and thus was able to save the bulk of his collections from the attentions of the piratical Pursh. Plants collected on this first expedition are accessioned primarily at the Academy of Natural Sciences of Philadelphia and the Royal Botanic Gardens, Kew, England. In Europe, Nuttall’s botanical fortunes flourished, and he became highly regarded professionally. He returned to the United States in late 1814 or early 1815 and completed further botanical studies at the Philadelphia Academy, eventually publishing a number of important books and papers on American plants, including his catalog of North American plants known up to 1817 (Nuttall, 1818). In 1823, Nuttall received an appointment as lecturer and curator at Harvard University, which he resigned in 1833 to undertake the third of his western trips (McKelvey, 1955), described later in this chapter. After this journey, he spent several years mostly at the Philadelphia Academy and at Harvard, before returning to England in 1842. Except for a short return to New England from 1847 to 1848, his remaining years up to his death in 1859 were spent there. Whereas the earliest accounts to include substantial observations on the vegetation and flora of Missouri (cited above) concerned travels up the Missouri River valley, the first naturalist to travel extensively in the Ozarks was Henry Rowe Schoolcraft (1793–1864), who would later gain fame as a geologist and noted expert on Native American languages and folklore (Rafferty, 1996). Although he collected no specimens, his observations of the geology, hydrology, vegetation, and human habitation of the region are the most detailed from that time period, and offer some of the best glimpses of what the Ozark landscape was like prior to large-scale colonization and alteration by settlers from the eastern United States. In the summer of 1818, Schoolcraft traveled from St. Louis to the mining town of Potosi (Washington County), where he completed observations on the mining and smelting activities in the area (Schoolcraft, 1819; Rafferty, 1996). Together with a companion, Levi Pettibone (1780–1881), Schoolcraft then hiked through portions of present-day Missouri and Arkansas with the intent of observing lead deposits in the James River area of southwestern Missouri. The pair left Potosi on November 6, 1818, and returned there on February 4, 1819, having traveled over 900 miles by Schoolcraft’s estimate. The full account of the journey was first published in 1821 (Rafferty, 1996). The next notable botanical activity in Missouri involved the “Yellowstone Expedition” of 1819–1820. This expedition consisted of a military portion intended to establish forts along the upper Missouri River (and possibly the mouth of the Yellowstone River) and an independent scientific portion, which was intended to map and explore a vast portion of the lands between the Mississippi River and the Rocky Mountains. The scientific portion was under the command of Major Stephen Harriman Long (1784–1864) and was the second of three major journeys he would take in the American West (the others are not pertinent to Missouri botany; see McKelvey [1955]). It was notable as the first scientific expedition funded by the U.S. government and included among its members specialists in zoology, geology, cartography, journalism, art, and botany. The initial botanist was the ill-fated William Baldwin (1779–1819; commemorated by Vernonia baldwinii Torr., among others), who had started his professional career as a physician before developing an interest in botany and who had already made important contributions to floristic knowledge of Georgia, Florida, and South America. Baldwin had a long his-

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tory of poor health (it has been speculated that he had tuberculosis) and apparently was under the mistaken notion that extended travel would benefit his condition (Benson, 1988). It is of interest to note that after arriving in St. Louis, Baldwin had the opportunity to spend time with the aging John Bradbury, who was living there. The expedition departed from St. Louis by steamboat on June 21, 1819, and made its way slowly upriver. Apparently most of the scientists were not prepared for the rigors of the trip and the poor living conditions aboard the ship, and Baldwin’s health declined quickly. By July 13, when the party reached Franklin (Howard County), Baldwin was too sick to continue with the expedition and (amazingly) spent his final few weeks making botanical observations in the area before his death on August 31. Baldwin’s specimens of about 100 species later made their way to the eminent botanist, John Torrey (1797–1873; commemorated by Juncus torreyi Cov., Pycnanthemum torrei Benth., and Schoenoplectus torreyi (Olney) Palla, among others), and now reside at the herbarium of the New York Botanical Garden. Baldwin’s written observations of the flora and vegetation were quite good and were drawn on heavily for inclusion in the official account of the expedition (Spaulding, 1908–1909; Benson, 1988). While Long returned east to recruit a new botanist (among other things), the expedition continued up the Missouri River. Eventually, Long rejoined the party at Council Bluffs, Iowa, accompanied by the young botanist Edwin James (1797–1861; commemorated by Pomaria jamesii (Torr. & A. Gray) Walp and Carex jamesii Schwein., among others). Except for a few collections around St. Louis, James did not have the opportunity to botanize in Missouri (Spaulding, 1908–1909) while a part of the expedition. It should be noted that although the expedition was important in many ways, including the first botanical trek across the lengths of Nebraska and Oklahoma and the first collections from alpine regions of Colorado, it ultimately was curtailed by unfortunate budget cutbacks from the U.S. Congress (setting a precedent that has been repeated frequently in subsequent scientific undertakings). The expedition ended on October 12, 1820, in Cape Girardeau, Missouri. James compiled the official account, which was first published in two volumes in 1823 (Benson, 1988). Among the noteworthy observations in this travelogue is the earliest report of numerous ticks in the eastern Ozarks, which plagued the travelers during the return portion of their trip. Several additional expeditions to the western United States traveled through Missouri (which was accorded statehood in 1820) from St. Louis during the next few decades. By this time, the Missouri River valley had been studied enough that this region was of only incidental interest to the botanists, who made few collections within the borders of present-day Missouri. Some of these expeditions were more successful than others. The Scottish botanist (and especially bryologist) Thomas Drummond (1807–1835; commemorated by several species in the Missouri flora, including an Aster, Cornus, Hypericum, Lipocarpha, Sapindus, Solidago, and an introduced Phlox) made two journeys in North America. The first trip (1825–1827) took him primarily through arctic portions of Canada. The second expedition (1831–1835), which was funded by the sale of sets of specimens to subscribers in England and America, was intended to take him from New York to Philadelphia and on to St. Louis, where he was supposed to join a fur-trading party going to Santa Fe. The immensity of his undertaking is reflected in the fact that he commissioned the delivery of two tons of collecting papers to St. Louis (McKelvey, 1955). After walking across the Allegheny Mountains from Philadelphia to Wheeling, Pennsylvania, Drummond traveled by steamboat to Louisville and then to St. Louis, where he arrived on July 19, 1831. However, an illness contracted on the boat forced him to remain in St. Louis until he could recover enough to travel, and thus he missed the opportunity to join the fur traders. While recovering, Drummond collected a number of plants on both the Illinois and Missouri sides of the Mississippi River, including (in Missouri) the type specimen of what is now called Lipocarpha drummondii (Nees) G.C. Tucker, which was named in his honor. Eventual-

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ly he was able to journey by boat to New Orleans, from which he sailed to eastern Texas the following year. His botanical activities in Texas made him famous in botanical circles of the time, and his specimens are widely distributed, including sets at the herbaria of Kew Gardens and the Missouri Botanical Garden. After departing from Texas in 1835, Drummond sailed to Florida with the intent of exploring this area before returning to England. Unfortunately, during a side trip to Cuba, a fatal illness cut short his brilliant career as a botanical collector. Another traveler in the Missouri River valley was Prince Maximilian Alexander Philipp of Wied-Neuwied (1782–1867; commemorated by Helianthus maximilianii Schrad., among others), the Prussian nobleman and naturalist. Although better known for an earlier voyage to Brazil, he also ascended the Missouri River into the northwestern United States with a furtrading expedition (1832–1834), reaching as far as Montana and retracing the route of Bradbury and Nuttall along the way. Maximilian’s travelogue originally was published in three volumes (including an atlas) in German (Maximilan, 1839–1841), followed in 1843 by an English translation. Some of his collections were destroyed by a fire on the boat from St. Louis on which they were being shipped home, but a large collection of birds and other animals and about 200 plant specimens survived. The plants (now accessioned at the Natural History Museum of Vienna) were of limited scientific value, however, as Maximilian recorded little or no information on the collection date or source of each specimen (McKelvey, 1955). Most of the specimens appear to have originated from lands to the west of Missouri. Maximilian’s travelogue is, however, notable for other reasons, including his detailed descriptions of the land and peoples of the Missouri River valley, particularly of the vegetation, the Native American cultures encountered, and the fur-trading industry. An ill-fated botanical journey was that of the Prussian botanist Heinrich Carl Beyrich (1796–1834; commemorated by Erigeron strigosus var. beyrichii (Fisch. & C.A. Mey.) Torr. & A. Gray), who accompanied an expedition of U.S. Army Dragoons into what was then Indian Territory. The expedition was intended as a show of power and to negotiate peace settlements with tribes hostile to settlers. Beyrich had botanized extensively in parts of the southeastern United States in 1833, before receiving permission to accompany the military expedition. In the company of a regiment of Dragoons and the famous painter of the American West, George Catlin (1796–1872), he left St. Louis in a wagon and with a servant during the spring of 1834. The main party was joined at Fort Gibson (in present-day northeastern Oklahoma) in July and the group then proceeded west (exactly how far is unknown, but perhaps into northern Texas) on an arduous journey through hostile territory. The poorly prepared expedition was plagued by extreme heat, drought, famine, disease, and Indian attacks, and many members did not survive the return to Fort Gibson. Beyrich and Catlin were both seriously ill. Catlin survived, but Beyrich died at Fort Gibson sometime around the end of August, perhaps of cholera. According to Catlin, Beyrich had assembled a large set of specimens during the trip, but what became of these is uncertain (McKelvey, 1955). Perhaps they are to be found with other Beyrich collections at the Botanical Institute of the University of Kiel, Germany. No account of the botany of the expedition was published. As noted previously, Thomas Nuttall’s second expedition to the west involved travels in Arkansas and Oklahoma, with no botanical activities in Missouri. However, his third trip again retraced the route of Lewis and Clark to the Pacific Northwest and involved extensive natural history activities in the state. Under the auspices of the American Philosophical Society, he joined a fur-trading expedition of Nathanial Jarvis Wyeth (1802–1856). Although not a botanist, Wyeth had made plant collections on his first expedition to the west, which Nuttall had studied and described, and Nuttall thus was invited along on the follow-up. In the company of the ornithologist John Kirk Townsend (1808–1851), he left St. Louis by foot in late March 1834, joining the rest of the group in Boonville (Cooper County) less than two weeks later. As no journal of Nuttall’s is known to exist, it is in Townsend’s (1839) narrative that

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most of the natural history information was published. Because of the earliness of the season, botanical activities were relatively minimal during the Missouri portion of the trip. From Boonville, the expedition proceeded by steamboat up the Missouri River and eventually reached the Pacific Ocean in September 1834. Subsequently, Nuttall traveled twice to Hawaii and explored portions of western California, returning to Philadelphia in 1836. The specimens collected on this immense journey are accessioned at the Academy of Natural Sciences of Philadelphia. The final major expeditions to travel through portions of Missouri during the 1830s were those of the French geographer and explorer Joseph Nicolas Nicollet (1786–1843; commemorated by Potentilla nicolletii (S. Watson) E. Sheld.). Several earlier explorers had attempted to trace the source of the Mississippi River, including Henry Rowe Schoolcraft (on two attempts both before and after his travels in the Ozarks). Nicollet was commissioned by the U.S. War Department in 1838 to map the major drainages of the Mississippi River, and he hired the German botanist Karl Andreas Geyer (1809–1853; commemorated by Chamaesyce geyeri (Engelm.) Small) to make a collection of botanical specimens during the trips. Also on the expeditions was Lieutenant John Charles Frémont (1813–1890; commemorated by Clematis fremontii S. Watson, among others), who would later gain fame for his explorations of the Rocky Mountains and the southwestern United States (including collections of some plant specimens). Geyer had arrived in America in 1835 with high hopes of achieving fame through botanical explorations in the West. Ill-equipped and inexperienced, his first few years of forays westward from St. Louis were relatively unproductive and are survived only by a few specimens at the Missouri Botanical Garden from the environs of St. Louis city. He met Nicollet on a boat trip from Kansas City to St. Louis in 1837 (McKelvey, 1955). The two expeditions left St. Louis in 1838 and 1839 and traveled up the Missouri River to present-day Minnesota and North Dakota, respectively. Many of the plants collected were lost when a mule carrying them toppled over a cliff, but the remainder were studied by John Torrey (see above), who published a catalog of the specimens as an appendix to Nicollet’s (1843) account of the expedition (which also contained important observations of the vegetation, geology, hydrology, etc. of the areas traversed). The specimens, nearly all from areas to the north of Missouri, are accessioned at the herbarium of the New York Botanical Garden. Geyer subsequently used St. Louis as a base of operations for botanical activities. In 1841, he joined a party led by John Frémont to explore the Des Moines River valley from Clark County, Missouri, to central Iowa. In 1842, he collected locally in areas around St. Louis in both Illinois and Missouri, financing his activities with the sale of sets of specimens (Gray, 1843; Engelmann, 1844). The collections were studied and distributed by George Engelmann (see below) and Asa Gray (1810–1888, famous botanist of Harvard University; commemorated by numerous species, including Carex grayi J. Carey). In 1843 and 1844, Geyer explored the Rocky Mountains with a group led by the wealthy Scottish sportsman and explorer, Captain (Sir) William Drummond Stewart (1794–1871), whose group was equipped in relative luxury and contained a number of botanists. He traveled from St. Louis as far north as Montana, but made no collections in Missouri. Instead of returning to St. Louis, however, he took the opportunity to reach Oregon and then to sail to England via Hawaii and around the tip of South America. Gray and Engelmann were thus deprived of the opportunity to work with Geyer’s 9,000–10,000 specimens, and these were studied and distributed by the dean of British botanists, William Jackson Hooker (1785–1865), at Kew Gardens. After a period in England, Geyer returned to Germany, married, taught botany lessons, and operated a plant nursery, but he never returned to America (McKelvey, 1955). In the 1840s, travelers from St. Louis who explored and botanized places to the west of

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Missouri, but made few or no collections in the state (summarized in McKelvey, 1955), included: William Gambel (1821–1849), who reached Santa Fe and then southern California in 1841, and died after repeating the voyage in 1849; John Frémont (see above), who made four expeditions through the Rocky Mountains to the western United States from 1842 to 1849, after his earlier work with Nicollet (see above); and the great ornithologist and artist, John J. L. Audubon (1780–1851), who painted detailed portraits of plants (some species unknown to science at the time) as perches for his birds, and whose North American travels included a trip up the Missouri River to the Yellowstone River in 1843. McKelvey (1955) also discussed Auguste Adolph Lucien Trécul (1818–1896), a French botanist who was sent to the United States in 1847 to study edible plants, and who botanized west of the Mississippi River in 1848–1850. Trécul spent much of his time on lengthy trips from St. Louis to the Rocky Mountains and Texas (and northern Mexico), but he apparently botanized in eastern Missouri during June and July of 1848. McKelvey cited (from Trécul’s unpublished notebook) collecting localities in Iron, Jefferson, St. Francois, Ste. Genevieve, and Washington Counties, as well as St. Louis, but these and other collections from 1848 were lost when the ship carrying them back to Europe sank. Only a few specimens from St. Louis that were given to George Engelmann (see below) have survived at the Missouri Botanical Garden herbarium. A few additional collections made during 1849 on the boat ride south from St. Louis to New Orleans were noted by McKelvey. These are preserved at the Natural History Museum in Paris and include a specimen of a spider flower (Cleome hassleriana Chodat) collected at Ste. Genevieve (Ste. Genevieve County) as an escape from cultivation (which was brought to my attention through the kindness of Dr. Hugh Iltis of the University of Wisconsin, Madison). Botanical exploration of the American west was far from over by 1850, when McKelvey’s volume concludes. However, by the mid-1840s, westward expansion had progressed to the point that St. Louis no longer was the gateway to the frontier. Subsequent expeditions often started from Fort Leavenworth, Kansas, and many of the notable journeys to the Southwest began in the port of Galveston, Texas. Thus, the history of Missouri botany shifted from activities of those from elsewhere who stopped to gather plants while on the way to points further west, to those who resided primarily in the state and carried out all or part of their botanical research here.

ST. LOUIS–AREA BOTANISTS Although he stayed but a short time, the first resident collector in Missouri was Lewis Caleb Beck (1798–1853; commemorated by Megalodonta beckii (Torr. ex Spreng.) Greene and the genus Neobeckia Greene). A physician by training, he moved to St. Louis toward the end of 1819 to join his brother Abraham Beck, a lawyer, and to set up a medical practice. During the year of 1820, he botanized extensively on both the Missouri and Illinois sides of the Mississippi River with the aim of summarizing the “mineral and vegetable riches” of the states through compilation of a gazetteer of the region (Beck, 1823). The summary of his botanical findings was the first such compilation of species for the area (Beck, 1826–1828) and contained several species new to science that are now considered to be common Missouri wildflowers, including Monarda bradburiana, Phlox bifida, Trillium recurvatum, and Trillium viride. He named the first of these for his friend, John Bradbury (see above), who was residing in St. Louis at the time of Beck’s arrival but was no longer active in botany. Beck did not like the frontier atmosphere of St. Louis and toward the end of 1820 he moved back to New York. Although he traveled to St. Louis again in 1821 to attend his brother’s funeral, he did no further botanizing in the region. His private herbarium eventually came to be accessioned

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at the New York State Museum, in Albany. Beck later went on to distinguish himself professionally, first as a physician and writer, then as a professor and lecturer in medicine, botany, and chemistry, and finally as state mineralogist of New York (Sebring, 1934). It was not until 1835 that another botanist took up residence in St. Louis. He was George Engelmann (1809–1884; commemorated by numerous species, including Isoetes engelmannii A. Braun, Ophioglossum engelmannii Prantl, and the genus Engelmannia Torr. & A. Gray) (Fig. 2), one of the most celebrated figures of all time in Missouri botany. Engelmann was broadly trained in botany and medicine in Germany. He immigrated to the United States in late 1832, where he joined relatives at the German settlement in Belleville, Illinois. He soon began to botanize the St. Louis Fig. 2. George Engelmann, from an oil painting by area. By 1835, Engelmann had established a R. A. Clifford, dated 1860. Photograph courtesy of the Missouri Botanical Garden Library Archives, medical practice in St. Louis, which he was to St. Louis. maintain successfully throughout the rest of his botanical career. Engelmann’s list of accomplishments is truly astounding (summarized by Wolf, 1988): He published over 100 botanical papers, including large monographs on difficult groups, such as Agave, Crataegus, Cuscuta, Euphorbia, Isoetes, Juncus, Quercus, Yucca, Cactaceae, and Pinaceae, among others; he published more than 600 species new to science; he founded the St. Louis Academy of Science; and he was largely responsible for the foundation of what would become the library and herbarium of the Missouri Botanical Garden. He was a colleague of most of the more prominent botanists of the eastern United States, as well as those in Europe, exchanging botanical observations and specimens with these individuals. Engelmann’s personal herbarium, which was incorporated into the Missouri Botanical Garden’s herbarium upon his death, contained more than 97,000 specimens. Aside from his own local botanical activities and excursions into portions of Arkansas, Illinois, the Appalachian and Rocky Mountains, the Lake Superior region, the Pacific Coast, and the Missouri Ozarks, Engelmann was instrumental in organizing the botanical activities of more than 30 expeditions to the western United States. He located botanists to accompany these expeditions and arranged for the distribution and sale of the resulting specimens, as well as the publication of summaries of the plants collected. It was this access to large numbers of newly collected specimens from the American West and his astuteness in their study that made George Engelmann one of the most important botanists of the nineteenth century. For most of his career, Engelmann had a close working relationship with Asa Gray (see above) and John Torrey (see above), and the three scientists were considered the foremost authorities on North American plants of their time (Shaw, 1986). Engelmann became a friend of the wealthy St. Louis businessman, Henry Shaw (1800–1889). Shaw had moved to St. Louis in 1819 from Sheffield, England, and amassed a fortune selling hardware and dry goods while also investing wisely in real estate. Shaw’s fortune allowed him to retire in 1839, at the relatively young age of 40. He developed an interest in botanical gardens during three lengthy trips to Europe between 1840 and 1852 (Rudolph, 1991), where he had the opportunity to visit several of the finest gardens there. It was William Jackson Hooker (see above), director of Kew Gardens and a towering figure in

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botany of the time, who suggested that Shaw consult with Engelmann for suggestions on the establishment of his own botanical garden. Engelmann persuaded Shaw to include a research component into his plans in order to lend scientific legitimacy to the endeavor. Although Engelmann was never formally on the staff of Shaw’s Garden, he remained Shaw’s botanical advisor for the rest of his life. With Shaw’s funding, he traveled to Europe for two years and acquired books and herbarium specimens that became the nucleus of the Garden’s research facilities (Shaw, 1986). The Missouri Botanical Garden was opened to the public in 1859 and included one museum building (still in use today) among its facilities. Upon Shaw’s death in 1889, it became a private institution governed by a board of trustees and a director. Shaw also established a school of botany at Washington University, in St. Louis, with an endowment to support the George Engelmann Professorship of Botany, which has been held by successive Garden directors. The research program begun by Engelmann and Shaw at the Garden became the most important center in the state for Missouri botany (and among the most important centers in the world for floristic and taxonomic research) and will be remarked on again repeatedly in the rest of this chapter. Soon after George Engelmann became established in St. Louis, other botanists also moved into the area. One of the first was the French botanist and plantsman Nicholas Riehl (1808–1852; commemorated by Clematis fremontii var. riehlii R.O. Erickson), who became a friend and admirer of Engelmann. Riehl settled in St. Louis in 1836 and opened what may have been the first plant nursery in the state of Missouri. Riehl’s nursery furnished to Henry Shaw some of the first trees that were planted in his botanical garden (Spaulding, 1908–1909). He kept a personal herbarium with both his own collections from Europe and Missouri and those he received on exchange from other European collectors. Eventually, this important collection was sold to Shaw and became part of the holdings at the herbarium of the Missouri Botanical Garden. Another early St. Louis botanist was the Jesuit missionary Jean-Baptiste Duerinck (1809–1857), who immigrated to the United States from Belgium in 1834 and lived in Florissant until 1840 while studying theology and teaching at St. Louis University. Born and educated in Belgium, Duerinck was an avid botanist and collected plants in the St. Louis area on both the Illinois and Missouri sides of the Mississippi River between 1836 and 1840 (Dorr, 1986). He exchanged specimens with George Engelmann and the southeastern botanist Stephen B. Buckley (1809–1884; commemorated by Solidago buckleyi Torr. & A. Gray, among others), whose herbarium also is now at the Missouri Botanical Garden. Buckley also botanized the St. Louis area (the two collected together in 1838), but he did most of his fieldwork during the trips to Illinois. Duerinck also sent specimens to the famous Belgian botanist Martin Martens (1797–1863), whose herbarium is at the Botanical Garden in Brussels. Martens described ten species as new to science based on specimens that Duerinck provided. Most of these have been relegated to synonymy under earlier names for the same taxa (Dorr, 1986), but Helianthus grosseserratus M. Martens is still considered the correct name for one showy sunflower. Duerinck continued his botanical activities after being transferred to Ohio in 1840 (later to Kentucky and Kansas). He was killed in a flatboat accident on the Missouri River near Independence while returning to St. Louis from his final posting in Kansas. The Prussian botanist Augustus Fendler (1813–1883; commemorated by Sphaeralcea fendleri A. Gray, among others) arrived in America in 1836 and eventually moved to St. Louis in 1839, where he was a lamp maker for a brief time before traveling to New Orleans and then to Texas for a year. Upon his return he taught school in Illinois during 1840 and 1841, later becoming a hermit on an island in the Missouri River near Wellington, Missouri, in Lafayette County (Stieber and Lange, 1986). A meeting with a botanist during a trip to Germany in 1844 got Fendler interested in

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botany. He moved back to St. Louis and began collecting the region’s flora under the tutelage of George Engelmann (see above), who determined his collections, which are accessioned at the Missouri Botanical Garden (Spaulding, 1908–1909). This experience prepared Fendler for a journey to Santa Fe in 1846 and 1847, where he collected numerous specimens for Asa Gray (see above). A return trip in 1849 was largely unsuccessful, because his collecting materials were lost in a flood. However, he completed other notable voyages, including four months in Panama later in 1849 and four years in Venezuela (1854–1858), making extensive collections on each of these journeys. Fendler eventually returned to Missouri in 1864 and settled in Allenton (St. Louis County). He lived with a brother there and was involved in the organization of the Missouri Botanical Garden’s herbarium (Spaulding, 1908–1909). Although few of his botanical collections from this time are preserved at the Garden, he apparently interacted with area botanists, meeting weekly with George Letterman (see below) to determine plants during the later portion of his time in Missouri (Spaulding, 1908–1909). In 1871, ill health forced his return to Europe. He returned to the eastern United States in 1873, settling in Delaware, but in 1877 he and his brother moved to Trinidad, where he spent the remainder of his life studying the local flora while suffering from rheumatism. The German physician Friedrich Adolph Wislizenus (1810–1889) moved to the St. Louis area in 1837 (Greger, 1935b) and eventually became the medical partner of George Engelmann (see above). Engelmann stimulated Wislizenus’s interest in plants, and he became an avid botanist and collector. He is best remembered for his travels and botanical activities in the southwestern United States and northern Mexico in 1846, but he also traveled to the Rocky Mountains, California, Panama, and Europe (Spaulding, 1908–1909; McKelvey, 1955). In between these trips, he resided in St. Louis. Wislizenus also was one of the founders of the Academy of Science of St. Louis (along with George Engelmann), and he tended Engelmann’s medical practice when Engelmann was off on botanical excursions. Most of his few Missouri collections were made during the earlier portion of his botanical career and are preserved at the Missouri Botanical Garden. Another physician/botanist was Theodore C. Hilgard (1828–1875), whose family emigrated from Bavaria to Belleville, Illinois, in 1836 (Spaulding, 1908–1909). Belleville at that time was a haven for political refugees from various parts of Germany, and relatives of Wislizenus and Engelmann also established households there. Hilgard eventually returned to Europe for his university training and studied medicine in Heidelberg, Vienna, and Berlin. He later became a pharmacist in Philadelphia, where a colleague, the American botanist and pharmacist, Elias M. Durand (1794–1873), stimulated his interest in plants. Hilgard returned to St. Louis in 1855, where he spent most of the remainder of his life practicing medicine. He was a charter member of the Academy of Science of St. Louis and published a number of papers in its Proceedings (Spaulding 1908–1909) and in other journals. Hilgard specialized mostly in nonvascular plants but also was very interested in developing an updated classification of angiosperms. His personal herbarium of about 12,000 specimens was donated to the University of California at Berkeley after his death, where it was destroyed during the great San Francisco fire of 1897. Another collection of lichens and bryophytes remained at the St. Louis Academy, where it also was destroyed by a subsequent fire. After Henry Shaw’s death in 1889, the Garden’s new board of trustees appointed William Trelease (1857–1945) as director, a position he would hold until 1912. Like Engelmann, Trelease had a broad interest in botany and published numerous taxonomic papers, including revisions of Acer, Agave, Quercus, Rumex, and Yucca. He also was one of the few directors who collected substantial numbers of specimens in Missouri (Table 1). Trelease received his doctorate from Harvard University in 1884 and was a professor at the University of Wisconsin before moving to the Garden and Washington University (Rudolph, 1991). He published on

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ecology, entomology, and mycology in addition to plant taxonomy. Trelease continued to develop the Garden’s scientific and educational programs, and the library and herbarium grew dramatically under his administration. After resigning in 1912, following a period of tension with the trustees, Trelease accepted a position as head of the Department of Botany at the University of Illinois at Champaign-Urbana. One way in which Trelease encouraged expansion of the research program at the Garden was to employ promising graduate students to work as assistants in the herbarium (Rudolph, 1991). Several of these became well-known botanists later, including Albert S. Hitchcock (1865–1935), who became the main authority on systematics of grasses at the Smithsonian Institution, and Louis H. Pammel (1862–1931), who became a leading botanist and conservationist at Iowa State College (later Iowa State University) and who made significant collections in the Rocky Mountains. Trelease also encouraged the activities of other local botanists, notably Hermann Von Schrenk (1873–1953), a mycologist and plant pathologist. Von Schrenk, who completed his doctorate at Washington University in 1897, was head of the Mississippi Valley Laboratory of the U.S. Department of Agriculture (which Trelease convinced him to locate at the Garden) from 1901 until 1907, when the government combined it with the U.S. Forest Service and moved the facility to Washington, D.C. (Spaulding, 1908–1909). He remained at the Garden for the rest of his career as staff pathologist and also did outside consulting work (Beilmann, 1953). Von Schrenk was a specialist on diseases of timber trees, particularly of Taxodium, and also an expert on the timber industry in the Midwest. However, his most enduring contribution to science and industry was his development of the process of impregnating wood with creosote to preserve it against decay, which was extremely important to both the timber industry and the railroad companies (which depend on wooden ties). The succession of directors that followed Trelease each brought a different emphasis to the Garden’s research programs. They are listed only briefly here. George T. Moore (1871–1956) was director from 1912 to 1953. Among his many accomplishments, he hired Jesse M. Greenman (1867–1951) to curate the herbarium and instructed him to oversee a program of study on the southwestern flora, including Missouri. Edgar Anderson (1897–1969) was an outstanding teacher, geneticist, and ethnobotanist, who was pressed into service from 1954 to 1957 after Moore’s retirement due to declining health. His research at Washington University and the Garden (Eisendrath, 1972) included several publications on the spiderwort genus, Tradescantia, notably a revision of the species of the United States (Anderson and Woodson, 1935) completed in collaboration with Robert E. Woodson (1904–1963), an eminent taxonomist and evolutionary biologist who was curator of the Garden’s herbarium from 1948 to 1963. Anderson later remained as a curator at the Garden until his sudden demise on the grounds. Frits W. Went (1903–1990), who was director from 1958 to 1963, was responsible for the Garden’s innovative “Climatron” geodesic dome greenhouse, as well as increased interest in cultural programs and the beginning of design and fundraising for a new facility to house the herbarium and library. David M. Gates (1921–), who was director from 1965 to 1971, was an ecologist who began the revitalization of the Garden’s flagging research program and also facilitated the completion of the new research facility initially conceived by his predecessor, Went. The Garden’s present director, Peter H. Raven (1936–), has had a profound effect on development of the research program. Born in Shanghai, China, raised in California, and educated in the University of California system, Raven took a very early interest in natural history and was immersed in botany at the age of eight through his membership in the Student Section of the California Academy of Sciences. By the time that he came to the Garden in 1971, he already had an extensive list of published papers and books on systematics, floristics, biogeography, and ecology and an international reputation as an outstanding researcher

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Table 1. Selected collectors of the Missouri flora, prior to January 1, 1998, listed alphabetically. Herbarium acronyms: A = Arnold Arboretum of Harvard University, Cambridge, MA; BEREA = Berea College, Berea, KY; DRURY = Drury College, Springfield; F = Field Museum of Natural History, Chicago; KANU = University of Kansas, Lawrence; KSP = Pittsburg State University, Pittsburg, KS; MICH = University of Michigan, Ann Arbor; MIL = Milwaukee Public Museum; MLT = Martha Lafite Thompson Nature Sanctuary, Liberty; MO = Missouri Botanical Garden; MODNR = Missouri Department of Natural Resources, Jefferson City; MOR = Morton Arboretum, Lisle, IL; MSSC = Missouri Southern State University, Joplin; MWSJ = Missouri Western State College, St. Joseph; NEMO = Truman State University, Kirksville; NMC = New Mexico State University, Las Cruces; NWMO = Northwest Missouri State University, Maryville; NY = New York Botanical Garden, New York; OKL = Oklahoma State University, Norman; SEMO = Southeast Missouri State University, Cape Girardeau; SIU = Southern Illinois University, Carbondale; SMITH = T. Smith personal herbarium, Jefferson City; SMS = Southwest Missouri State University, Springfield; SOTO = College of the Ozarks, Point Lookout; UARK = University of Arkansas, Fayetteville; UMKC = University of Missouri–Kansas City (now at Powell Gardens, Kingsville); UMO = University of Missouri–Columbia; US = U.S. National Herbarium, Washington, D.C.; WARM = Central Missouri State University, Warrensburg; WJC = William Jewell College, Liberty. Approx. Number Collected

Collector

Years of Collection

Collection Sites

Principal Repository(ies)

Craig Anderson

1993–

Statewide

MO