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Biosystematic Study of the Spiranthes cemua Complex MAR i i 1983 BOTANICAL GARDEN Charles J. Sheviak Curator of Botany New York State Museum 1982 Bulletin No. 448 New York State Museum The University of the State of New York THE STATE EDUCATION DEPARTMENT Albany, New York, 12230 Digitized by the Internet Archive in 2017 with funding from IMLS LG-70-15-0138-15 https://archive.org/details/bulletinnewyorks4481newy Biosystematic Study of the Spiranthes cemua Complex Charles J. Sheviak Curator of Botany New York State Museum The State Education Department does not discriminate on the basis of age, color, creed, disability, marital status, veteran status, national origin, race, or sex in the educational programs and activities which it operates. This policy is in compliance with Title IX of the Education Amendments of 1972. Inquiries concerning this policy may be referred to the Department’s Affirmative Action Officer, Education Building, Albany, NY 12234. 1982 Bulletin No. 448 New York State Museum The University of the State of New York THE STATE EDUCATION DEPARTMENT Albany, New York, 12230 , THE UNIVERSITY OF THE STATE OF NEW YORK Regents of The University (with years when terms expire) 1988 Willard A. Genrich, Chancellor, LL.B., L.H.D., LL.D., 1986 Litt.D., D.C.S., D.C.L. Buffalo 1988 J. Edward Meyer, Vice Chancellor, B.A., LL.B., L.H.D. Chappaqua Kenneth B. Clark. A.B.. M.S., Ph.D.. LL.D., L.H.D. D.Sc. Hastings on Hudson 1989 Emlyn I. Griffith. A.B., J.D Rome 1983 Mary Alice Kendall. B.S. Rochester 1984 Jorge L. Batista. B.A.. J.D., LL.D. Bronx 1986 Laura Bradley Chodos. B.A., M.A. Vischer Ferry 1987 Martin C. Barell, B.A., I.A., LL.B. Kings Point 1984 Louise P. MattEONI. B.A., M.A., Ph.D. Bayside 1987 R. CARLOS CARBALLADA, B.S., L.H.D. Rochester 1988 Floyd S. Linton, A.B., M.A., M.P.A.. D.C.I Miller Place 1988 Salvatore J. Sclafani, B.S., M.D Staten Island 1989 MlMI LlEBFR. B.A., M.A. New York 1985 Shirley C. Brown, B.A., M.A., Ph.D. Albany 1983 Robert M. Best, B.S. Binghamton President of The University and Commissioner of Education Gordon M. Ambach Executive Deputv Commissioner of Education Robert J. Maurer Deputy Commissioner for Cultural Education Carole F. Huxley Assistant Commissioner for the State Museum Director, State Science Service Richard H. Monheimer Chief Scientist, Biological Survey Norton G. Miller State Botanist Richard S. Mitchell CONTENTS Acknowledgements vi Introduction 1 Taxonomic Review 2 Synopsis of Characters 5 Roots 5 Leaves 5 Inflorescence 6 Floral Bracts 7 Flowers 7 Seeds 9 Cytology 9 Guard Cell Measurements II Apomixis 11 Teratologies 13 Pollination Biology and Hybridization 13 Ecology 14 Discussion 15 Taxonomic Treatment 18 Key 19 Spiranthes odorata 18 Spiranthes ochroleuca 20 Spiranthes magnicamporum 20 Spiranthes cernua 20 Nomina Ohscura 22 Literature Cited 23 Tables 27 Figures 33 iii Spiranthes is the most perplexing orchid genus in our flora. It is the least understood and the one that furnishes to authors who grow impatient under the restraints imposed by cautious progress, the best opportunities for the multiplication of species. It is a genus that repays intensive observation in the field and prolonged contempla- tion in the herbarium. Oakes Ames, 1921 ACKNOWLEDGEMENTS Many persons have contributed to this study, and it would not have been possible without their assistance. For their support, advice, and criticism of much of this work, I am indebted to Alan Haney, University of Illinois; and Richard E. Schultes, Leslie Garay, Reed Rollins, and Carroll Wood, Harvard University. Paul Catling, Agriculture Canada, has been most generous in the sharing of information, ideas, and materials and has contributed valuable criticism and review. The contribution of Paul Shildneck, Mt. Zion, Illinois, has been invaluable, due to his repeated provision of critical live material, notes, and assistance in the field. Others who have generously provided live material, photographs, data, and field assistance include Marlin Bowles, Illinois Na- ture Preserves Commission; Eric DeLin, Ft. Montgomery, New York; Carlyle Luer, Marie Selby Botanical Gardens; Timothy McCabe, New York State Museum; Lawrence Magrath, University of Science and Arts of Oklahoma; Max Medley, University of Louisville; David Niemann, Iowa State University; and Thomas Todsen, Las Cruces, New Mexico. I wish to thank the Curators and staffs of the herbaria which lent specimens for study or provided facilities during my visits to their collections, including AMES, BUF, CM, DEK, EIU, F, ILL, ILLS, ISM, KANU, MIL, MO, MWI, NEBC, NDA, NY, NYS, OKL, OKLA, PH, SIU, US, VDB, WIS; particular thanks are due Robert Evers, Illinois Natural History Survey, and Ronald McGregor, State Biological Survey of Kansas, for their generous loan arrangements and provision of live specimens, notes, and criti- cism. I am indebted to Timothy McCabe, New York State Museum, for pollinator determinations. Drawings are by Kathryn M. Conway, New York State Museum. This study has been supported in part through grants from the American Orchid Society Fund for Education and Research, the Atkins Fund, Harvard University, and the Femald Fund for Field Work in Systematic Botany, Harvard University; and through the research programs of the Department of Botany, University of Illinois; Department of Biology, Harvard University; and Biological Survey, New York State Museum. Additionally, the expenditure of personal funds by many of the persons above in the course of their contributing to this study is most noteworthy and is deeply appreciated. vi INTRODUCTION The genus Spiranthes has gained considerable noto- treat western populations within an eastern system riety as a taxonomically perplexing group. The history that did not in fact recognize their existence. of its study includes alternating periods of splitting and It was under such conditions that I began a study of lumping, and a general lack of acceptance of any one the complex in the Midwest. In 1968, I began to at- treatment has been apparent. In large part this situa- tempt to determine specimens of Spiranthes cernua tion seems to reflect the northeastern viewpoint from from the region, both in the herbarium and the field. which much of this work was done. Some of the most It eventually became evident that two of the taxa re- confusing species are wide-ranging; treatments which ported by various workers did not occur there, al- appeared appropriate in a limited geographic context though it was clear that the populations referred to S. often were inadequate and misleading elsewhere. This cernua were not uniform to any reasonable degree, has been particularly true of the S. cernua complex, a and that taxonomically distinct entities were present. group harboring some of the most intractable problems It was also clear that some were not members of any in the genus. The present study has benefited from an currently recognized taxa, and in 1973 I described the initial focus on the largely neglected western portions new species, S. magnicamporum. of the complex, where a combination of unique charac- The recognition of a new species in the complex was ter states and striking habitat differences provided significant beyond the addition of another taxon, for some early insight; subsequent work in the East has the emphasis during this century had been on a reduc- been guided by the Midwestern findings and has tion of taxa from specific rank to varietal status under shown the complex to behave similarly in both areas, S. cernua. The publication of S. magnicamporum was and, ultimately, has revealed its intrinsic nature. thus contrary to established precedent. Significantly, it was in part the lack of understanding of the situation In 1870, E. L. Greene, in his paper on the vegeta- in the Midwest that had led eastern workers to aban- tion of central Illinois, reported from the prairies of don specific status for some of the eastern taxa; the region “a Spiranthes of doubtful species.” He was Midwestern specimens were viewed as serving to obs- thus the first worker to recognize the systematic intri- cure the distinctions between eastern species. This sit- cacies of the Spiranthes cernua complex in the uation suggested that a reassessment of the entire prairies. It was not until very recently, however, that complex was in order. anyone seriously began to study the problem. Over Another important factor contributing to the confu- one hundred years elapsed between Greene’s publica- sion in the group has been the lack of information on tion and the description of S. magnicamporum, the the reproductive biology of the plants. Although poly- first Midwestern member of the complex to be sepa- embryony has long been known in some members of rated from S. cernua. the complex and its correlation with agamospermy es- This lack of treatment of the group stems in part tablished, the presence or absence of apomixis, al- from the difficulty encountered in the use of herba- though frequently cited in the literature, has not been rium material. Spiranthes does not provide many utilized nearly to its potential for elucidation of the characters of suitable quality for systematic work, and evolution and relationships of the various members of many that are present in living material, including the complex and of the intrinsic nature of the complex some of the most important, are lost when preserved as a whole. This failure on the part of earlier workers as dried specimens. The situation was made more diffi- no doubt arose through the almost complete lack of cult by the poor representation of midwestern material cytological study, although the evidence for polyploidy in the major eastern herbaria. The pioneering work in the genus should have stimulated this line of inves- that was conducted in the East thus suffered from a tigation.

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