\ Monograph of the Neotropical Fern Genus Polybotrya (Dryopteridaceae) RobbinC. Moran Volume 34, Article 1 November 1987 Monograph of the Neotropical Fern Genus Polybotrya (Dryopteridaceae) RobbinC. Moran Illinois Natural History Survey Illinois Department of Energy and Natural Resources Department of Plant Biology University of Illinois at Urbana-Champaign Illinois Natural History Survey Bulletin Volume 34. Article 1 November 1987 This work is dedicated to my parents, Livia Ann Moran and John Howard Moran. Dr. Robbin C. Moran currently works at the Missouri Botanical Garden in St. Louis. Illinois Natural History Survey, Lorin I. Nevling, Chief A Division of the Illinois Department of Energy and Natural Resources Printed by Authority of the State of Illinois (62997—1,200—11-87) No charge is made for most publications of the Illinois Natural History Survey, and a list of those publications is available upon request. Single copies of most publications are available to anyone request- ing them. Requests for multiple copies should be made in writing and should explain the use to be made of the publications. Address correspondence to the Office of the Chief at the address below. Illinois Natural History Survey Natural Resources Building 607 East Peabody Drive Champaign, Illinois 61820 Citation: Moran, R.C. 1987. Monograph ofthe Neotropical Fern Genus /'o/)*o/r>'<'(Dryopteridaceae). Illinois Natural History Survey Bulletin 34 (I): U1.18. US ISSN 0073-4918 ... Contents Acknowledgments v Part One: Introduction and Discussion Materials and Methods 1 Taxonomic History of the Genus 3 Ecology 6 Geography 7 Morphology and Anatomy II Chromosome Numbers 27 Cladistic Analysis of the Species 27 Subdivision of the Genus 31 Relationships with Other Groups 34 Part Two: Taxonomic Treatment Notes on the Presentation of Data 37 Description of the Genus 38 Key to the Species of Polybotrya 39 Species Descriptions 43 Polybotrya subgenus Soromanes 1 Polybotrya serratifolia (Fee) Klotzsch 43 2. Polybotrya polybotryoides (Baker) Christ 46 3. Polybotrya suberecta (Baker) C. Chr 50 4. Polybotrya andina C. Chr 50 Polybotrya subgenus Sorbifolia 5. Polybotrya sorbifolia Kuhn 53 6. Polybotrya fractiserialis (Baker) J. Smith 54 7. Polybotrya crassirhizoma Lellinger 58 8. Polybotrya espiritosantensis Brade 60 Polybotrya subgenus Polybotrya 9. Polybotrya caudata Kunze 60 10. Polybotrya goyazensis Brade 66 1 1 Polybotrya pubens Martius 68 12. Polybotrya glandulosa Kuhn 71 13. Polybotrya lechleriana Mettenius 71 14. Polybotrya attenuata Moran 74 15. Polybotrya stolzei Moran 76 16. Polybotrya alfredii Brade 78 17. Polybotrya botryoides (Baker) C. Chr 80 18. Polybotrya lourteigiana Lellinger 82 19. Polybotrya pittieri Lellinger 84 20. Polybotrya cylindrica Kaulfiiss 86 2 1 Polybotrya hickeyi Moran 88 22. Polybotrya puberulenta Moran 90 23. Polybotrya alata Moran 92 . 24. Polybotrya aequatoriana Moran 94 25. Polybotrya appressa Moran 94 26. Polybotrya altescandens C. Chr 97 27. Polybotrya gomezii Moran 99 28. Polybotrya osmundacea Willd 101 29. Polybotrya cyathifolia Fee 106 30. Polybotrya latisquamosa Moran 108 31. Polybotrya sessilisora Moran 108 32. Polybotrya canaliculata Klotzsch Ill 33 Polybotrya semipinnata Fee 113 34. Polybotrya speciosa Schott 115 35. Polybotrya pilosa Brade 117 Names of Uncertain Application 119 Excluded Taxa 120 Literature Cited 122 Taxa and Distribution of Polybotrya 1 24 Distribution Maps 124 Index to Collectors' Numbers 132 Index to Taxonomic Names 137 Acknowledgments No one helped me more than the five pteri- the first at Flathead Lake Biological Station, dologists who carefully reviewed every page Montana, and the second at Mountain Lake of this monograph: Dr. David B. Lellinger, Biological Field Station, Virginia. I thank Smithsonian Institution, Washington, D.C.; them for these memorable summers. Dr. Alan R. Smith, University of California, Dr. Kenneth R. Robertson of the Illinois Berkeley; Mr. Robert G. Stolze, Field Mu- Natural History Survey, Champaign, was my seum of Natural History, Chicago, Illinois; major advisor and provided much help and Dr. Rolla M. Tryon, Harvard University, encouragement. Others who served on my Cambridge, Massachusetts; and Dr. Warren doctoral committee were Dr. J. Leland H. Wagner, Jr., University of Michigan, Ann Crane, Illinois Natural History Survey, Arbor. Not only did they spend long hours Champaign; Drs. Johannes M. J. de Wet, reading my rough manuscripts, but they also Zane B. Carothers, and Malcolm L. Sargent, patiently answered my numerous queries University of Illinois at Urbana-Champaign; about the intricacies of fern taxonomy. Dr. Rolla M. Tryon, Harvard University, My first field trip—to Costa Rica—was Cambridge; and Dr. Warren H. Wagner, Uni- made on funds awarded by the Tinker Foun- versity of Michigan, Ann Arbor. Dr. Tom dation. Doctoral Dissertation Improvement L. Phillips, University of Illinois at Urbana- Grant # 83-06990 from the National Science Champaign, provided numerous helpful com- Foundation provided most of my financial ments, esp)ecially on the Ecology and Geog- support, primarily for three additional col- raphy sections. lecting trips in Latin America and for a semes- 1 completed most of the work for this ter of study at Harvard University. monograph in the herbarium at the Illinois Two people were particularly helpful dur- Natural History Survey, and I am greatly in- ing my fieldwork in Latin America. First, debted to that institution for its support and Mr. Luis D. Gomez P. of the Museo Nacional for the use of its facilities. Several biologists de Costa Rica, San Jose, gave me lodging in at the Survey helped me in various ways. Mr. his house and extended numerous other cour- John Taft and Ms. Mary Kay Solecki provid- tesies while I worked in Costa Rica. Second, ed useful discussion and patiently tested my Professor Francisco Ortega, UNELLEZ, keys. Dr. David Swofford was helpful with Guanare, Venezuela, gave me the use of his the cladistic analysis. Mr. Bill N. McKnight herbarium and plant-drying facilities and took was a constant friend and critic throughout me on collecting trips through the Andes of the study and assisted with the reproduction western Venezuela. I thank them both for of the illustrations. 1 am also indebted to the their hospitality and for sharing their knowl- editorial staff at the Survey, especially Patty edge of ferns. Duzan and Eva Steger who typeset the man- During my semester of study at Harvard uscript, Audrey Hodgins who edited it. and University, Dr. Alice F. Tryon was extremely Molly Scott who helped with the production. helpful in showing me how to obtain high I am grateful to the directors and curators quality scanning electron microscope photo- of the following herbaria for the large quan- graphs of spores. Dr. Rolla M. Tryon helped tities of valuable material they made available with nomenclatural, systematic, and other to me, often for a considerable period of time. matters. The abbreviations used in the text follow the I learned much about pteridophyte biol- names of the herbaria; Arnold Ariwretum, ogy from two summer field courses with Drs. Harvard University (A), Cambridge, Mas- Warren H. Wagner and Rorence S. Wagner, sachusetts; Herbarium Jutlandicum, Univer- sity of Aarhus (AAU), Risskov, Denmark; New York; Museum National d'Histoire Herbarium Amazonense, Universidad Na- Naturelle (P), Paris, France; Academy of cional de la Amazonia Peruana (AMAZ), Natural Sciences of Philadelphia (PH), Iquitos, Peru; Botanischer Garten und Philadelphia, Pennsylvania; Herbario Uni- Botanisches Museum (B), Berlin, Federal versitario, Universidad Nacional Experimen- Republic of Germany; British Museum of tal de los Llanos Occidentals "Ezequiel Natural History (BM), London, England; Zamora" (PORT), Portuguesa, Venezuela; Herbier (CAY), Cayenne Cedex, French Herbario del Instituto de Ciencias Naturales, Guiana; Herbario Nacional Colombiano, Universidad Central (Q), (Juito. Ecuador; In- Museo de Historia Natural, Universidad Na- stituto de Ciencias, Pontificia Universidad cional (COL), Bogota, Colombia; Herbario Catolica del licuador (C?CA), Quito, Ecua- Nacional de Costa Rica, Museo Nacional dor; Jardim Botanico do Rio de Janeiro (RB), (CR), San Jose, Costa Rica; John G. Searle Rio de Janeiro, Brazil; Department of Bot- Herbarium, Field Museum of Natural History any, University of California (UC), Berke- (F), Chicago, Illinois; Conservatoire etJardin ley, California; United States National Her- botaniques de la Ville de Geneve (G), Swit- barium, Smithsonian Institution (US), Wash- zerland; Gray Herbarium, Harvard Univer- ington, D.C.; Herbario San Marcos, Museo sity (GH), Cambridge, Massachusetts; de Historia Natural, Universidad Nacional Rijksherbarium (L), Leiden, Netherlands; In- Mayor de San Marcos de Lima (USM). Lima, stituto Miguel Lillo de la Fundacion Miguel Peru; Instituto Botanico (VEN), Caracas, Lillo (LIL), Tucuman, Argentina; Facultad Venezuela; Pringle Herbarium, University of de Ciencias Naturales y Museo, Division Vermont (VT), Burlington, Vermont; Institut Plantas Vasculares, Universidad Nacional de fiir systematische Botanik der Universitat La Plata (LP), La Plata, Argentina; Herbario Zurich (Z), Zurich, Switzerland. Nacional de Bolivia, Universidad Mayor de This work was originally part of a disser- San Andres (LPB), La Paz, Bolivia; Her- tation submitted to the Graduate College of barium of the University of Michigan the University of Illinois at Urbana-Cham- (MICH), Ann Arbor, Michigan; Missouri paign in partial fulfillment