Brigham Young University Science Bulletin, Biological Series Volume 19 Number 2 Article 1 3-1974 Taxonomy of the genus Chloris (Gramineae) Dennis E. Anderson Department of Biology, Humboldt State College, Arcata, California 95521 Follow this and additional works at: https://scholarsarchive.byu.edu/byuscib Part of the Anatomy Commons, Botany Commons, Physiology Commons, and the Zoology Commons Recommended Citation Anderson, Dennis E. (1974) "Taxonomy of the genus Chloris (Gramineae)," Brigham Young University Science Bulletin, Biological Series: Vol. 19 : No. 2 , Article 1. Available at: https://scholarsarchive.byu.edu/byuscib/vol19/iss2/1 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Brigham Young University Science Bulletin, Biological Series by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. S'/U/?- FJrpya Brigham Young University Science Bulletin *'''^ubrTry°°'" MAY271P74 HARVARD UNIVERSITAXONOMY OF THE GENUS CHLORIS (GRAMINEAE) by Dennis E. Anderson BIOLOGICAL SERIES — VOLUME XIX, NUMBER 2 MARCH 1974 /ISSN 0068-1024 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES Editor: Stanley L. Welsh, Department of Botany, Brigham Young University, Provo, Utah Acting Editor: Vernon J. Tipton, Zoology Members of the Editorial Board: Ferron L. Andersen, Zoology Joseph R. Murdock, Botany WiLMER W. Tanner, Zoology Ex officio Members: A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Director, Brigham Young University Press The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology. Separate numbers and back volumes can be purchased from University Press Marketing, Brigham Young University, Provo, Utah 84602. All remittances should be made payable to Brigham Young University. Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84602. Brigham Young University Science Bulletin TAXONOMY OF THE GENUS CHLORIS (GRAMINEAE) by Dennis E. Anderson BIOLOGICAL SERIES — VOLUME XIX, NUMBER 2 MARCH 1974 / ISSN 0068-1024 TABLE OF CONTENTS ABSTRACT 1 ACKNOWLEDGMENTS 1 INTRODUCTION 1 NOMENCLATURAL HISTORY OF THE GENUS 2 TAXONOMIC RELATIONSHIPS OF CHLORIS 2 Cytology 3 Leaf Epidermis 5 Anatomy, Embryology, and Seedling Morphology 6 Relationships of Chloris and Other Genera 9 Relationships within Chloris 13 Chloris Swartz 15 Key to the species of Chloris 16 1. Chloris roxlmrghiana Schultes 22 2. Chloris puniculata Scribner in Robinson 24 3. Chloris herroi Arechavaleta 25 4. Chloris ciliata Swartz 27 5. Chloris lamproparia Stapf in Chevalier 30 6a. Chloris canterai Arechavaleta var. canterai 30 6b. Chloris canterai Arechavaleta var. grandiflora ( Rosengurtt & Izaguirre de Artucio) Anderson 32 7. Chloris dantUjiina C. D. Adams 34 8. Chloris cruciata (Linnaeus) Swartz 37 9. Chloris filiformis (Vahl) Poiret in Lamarck 39 10. Chloris suringari Hitchcock in Urban 39 11. Chloris scariosa F. von Mueller 40 12. Chloris prieurii Kunth 42 13. Chloris rohusta Stapf in Chevalier 43 14. Chloris mollis (Nees) Swallen 45 15. Chloris longiaristata Napper 46 16. Chloris quinque.sctica Bhide 48 17. Chloris wightiana Nees ex Steudel 48 18. Chloris montana Roxburgh 51 19. Chloris bournci Rangachariar & Tadulingam 53 20. Chloris inflata Link 53 21. Chloris formosana (Honda) Keng 57 22. Chloris pilosa Schumacher 58 23. Chloris virgata Swartz 60 24. Chloris gai/aiu/ Kiuith 65 25. Chloris castilloniatui Lillo & Parodi 69 26. Chloris orthonoton Doell in Martins 69 27. Chloris aristata (Cervantes) Swallen 70 28. Chloris hrandegei (Vasey) Swallen 72 29. Chloris chloridca ( Presl) Hitchcock 74 30. Chloris ventricosa R. Brown 76 31. Chloris truncate R. Brown 78 32. Chloris pumilio R. Brown 80 33. Chloris lobata Lazarides 82 34. Chloris divaricata R. Brown 83 35. Chloris pcctinata Bentham 86 36. Chloris crinita Lagasca 87 37. Chloris pluriflora (Foumier) Cla)'ton 89 38. Chloris cticullata Bischoff 91 39. Chloris verticillata Nuttall 93 40. Chloris andropogonoidcs Foumier 95 41. Chloris texensis Nash 103 42. Chloris sesqiiiflora Burkart 104 43. Chloris humhcrtiana A. Camus 104 44. Chloris amethi/stca Hoclistt-tter 106 45. Chloris subniutica Humboldt, Bonplantl. Kunth 106 46. Chloris somalewis Rendle 109 47. Chloris cleimntis Merrill 109 48. Chloris radiata (Linnaeus) Swartz 110 49. Chloris pycnotlirix Trinius 114 50. Chloris halophila Parodi 1 17 .51. Chloris ekmanii Hitchcock 119 52. Chloris arcmiria Hitchcock & Ekman in Hitchcock 119 53. Chloris sagnwuna Achille Richard in .Sagra 121 54. Chloris cubensis Hitchcock & Ekman in Hitchcock 123 55. Chloris mossambicensis Schumann 123 56. Chloris hurrnetuis Ander.son 125 UNIDENTIFIED, REJECTED, OR REASSIGNED NAMES 126 LITERATURE CITED 131 TAXONOMY OF THE GENUS CHLORIS (GRAMINEAE) by Dennis E. Anderson' ABSTRACT This study is a worldwide taxononiic treat- of unidentified or rejected names. Distributions ment utilizing endomorpliic characters such as and maps are given for each species, along with cytology, histology, and ciiihrNology, in addition lists of specimens examined dining studies at to the traditional cxoniorphic features. New various iu-rbaria. Reports of field studies in- chromosome numbers and histological informa- volving introgression among certain species are tion are reported. Relationships between Chloris also included. A new species, Chloris hurmen- and other genera in the Chlorideae are dis- sis, is described from Bumia, and a new com- cussed. A total of r6 species is recognized in the bination, Chloris canterai Arech. var. graiuliflora genus; each is described and illustrated. Com- ( Rosengurtt & Izaguirre de Artucio ) Anderson, plete synonymies are included as well as a list is also proposed. ACKNOWLEDGMENTS These studies were begun in the summers A special debt is owed to Dr. W. D. Clayton of 1963 and 1964 as part of a National Science of the Royal Botanic Gardens, Kew, England, Foundation program in l\esearch Participation who provided advice, cooperation and friend- for College Teachers sponsored b\' the Depart- ship, and thus made my stay at Kew most pro- ment of Botan\-, University of Texas. Additional ductive. financial support was provided b\' this program California State University, Humboldt, has through grants ( NSF GY-3002, NSF GE-69:8). generously provided released time and leaves in The assistance of Dr. Harold Bold, director of support of these studies, and the continued as- the summer program, is also gratefully acknowl- sistance of President Cornelius Siemens is ac- edged. Initialh', Dr. Walter V. Bro\\'n pointed knowledged with many thanks. out to me many of the biosystematie problems I would also like to thank Mr. Rupert C. of Chloris in Texas, and I am grateful to him Barneb\- for his aid in the preparation of the for his continued encouragement. Latin description of Chloris btirmsiisis. .additional direct assistance was received Michael Lazarides kindly let me examine his from the National Science Foundation in 1969 manuscript treatment of Chloris in Queensland, (NSF GB-7235), enabling me to visit a number prior to its publication. of major overseas herbaria and to do additional I am grateful to my wife, Nancy, for the end- field work. The curators were most helpful and less hours she contributed in filing and organiza- allowed me to borrow man\- specimens. Their tion, in establishing and verifying geographical cooperation is much appreciated. locations, and, finally, in cartography. INTRODUCTION With its delicate, digitate clusters of spikes, of Nebraska, from the Cape of Good Hope to Chloris must certainly be among the most strik- the sands of the Sahara, from Afghanistan to ing of grass genera. The distribution spans the island archipelagoes of the South Pacific, the warmer regions of most of the continents, and from the subcontinental islands of Ne.v from the pampas of Argentina to the sandhills Guinea and Borneo through the continent of ^Department of Biology, Humboldt State College, Areata, California 95521. Bricham Young University' Science Bulletin Australia. Thus, it is likely that taxonomists in me to conclude that a classical studv was needed these areas have seen (me or more of the taxa and that experimental studies could then be in the genus. Though these attributes would made on a sounder foundation. seem to insiue popularity among agrostologists, Well over 15,000 herbarium specimens and Chloris nevertheless has not been subjected to many mass collections from hybridizing popula- any systematic review for nearly a centuiy. This tions were studied. The numbers of specimens has been an era characterized by extensive ex- available van' widely. Some taxa are repre- ploration and collection of hitherto unknown sented only by the holot\'pe. Others are known areas, as well as the development of revolu- from such large numbers of specimens that tionary systematic thought. population descriptions may be advanced with My interest in Chloris began with studies in confidence. Still others ha\'e been collected so hybridizing species in Texas; this interest spread thoroughly that the enormous complexit)' of the later to similar studies in the Southeast; and ulti- population has revealed the necessity for further mately it encompassed more classical investiga-