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(Poaceae: Panicoideae) in Thailand

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(Poaceae: Panicoideae) in Thailand Systematics of Arundinelleae and Andropogoneae, subtribes Chionachninae, Dimeriinae and Germainiinae (Poaceae: Panicoideae) in Thailand Thesis submitted to the University of Dublin, Trinity College for the Degree of Doctor of Philosophy (Ph.D.) by Atchara Teerawatananon 2009 Research conducted under the supervision of Dr. Trevor R. Hodkinson School of Natural Sciences Department of Botany Trinity College University of Dublin, Ireland I Declaration I hereby declare that the contents of this thesis are entirely my own work (except where otherwise stated) and that it has not been previously submitted as an exercise for a degree to this or any other university. I agree that library of the University of Dublin, Trinity College may lend or copy this thesis subject to the source being acknowledged. _______________________ Atchara Teerawatananon II Abstract This thesis has provided a comprehensive taxonomic account of tribe Arundinelleae, and subtribes Chionachninae, Dimeriinae and Germainiinae of the tribe Andropogoneae in Thailand. Complete floristic treatments of these taxa have been completed for the Flora of Thailand project. Keys to genera and species, species descriptions, synonyms, typifications, illustrations, distribution maps and lists of specimens examined, are also presented. Fourteen species and three genera of tribe Arundinelleae, three species and two genera of subtribe Chionachninae, seven species of subtribe Dimeriinae, and twelve species and two genera of Germainiinae, were recorded in Thailand, of which Garnotia ciliata and Jansenella griffithiana were recorded for the first time for Thailand. Three endemic grasses, Arundinella kerrii, A. kokutensis and Dimeria kerrii were described as new species to science. Phylogenetic relationships among major subfamilies in Poaceae and among major tribes within Panicoideae were evaluated using parsimony analysis of plastid DNA regions, trnL-F and atpB- rbcL, and a nuclear ribosomal DNA region, ITS. A total of 132 grasses from six subfamilies and representatives from five tribes of Panicoideae, Andropogoneae (11 subtribes), Arundinelleae (4 genera), Eriachneae (1 genus), Isachneae (2 genera), and Paniceae (4 subtribes), were analysed. The PACCMAD clade, including Panicoideae, Arundinoideae, Centothecoideae, Chloridoideae, Micrairoideae, Aristidoideae and Danthonioideae, was well resolved. A close relationship between Aristidoideae and Chloridoideae was found. The monophyly of Micrairoideae was resolved but the relationships of three tribes (Eriachneae, Isachneae and Micraireae) within Micrairoideae were not fully resolved, only Eriachne and Isachne were resolved as monophyletic. Panicoideae s.s. were supported as monophyletic and sister to a clade of Danthoniopsis and Tristachya. Within Panicoideae, only a clade of tribe Andropogoneae + Arundinella + Garnotia was supported by bootstrapping. None of the analyses supported the monophyletic status of tribe Paniceae. Within Paniceae, the bristle clade (excluding Cenchrus) + a rare Australian non bristle-bearing genus, Alexfloydia, and the forest shade clade, were found, but their circumscription remains ambiguous. A sister relationship between Homopholis and Walwhalleya, which are endemic and rare Australian Paniceae grasses, was also resolved. Tribe Arundinelleae was found to be polyphyletic. This study supported the separation of Arundinella and Garnotia from the remaining Arundinelleae and the inclusion of both genera in their own subtribes (Arundinellinae Honda s.s. and Garnotiinae Pilger) within the III Andropogoneae. Tribe Arundinelleae should be abandoned as a taxonomic group within the centothecoid + panicoid clade. Tribe Andropogoneae were found to be monophyletic only if Arundinella and Garnotia were included. Only five out of a total of 11 subtribes from Andropogoneae, including Chionachninae, Coicinae, Dimeriinae, Germainiinae and Tripsacinae, were supported as monophyletic. This was the first time that Dimeriinae and Germainiinae were included in a molecular study. Five distinct groups within Andropogoneae corresponding to three genera from subtribe Rottboelliinae (Hackelochloa, Hemarthria and Mnesithea), an agamic complex (Bothriochloa, Capillipedium and Dichanthium), a sub-basal awn group (Arthraxon and Thelepogon), the core Andropogoneae genera (Hyparrhenia Cymbopogon, Schizachyrium and Andropogon), and a Themeda group, were supported as monophyletic. Plastid microsatellites or simple sequence repeat (SSR) DNA markers in conjunction with Nei’s (1978) unbiased genetic distance and UPGMA were employed to assess the intrapopulational and interpopulational variations of two forest grasses with contrasting breeding system, Arundinella setosa and Garnotia tenella. A total of seven haplotype and four groups from 11 populations of A. setosa, and a total of 11 haplotypes and four groups from eight populations of G. tenella, were defined. The patterns of haplotype distributions of both grasses reflect patterns of plant dispersal via seed. The low population genetic differentiation among populations in both grasses (Arundinella = 0.049; Garnotia = 0.155) suggested that the most variation in haplotypes is distributed within populations and not among populations. The high value of genetic variation within A. setosa and G. tenella populations suggests that they consist of multiple genets. The high value of genetic diversity but low gene flow estimates of both grasses, found in Northern Thailand, can be affected by topographic barriers and habitat fragmentation. No obvious correlations of biogeographical distribution in A. setosa and G. tenella were found that were consistent with previously defined Thai floristic regions but groupings in the analysis indicated a general east-west divide. IV Acknowledgements First of all, I would like to thank my family for their support, encouragement, and unshakable faith in my abilities during the course of my studies. I wish to thank my great supervisor Dr. Trevor Hodkinson for his enthusiasm, support, friendship and for all the encouragement he gave me during my Ph.D. study. I also wish to thank those who helped me in collecting grass specimens Drs. Simon Laegaard, Surrey Jacobs, Trevor Hodkinson, Sarawood Sungkaew and Ms Narumon Kritsanachandee. I am also grateful to thank Dr. David Simpson and Stephen Renvoize who helped me when I was working in the herbarium at the Royal Botanic Gardens, Kew, England and to Dr. Vincent Savolainen and Mr. Laszlo Csiba when I was in the Jodrell Laboratory of Kew. I would like to express my gratitude to Prof. Dr. Pranom Chantaranothai for his kindness, comments and useful suggestions and Drs. Pimwadee Pornpongrungrueng, Paweena Traiperm for their support of the work, and Prof. Dr. Benjamin Øllgaard for his kindly suggestions and for supplying the Latin diagnosis. Many thanks to all curators and the staff of the following herbaria: AAU, ABD, B, BK, BKF, BKU, BM, C, CMU, E, GH, K, KKU, L, NY, QBG, SING and TCD for the use or loan of specimens. Thanks very much to all members of the Botany Department from Trinity College. I will not forget Ireland! Last, but not least, a special thanks to Prof. Dr. John Parnell, for his very kind help, encouragement, and support throughout my research. This work was supported by the TRF/BIOTEC Special Program for Biodiversity Research and Training Grant BRT_148026, the Trinity College Dublin Postgraduate Travel Reimbursement Fund, Natural History Museum, National Science Museum, Technopolis, Pathum Thani, Thailand. This thesis and the work carried out herein are dedicated to the memory of Dr. Jarujin Nabhitabhata (Director, Natural History Museum, Thailand) 1950-2008. V Table of Contents Declaration …………...……………………………………………………………………. II Abstract …………………………………………………………………………………... III Acknowledgments ……………………………………………………………….……….... V CHAPTER 1 Introduction ……………………………………………………………….. 1 1.1 General introduction ……………………………………………………….………… 1 1.1.1 Morphology ………….………………………………………………………...…. 1 1.1.2 Major groups of grasses and their classification ……………………………............. 8 1.2 Poaceae and the Flora of Thailand project ………………………...………………..... 20 1.3 Aims of this thesis …………………………………………………………………... 21 CHAPTER 2 Floristic treatments ……………………………………………………… 28 2.1 Introduction ………………………………………………………………………… 28 2.1.1 Taxonomic history of Arundinelleae ………………………………………….…. 28 2.1.2 Taxonomic history of Chionachninae ……………………………..…….……….. 31 2.1.3 Taxonomic history of Dimeriinae …………………..…………………………… 33 2.1.4 Taxonomic history of Germainiinae ………………………………..…….…...…. 34 2.2 Aims ………………………………………………………………………................. 36 2.3 Materials and methods ……………………………………………………………..... 36 2.4 Species concept ……………………………………………………………................. 38 2.5 Results ……………………………………………….…………………………….... 40 Tribe Arundinelleae ………………………………………………………………....… 40 Key to genera of Thai Arundinelleae ……….…………………….……………….… 40 Arundinella ……...………………………………………………………………...... 41 Key to species of Thai Arundinella ……………………………...……………..…. 43 Arundinella bengalensis (Spreng.) Druce ……………………………………….… 44 Arundinella cochinchinensis Keng ………………………………………………… 47 Arundinella decempedalis (Kuntze) Janowski ……………………………………... 48 Arundinella holcoides (Kunth) Trin. …………………………….………………... 50 Arundinella kerrii C. Hambananda ex Teerawat. & Sungkaew ………………....... 52 Arundinella kokutensis Teerawat. & Sungkaew ……………..…………………… 54 Arundinella nepalensis Trin. ………………….………………….…………..….... 56 Arundinella setosa Trin ………………………………….………………….…… 58 VI Key to the varieties of Arundinella
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