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Systematics of Elatostema (Urticaceae)

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Systematics of Elatostema (Urticaceae) SYSTEMATICS OF ELATOSTEMA (URTICACEAE) Julisasi Tri Hadiah A thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy University of New South Wales July 2007 Acknowledgements I would like to express my gratitude to my supervisors, A/Prof Paul Adam (University of New South Wales), Dr Barry Conn (NSW), and Dr Chris Quinn (NSW) for providing guidance and expert advice throughout my thesis, with a special thanks to Chris who continued supporting me even after his retirement. I would especially like to thank Drs Dedy Darnaedi, (former Director Kebun Raya Indonesia) and Irawati (current Director) for giving me permission to study overseas and for their continuing support throughout this project. Likewise, I wish to thank Drs Tim Entwisle (Director Royal Botanic Gardens Sydney) and Brett Summerell (Director Plant Sciences and Public Programs, NSW) for generously allowing me full access to the facilities of their organisation. I would also like to thank the staff of the Royal Botanic Gardens, Sydney and, particularly, those of the National Herbarium of New South Wales, who have provided assistance to ensure that my research on the systematics of Elatostema was successful. It has always been an enjoyable and excellent organisation in which to work. I have met many friendly people. Especially, I wish to thank Hannah McPherson, Lucy Nairn, Gillian Towler, Andrew Perkins and Nick Yee for their warm friendship, their help when moving to new homes, taking me to hospital when sick, for great weekends together, for having fun collecting orchids and learning about freshwater ecology, up to my knees in the freezing water of the Kangaroo River. They have always cheered me up when I needed it most and have helped me to feel at home in Australia. I would like to give special thanks to Andrew Perkins and Margaret Heslewood for their introduction to molecular techniques and assistance in the laboratory. Margaret also provided additional sequences used in this project. Caroline Porter and Adam Marchant generously provided additional support in the molecular laboratory at NSW. Kathi Downs kindly inducted me into the processes and procedures of NSW. Louisa Murray assisted with the management of the loan herbarium material. Zonda Erskine kindly ensured the prompt processing of all loan material upon its arrival at NSW. I v thank Miquel Garcia for helping me to understand the cataloguing system use in the library at NSW and for promptly obtaining important references that I was unable to locate. Thanks to Peter Hind for introducing me to the flora of the Sydney region and for making me feel welcome in Sydney. Lesley Elkan and Catherine Wardrop provided the excellent botanical illustrations and diagrams used in this thesis. Gary Chapple expertly re-formatted the various data matrices exported from PAUP* software into a more suitable format for inclusion in the Appendices. And, to Andrew, thanks for the frequent cups of coffee; to Elizabeth and Chris, a special thanks for allowing me to take-over their computers for hours, days, in fact, for weeks at a time. Esti Ariyanti (Kebun Raya Purwodadi), Barry Conn (NSW), Tjetjep Rayadi (Kebun Raya Bogor), Adjun, Rustandi, Nanang Suryana, (latter three Kebun Raya Cibodas) and Frank Zich (then Kebun Raya Eka Karya, Bali) provided field assistance and companionship on my various field work in Indonesia. Elizabeth Brown, Barry Conn, Andrew Perkins and David Orr and Bruce Casler (both Waimea Valley Audubon Center, Hawaii, USA) kindly provided additional collections for DNA analysis. Ruspandi (Kebun Raya Bogor) assisted with initial identification of field collections. I also thank the Cibodas staff for maintaining a living collection of wild-sourced material of Elatostema for this project and for future study. Dick Brummit, Petra Hoffman (both K), James Reveal (University of Maryland, USA), Nicholas Turland (MO), Peter Wilson (NSW) kindly provided informative comments on the nomenclature of the tribal names of the Urticaceae. I am very grateful for the four-year funded AusAID scholarship, from 1999 until 2003. I wish to thank the staff of the Australian Development Scholarship (ADS) agency for making this possible. I thank the Directors and staff of the following herbaria for access to their collections and/or for providing loan specimens: BO, BRI, CANB, GH, K, KRB, L, LAE, MEL, NSW and NY. vi I am very indebted to Helen and Barry Conn for their continuous support throughout my time in Australia. They welcomed me into their home, provided mental and material support, and assisted me academically and personally during my study and especially during the writing phase of my thesis. They keep encouraging me to remain positive when it all seemed to be too hard and they helped me stay cheerful when I was feeling overwhelmed by what had to be completed. Sometimes I thought that I would not be able to put a smile on my face any more, but they always found a way to make me laugh through the hardest times. They introduced me to ‘NCIS’, ‘Life on Mars’, ‘House’, ‘Medium’, the new ‘Robin Hood’ and Rugby League. I shall never be able to repay them for everything they have done for me. Finally, I would like to thank my beloved family, especially my husband Tansen, for their endless love and support, and for always believing in me. vii viii ABSTRACT Elatostema J.R.Forst. & G.Forst. (Urticaceae) is a taxonomically problematic genus of approximately 300 species that is widespread throughout the tropical, subtropical and sub-temperate regions of Africa through to SE Asia, Australasia to Polynesia. Morphological and molecular analyses were conducted to evaluate the infra-familial classification of the Urticaceae, to test the monophyly of Tribe Elatostemeae, to define generic limits of Elatostema and assess its relationship within the tribe, and to examine the current infrageneric grouping within Elatostema. Phylogenetic analyses based on choloroplast DNA sequences of rbcL and trnL-F do not provide support for the monophyly of Urticaceae, because of the position of Poikilospermum (currently Cecropiaceae) within the tribe Urticeae. Although the status of Cecropiaceae is equivocal, there is support for the inclusion of this family in the Urticaceae, with Cecropia and Coussapoa (Cecropiaceae) having close affinities to the Boehmerieae and Parietarieae. The phylogenetic position of Myriocarpa is unresolved, but is excluded from the Boehmerieae, as currently classified. The Elatostemeae is paraphyletic with Pilea placed sister to the Urticeae. Evaluation of the infrageneric classification of Elatostema, based on phylogenetic analyses of both morphological and molecular data (trn and ITS) does not support the current subgeneric classification as proposed by Schröter and Winkler (1935, 1936). The analyses support two main infrageneric grouping: (1) a group consisting of Elatostema subg. Pellionia and Procris, and (2) a group consisting of the remaining members of Elatostema (including E. griffithianum – subg. Pellionia). The molecular data are regarded as a more accurate estimate of the phylogeny than provided by morphology, with molecular data having a higher Rescaled Consistency Index on the most parsimonious trees, together with a much greater level of resolution and support than that of the morphological analyses. ix x Table of Contents TABLE OF CONTENTS Title Page .................................................................................................................................. i Originality Statement .............................................................................................................iii Acknowledgements .................................................................................................................. v Abstract ................................................................................................................................... ix Table of Contents .................................................................................................................... xi List of Figures ....................................................................................................................... xix List of Tables.......................................................................................................................xxiii Chapter 1 Introduction ........................................................................................................ 1 1.1 The family Urticaceae................................................................................................ 1 1.2 Rationale for project .................................................................................................. 2 1.3 Objectives of project ....................................................................................... 4 Chapter 2 Taxonomic history .............................................................................................. 5 2.1. Taxonomic history of Urticaceae A.L.Juss. noms. cons. ........................................... 5 2.1.1. Early taxonomic history of family........................................................................ 5 2.1.2. Circumscription of tribes of Urticaceae................................................................ 7 2.1.3. Modern taxonomic history of Urticaceae ............................................................. 8 2.2. Phylogeny of Urticaceae ............................................................................................ 9 2.3. Nomenclature of tribe
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