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Jane Herbert Phd Thesis @C@A4;0A82@ 0<3 18=6?0>7C =5 ;C?8202404 9DPH 7HSEHSU 0 AKHTLT @VEOLUUHG IQS UKH 3HJSHH QI >K3 DU UKH BPLWHSTLUZ QI @U$ 0PGSHXT (&&* 5VNN OHUDGDUD IQS UKLT LUHO LT DWDLNDENH LP ?HTHDSFK/@U0PGSHXT.5VNNAHYU DU. KUUR.%%SHTHDSFK#SHRQTLUQSZ$TU#DPGSHXT$DF$VM% >NHDTH VTH UKLT LGHPULILHS UQ FLUH QS NLPM UQ UKLT LUHO. KUUR.%%KGN$KDPGNH$PHU%'&&()%(+-, AKLT LUHO LT RSQUHFUHG EZ QSLJLPDN FQRZSLJKU Systematics and biogeography of Myricaceae Jane Herbert A thesis submitted to the University of St Andrews for the degree of Doctor of Philosophy School of Biology University of St Andrews November 2004 Abstract Two molecular phylogenetic studies were undertaken to examine relationships within Myricaceae. Analyses of DNA sequences of the plastid rbcL gene, trnL-Fregion and nuclear ITS region showed the family to be monophyletic. In all analyses Canacomyrica, a monotypic genus endemic to New Caledonian that bears several distinctive features such as staminodes in the female flowers, fell into a well­ supported clade sister to the rest ofMyricaceae. Phylogenetic analyses ofITS and trnL-F sequence data, representing all genera and subgeneric groups, were undertaken using maximum parsimony and Bayesian methods. The following relationships were strongly supported: (Canacomyrica (Comptonia (Myrica, Morella))). The clade containing all species formerly considered to comprise Myrica s.l. was split into two strongly supported clades corresponding to Myrica s.s. and Morella; this finding strengthens the argument for recognition of these as separate genera. Within Morella, two clades corresponded to previously recognized subgenera. Molecular dating analyses were performed using Penalized Likelihood. Close correlations between lineage-specific diversification and major orogenic or climatic events were inferred. This study suggests that much of the diversity in Morella arose during the Neogene and seed-dispersal by birds has been a significant factor in determining the modem distribution. A study of the conservation status of Canacomyrica was conducted using field observations and data from herbarium specimens. This species was found to occur in just eleven fragmented localities: six outside protected areas and three threatened by mining or bush fires. IUCN Red List status of Endangered was recommended. The morphology and ecology of Canacomyrica was studied to enhance knowledge of this poorly known species and provide comparative data for use in a study of the morphology of the entire family. A new classification scheme with keys was presented including, for the first time, Canacomyrica. New combinations in Morella were recommended. Declaration I, Jane Herbert, hereby certify that this thesis, which is approximately 44,000 words in length, has been written by me, that it is a record of work carried out by me and that it has not been submitted in any previous application for a higher degree. Jane Herbert November 2004 Statement I was admitted as a research student to the School of Biology, University of St Andrews, in January 2001 and as a candidate for the degree of PhD in January 2002; the higher study for which this is a record was carried out in the University of St Andrews between 2001 and 2004. Jane Herbert November 2004 Certificate I hereby certify that the candidate has fulfilled the conditions of the Resolution and Regulations appropriate for the degree of PhD in the University of St Andrews and that the candidate is qualified to submit this in application for that degree. Richard J. Abbott Jeff Graves November 2004 Copyright In submitting this thesis to the University of St Andrews I understand that I am giving permission for it to be made available for use in accordance with the regulations of the University Library for the time being in force, subject to any copyright vested in the work not being affected thereby. I also understand that the title and abstract will be published, and that a copy of the work may be made and supplied to any bona fide library or research worker. Jane Herbert November 2004 Acknowledgements I would like to thank my supervisors Richard Abbott and Jeff Graves and the members of staff at the University of St Andrews, past and present, who have assisted me during the course of this project: Pat Willmer, Robert Crawford, Tom Meagher, Alyson Tobin, Claire Peddy, Amanda Gillies, Dave Forbes, Harry Hodge. Staff at the Royal Botanic Garden, Edinburgh have been a great source of help and inspiration to me and I would like to thank them all, in particular: Ian Hedge, Fatima Sales, Michael Moeller, Mark Hughes, Frieda Christie, Louis Ronse Decraene, Toby Pennington, Vanessa Plana, Pete Hollingsworth, Martin Gardner, David Harris, Helen Hoy, Hannah Atkins, Alex Ponge, Greg Kenicer, Olive Hilliard, Bill Burtt, the library staff. I am also grateful to the staff of the herbarium and J odrell Laboratory at the Royal Botanic Gardens, Kew, and particularly Mark Chase whose contribution to part of this project has been invaluable. The curators and staff of the following herbaria have kindly supplied loans of specimens and digital images of types: Royal Botanic Garden, Edinburgh (E); Compton Herbarium, Cape Town (NBG); Harvard University Herbaria (GH); Royal Botanic Gardens, Kew (K); Natural History Museum, London (BM); Noumea Herbarium, New Caledonia (NOD); New York Botanic Garden (NY); Paris (P). I would like to thank particularly, Peter Phillipson (P) for checking many types and John Rouke (NBG) for his kind and inspiring conversation. Thanks are due to the many people who sent plant material to me or permitted the use of their sequence data: Carlos Parra-O. (Universidad Nacional de Colombia), Mike Pirie (NHN, Utrecht), Joe Dahl (Regional Parks Botanic Garden, Berkeley), Chen Zhiduan and Professor Song Ge (Institute of Botany, Beijing), Dorothy Steane (University of Tasmania), John Lavranos. I have corresponded with numerous people who have had interesting things to say about Myricaceae, nitrogen-fixation, lab techniques etc.: Peter Linder (University of Zurich), Richard Cowling (University of Port Elizabeth), Prof. David Ferguson (University of Vienna), Roger Reeves (Massey University), Keith Skene and Janet Sprent (University of Dundee), Joe Pollard (Furman University), Matt Lavin (Montana State University), Rosemary Paull and Bob Hill (Adelaide University). The study was made possible thanks to a NERC studentship. During the project I made a number of excursions to some exotic places with the help of the Russell Trust (University of St Andrews, UK), the Davis Expedition Trust (University of Edinburgh, UK), the Merlin Trust (Kettering, UK), and members of my family. The help of the following people, in the field and on subsequent occasions, is gratefully acknowledged: Tanguy Jaffre (JRD, Noumea, New Caledonia); Joseph Manaute (Province Sud, New Caledonia); the Administration, Province Sud, New Caledonia; the Administration ofthe Western Cape Nature Conservation Board; Roy Ernstzen, Cape Peninsula National Park, South Africa; Sue Brown, Anthony Hitchcock and the staff at Kirstenbosch and the Harold Porter Botanic Garden, South Africa; staff of Medio Ambiente (Tenerife, La Gomera and EI Hierro); Arnoldo Santos Guerra (Jardin de Ac1imatacion de La Orotava, Tenerife). I want to thank my friends at St Andrews, particularly Lynne Birrell, lain Matthews and Clive Nuttman for all their help and kindness. And for everything, but especially for being able to make me smile, I am forever grateful to Simon Gaskell. Many people have been involved in helping me to define the proj ect as it is presented here but I am indebted to Ian Hedge, for sowing the seed of an idea and for being supportive of my endeavours ever since. Jane Herbert November 2004 Dedicated to the memory of my grandmother Iris Bradbury 1918 - 2002 Contents Chapter 1 Introduction ................................................................................................................ 1 Chapter 2 Nuclear and plastid DNA sequences confirm the placement ofthe enigmatic Canacomyrica monticola in Myricaceae .......................................................................................................................... 37 Chapter 3 Distribution, habitat and Red List status of the New Caledonian endemic tree Canacomyrica monticola (Myricaceae) .......................................................................................................................... 62 Chapter 4 Aspects of the morphology and ecology of Canacomyrica .......................................................................................................................... 73 Chapter 5 Myricaceae: a morphological study .......................................................................................................................... 89 Chapter 6 Phylogenetic relationships and biogeography ofMyricaceae based on nuclear and plastid DNA sequences ........................................................................................................................ 113 Chapter 7 New combinations and a new species in Morella (Myricaceae) ........................................................................................................................ 146 Chapter 8 Conclusions and classification scheme for Myricaceae ........................................................................................................................ 152 Appendices 1) Species List ................................................................................... 166 2) Distribution Maps ........................................................................
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