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Systematics and Rarity of Australia's Tassel-Ferns (Lycopodiaceae : Lycopodiophyta)

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Systematics and Rarity of Australia's Tassel-Ferns (Lycopodiaceae : Lycopodiophyta) ResearchOnline@JCU This file is part of the following reference: Field, Ashley Raymond (2011) Systematics and rarity of Australia's tassel-ferns (Lycopodiaceae : Lycopodiophyta). PhD thesis, James Cook University. Access to this file is available from: http://researchonline.jcu.edu.au/44641/ The author has certified to JCU that they have made a reasonable effort to gain permission and acknowledge the owner of any third party copyright material included in this document. If you believe that this is not the case, please contact [email protected] and quote http://researchonline.jcu.edu.au/44641/ Systematics and rarity of Australia’s tassel-ferns (Lycopodiaceae : Lycopodiophyta) Thesis submitted by Ashley Raymond FIELD BSc (Hons) JCU in July 2011 for the degree of Doctor of Philosophy in the School of Marine & Tropical Biology James Cook University ii Statement of Access I, the undersigned, the author of this thesis, understand that James Cook University will make it available for use within the University Library and will by other means allow access to users in other approved libraries. I place a specific restriction on access to and the distribution of text and images of Chapter 5 of this thesis including the localities of the blue-tassel-fern, Huperzia dalhousieana. All users consulting this thesis will have to sign the following statement: In consulting this thesis I agree not to copy or closely paraphrase it in whole or in part without the written consent of the author; and to make proper public written acknowledgement for any assistance that I have obtained from it. _______________________ Ashley R. Field 15th July 2011 Declaration I declare that this thesis is my own work and has not been submitted in any form for another degree or diploma at any university or other institution of tertiary education. Information derived from the published or unpublished work of others has been acknowledged in the text and a list of references is given. _______________________ Ashley R. Field 15th July 2011 iii Statement of contribution of others Nature of assistance Contribution Contributor Intellectual support Advice on Bayesian Prof Ross Crozier inference analysis Dr Ellen Schluens Cartography assistance Mr Peter Bostock Mrs Adella Edwards Database assistance Mr Ainsley Calladine Mrs Holly Field Editorial assistance Mr Peter Bostock Dr Joseph Holtum Ass Prof Betsy James Prof Michelle Waycott Financial support Laboratory funding Queensland Government Growing the Smart State PhD funding program James Cook University (JCU) Teaching scholarship Prof Michelle Waycott Write-up grant Prof Richard Pearson Infrastructure support Plant growth facilities Mrs Holly Field Indena S.p.A, Milan James Cook University Takarah Gardens, Mackay Townsville Tassel-ferns Molecular analysis Genetic Analysis Facility (JCU) facilities Advanced Analytical Centre (JCU) Molecular Ecology and Evolution Laboratory (JCU) Other support Thesis printing James Cook University (JCU) Mr Ainsley Calladine Photographs, illustrations and figures in this thesis are my own except where specifically acknowledged. iv Dedication I dedicate this thesis to my wife Holly Field. No one has suffered more cuts and scratches or sorer feet and legs for my passions than has the girl who walks with me. I love you Holly, Ashley Acknowledgements I acknowledge and wholeheartedly thank my James Cook University supervisors Joseph Holtum and Michelle Waycott for their academic input, supervision and for providing advice on my manuscripts. In a moment of folly, after many drinks had been consumed, they agreed to supervise my PhD. Poor sods. I would especially like to thank my ‘honorary’ supervisors Peter Bostock and Betsy Jackes for their guidance in botanical Latin and taxomonic nomenclature, Ainsley Calladine for his instructions on DNA laboratory techniques, Faye Christidis and Ellen Schluens for critical review of my molecular methods and the late Ross Crozier for his tutelage on the subject of mixed-model Bayesian inference phylogenetic analysis. I would also like to thank the people who helped me to establish the large collection of living and preserved tassel-ferns and literature that I worked from in this study. They include Arne Anderberg, Dick Anderson, William Archer, Peter Bostock, Jeremy Bruhl, Andi Cairns, Christine Cargill, Darren Crayn, Paddy Dalton, Holly Field, Ray & Helen Field, Andrew Ford, Trevor Ford, Paul Forster, Bruce Fuhrer, Bruce Gray, Joseph Holtum, Garry Kenning, Chee Kin, Rita & Harry Kupke, David & Iris Liddle, Wei-Han Lim, Ray Lloyd, Gerald McCormack, Arthur Murray, Cyril Murray, Keith & Yvonne Nunn, Tony Orchard, Owen & Coral Rawlins, Andrew Smith, Chris & Jenny Smith, Miriam Supuma, Ing Toh, Low-Yee Wen, Jeff Wilkinson, Niklas Wikström, Henk van der Werf, Frank Zich, Sandra Zimmerman and the Ayapathu, Jirball, Kaanju, Kuku Yalanji, Lamalama and Umpila aboriginal people. I especially acknowledge Richard Pearson and Michelle Waycott for together providing a two-year teaching scholarship. This project was supported by a $10,500 research grant from the Queensland Government Growing The Smart State PhD Funding Program for investigation into the rarity and conservation of the blue-tassel-fern. The research was also supported by my wife Holly Field, my parents Ray & Helen Field, and my friends Owen & Coral Rawlins of Townsville Tassel-Ferns and Harry & Rita Kupke of Takarah Gardens. The above people provided many resources including land, a big ‘bush-house’ full of tassel-ferns, quarantine facilities, travel and laboratory funding and the best resource of all – their friendship. The deepest thanks to all of you, Ashley v vi Abstract Tassel-ferns belong to the Lycopodiaceae, a globally distributed family of homosporous, seedles vascular plants including species that exhibit diverse shoot forms relating to their varied habitats. Hypotheses regarding the generic placement and relationships of tassel-ferns have been unstable due to a paucity of morphological characters for systematic analysis which contrasted with relationships uncovered in the first molecular phylogeny of the group. This study investigates the monophyly, placement and global diversification of tassel-ferns (hitherto genus Huperzia Bernh.) using mixed-model Bayesian inference phylogenetic analysis of DNA-sequences from five chloroplast loci in combination with 70 non-molecular characters derived from a large global collection of plants grown under controlled conditions. The phylogenetic hypotheses generated from these analyses support the monophyly of Lycopodiaceae its subfamilies Lycopodioideae and Huperzioideae. These subfamilies exhibit significant synapomorphies that are associated with their differing habits and life-history strategies. The Lycopodioideae are anisotomously branching plants that spread laterally over the terrestrial habitat, contrasting with the Huperzioideae which are isotomously branching plants that have a tufted and generally localised habit. These features are corroborated by anatomical synapomorphies such as stem stele type and microscopic synapomorphies such as spore type and gametophyte types. Within the Huperzioideae are three general morphological and anatomical groups: Phylloglossum drummondii Kunze which is the only wholly deciduous Lycopodiaceae that regenerates from underground tubers, fir-moss type Huperzia s.l. which are disperse by detachable bulbils and tassel-fern type Huperzia s.l. which disperse primarily by spores or via layering of fertile shoot tips. The monophyly of the genus Huperzia s.l. is poorly supported and in several analyses it is rendered paraphyletic with the morphologically divergent Phylloglossum drummondii Kunze being embedded between within it. Based on molecular data, the genus Huperzia s.l. contains a clearly divergent well-supported fir-moss clade and tassel-fern clade. Both are corroborated by morphological synapomorphies that can be identified in the field. Based on the relationships between these major clades, a recommendation is made to continue to recognise the genus Phylloglossum Kunze and to restrict the genus vii Huperzia Bernh to the fir-moss clade and to recognise a third genus, Phlegmariurus Holub, for the species rich tassel-fern clade. Phlegmariurus (tassel-ferns) are predominantly epiphytic and epilithic tropical plants. They form tufted pendulous shoots and exhibit varied specializations related to the epiphytic habit including re-oriented lycophylls, succulence, glaucousness and multibranched strobili. New combinations with the genus Phlegmariurus Holub are proposed for the Australian tassel-fern species H. carinata (Desv. ex Poir.) Trevis., H. dalhousieana (Spring) Trevis., H. filiformis (Sw.) Holub, H. lockyeri (D.L. Jones & B. Gray) Holub, H. marsupiiformis (D.L. Jones & B. Gray) Holub, H. phlegmaria (L) Rothm., H. phlegmarioides (Gaudich.) Rothm., H.prolifera (Blume) Trevis., H. squarrosa (G. Forst.) Trevis. and H. varia (Brown) Trevis. In contrast, Huperzia s.s. (fir-mosses) are terrestrial, temperate and predominantly northern hemisphere plants and only one species, H. australiana (Herter) Holub, is confirmed as occurring in Australia. The Australian tassel-ferns P. carinatus (Desv. ex Poir.) Ching, P. dalhousianus (Spring) A.R.Field comb. nov., P. filiformis (Sw.) W.H.Wagner, P. lockyeri (D.Jones & B.Gray) A.R.Field comb. nov., P. marsupiiformis (D.Jones & B.Gray) A.R.Field comb. nov. and P. phlegmarioides (Gaudich.) A.R.Field comb. nov. are monophyletic. Based on an examination of their type materials, no recommendations
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