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Boraginaceae Varronia Rupicola (Urb.) Brtton

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Boraginaceae Varronia Rupicola (Urb.) Brtton ORBIT-OnlineRepository ofBirkbeckInstitutionalTheses Enabling Open Access to Birkbeck’s Research Degree output Boraginaceae Varronia rupicola (Urb.) Brtton : bio- geography, systematic placement and conservation genetics of a threatened species endemic to the Caribbean https://eprints.bbk.ac.uk/id/eprint/40179/ Version: Full Version Citation: Hamilton, Martin Allen (2016) Boraginaceae Varronia rupicola (Urb.) Brtton : biogeography, systematic placement and conservation genetics of a threatened species endemic to the Caribbean. [Thesis] (Unpublished) c 2020 The Author(s) All material available through ORBIT is protected by intellectual property law, including copy- right law. Any use made of the contents should comply with the relevant law. Deposit Guide Contact: email Boraginaceae Varronia rupicola (Urb.) Britton Biogeography, systematic placement and conservation genetics of a threatened species endemic to the Caribbean Thesis submitted for the degree of Doctor of Philosophy by Martin Allen Hamilton School of Biological and Chemical Sciences Birkbeck, University of London and Conservation Science Department Royal Botanic Gardens, Kew 11 December 2015 Declaration I hereby confirm that this thesis is my own work and that any materials from other sources used in this work have been fully and appropriately acknowledged. Martin Allen Hamilton 11 December 2015 2 Abstract In the Caribbean region, Varronia rupicola (Boraginaceae) is a medium to large, woody shrub endemic to the Puerto Rican Bank where it is threatened with extinction due to its limited area of occupancy, small populations and on-going threats. The greatest of these is currently loss of suitable habitat through development and degradation. These are caused by human activities that are expected to continue and possibly worsen. The species is also threatened by sea level rise and drought as well as natural disasters, particularly hurricanes and tsunamis. Combined, the effects of anthropogenic and climate change induced threats could push the species to extinction over the coming century. Through interrogation of the findings of cyto-, phylo- and population genetic as well as biogeographical research, it is clear that V. rupicola is a distinct species that is endemic to the islands of Puerto Rico, Vieques and Anegada where five populations were detected. The species has lost genetic diversity in the wild through a reduction in population size with allelic diversity proportional to the size of the population. The five populations were found to have lower than expected levels of heterozygosity as well as significant genetic differentiation and inbreeding. Varronia rupicola plants were found in an extremely limited area of intact habitat (<90 km2) overlying substrates that cover <200 km2 across the three islands. Protected areas contain less than a third (<30 km2) of the remaining intact habitat that supports the species and established ex-situ collections capture less than half of the private alleles found in the wild. An integrated approach to the species conservation is needed to maximise genetic diversity and potentiality allow adaptation of V. rupicola to environmental change and new threats. 3 Table of Contents Declaration .................................................................................................................................... 2 Abstract ......................................................................................................................................... 3 Table of Contents .......................................................................................................................... 4 Index of Tables .............................................................................................................................. 9 Index of Figures ........................................................................................................................... 12 Acknowledgements ..................................................................................................................... 20 Thesis Committee ................................................................................................................... 20 British Virgin Islands ................................................................................................................ 20 Department of Agriculture .................................................................................................. 20 National Parks Trust of the Virgin Islands ........................................................................... 20 Cuba ........................................................................................................................................ 20 Puerto Rico .............................................................................................................................. 20 Department of Natural and Environmental Resources ...................................................... 20 International Institute for Tropical Forestry ....................................................................... 20 University of Puerto Rico .................................................................................................... 20 US Fish and Wildlife Service ................................................................................................ 20 UK ............................................................................................................................................ 20 Fera Science ........................................................................................................................ 20 Royal Botanic Gardens, Kew ............................................................................................... 21 USA .......................................................................................................................................... 21 Fort Worth Zoo ................................................................................................................... 21 Fairchild Tropical Botanical Garden .................................................................................... 21 Herbaria .................................................................................................................................. 22 Previous workers ..................................................................................................................... 22 Funding organisations ............................................................................................................. 22 Fieldwork companions ............................................................................................................ 22 Chapter 1. Introduction .............................................................................................................. 24 1.1 The Puerto Rican Bank ...................................................................................................... 25 4 1.1.1. Physical environment ................................................................................................ 26 1.1.2. Climate ...................................................................................................................... 27 1.1.3. Population and development ................................................................................... 28 1.1.4. Vegetation and flora ................................................................................................. 29 1.2. Caribbean dry forest ........................................................................................................ 30 1.2.1. Dry forest composition and protection .................................................................... 30 1.2.2. Threats to the dry forest ........................................................................................... 31 1.3. Legislation and protected areas ....................................................................................... 35 1.3.1. Legislation ................................................................................................................. 36 1.3.2. Protected areas in Puerto Rico ................................................................................. 37 1.3.3. Protected areas in the British Virgin Islands ............................................................. 39 1.4. Boraginaceae systematics, biogeography and morphology ............................................ 39 1.4.1. Family divisions ......................................................................................................... 39 1.4.2. Subfamily Cordioideae .............................................................................................. 41 1.4.3. The genus Varronia ................................................................................................... 41 1.4.4. Varronia rupicola ...................................................................................................... 47 1.5. Thesis outline ................................................................................................................... 52 Chapter 2: Biogeography of Varronia rupicola ........................................................................... 53 2.1. Introduction ..................................................................................................................... 53 2.1.1. Geology ..................................................................................................................... 56 2.1.2.
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