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Borneo and Indochina Are Major Evolutionary Hotspots for Southeast Asian Biodiversity Mark De Bruyn Bangor University

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Borneo and Indochina Are Major Evolutionary Hotspots for Southeast Asian Biodiversity Mark De Bruyn Bangor University View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Research Online University of Wollongong Research Online Faculty of Science, Medicine and Health - Papers Faculty of Science, Medicine and Health 2014 Borneo and Indochina are major evolutionary hotspots for Southeast Asian biodiversity Mark de Bruyn Bangor University Bjorn Stelbrink Humboldt-Universitat zu Berlin Robert J. Morley Royal Holloway University of London Robert Hall Royal Holloway University of London Gary R. Carvalho Bangor University See next page for additional authors Publication Details de Bruyn, M., Stelbrink, B., Morley, R. j., Hall, R., Carvalho, G. R., Cannon, C. H., van den Bergh, G., Meijaard, E., Metcalfe, I., Boitani, L., Maiorano, L., Shoup, R. & Von Rintelen, T. (2014). Borneo and Indochina are major evolutionary hotspots for Southeast Asian biodiversity. Systematic Biology, 63 (6), 879-901. Research Online is the open access institutional repository for the University of Wollongong. For further information contact the UOW Library: [email protected] Borneo and Indochina are major evolutionary hotspots for Southeast Asian biodiversity Abstract Tropical Southeast Asia harbors extraordinary species richness and in its entirety comprises four of the Earth's 34 biodiversity hotspots. Here, we examine the assembly of the Southeast Asian biota through time and space. We conduct meta-analyses of geological, climatic and biological (including 61 phylogenetic) datasets to test which areas have been the sources of long-term biological diversity in SE Asia, particularly in the pre-Miocene, Miocene and Plio-Pleistocene, and whether the respective biota have been dominated by in situ diversification, immigration and/or emigration, or equilibrium dynamics. We identify Borneo and Indochina, in particular, as major 'evolutionary hotspots' for a diverse range of fauna and flora. While most of the region's biodiversity is a result of both the accumulation of immigrants and in situ diversification, within-area diversification and subsequent emigration have been the predominant signals characterizing Indochina and Borneo's biota since at least the early Miocene. In contrast, colonization events are comparatively rare from younger volcanically active emergent islands such as Java, which show increased levels of immigration events. Few dispersal events were observed across the major biogeographic barrier of Wallace's Line. Accelerated efforts to conserve Borneo's flora and fauna in particular, currently housing the highest levels of Southeast Asian plant and mammal species richness, are critically required. Disciplines Medicine and Health Sciences | Social and Behavioral Sciences Publication Details de Bruyn, M., Stelbrink, B., Morley, R. j., Hall, R., Carvalho, G. R., Cannon, C. H., van den Bergh, G., Meijaard, E., Metcalfe, I., Boitani, L., Maiorano, L., Shoup, R. & Von Rintelen, T. (2014). Borneo and Indochina are major evolutionary hotspots for Southeast Asian biodiversity. Systematic Biology, 63 (6), 879-901. Authors Mark de Bruyn, Bjorn Stelbrink, Robert J. Morley, Robert Hall, Gary R. Carvalho, Charles H. Cannon, Gerrit D. van den Bergh, Erik Meijaard, Ian Metcalfe, Luigi Boitani, Luigi Maiorano, Robert Shoup, and Thomas Von Rintelen This journal article is available at Research Online: http://ro.uow.edu.au/smhpapers/2361 Running head: MAJOR SOUTHEAST ASIAN EVOLUTIONARY HOTSPOTS Title: Borneo and Indochina are Major Evolutionary Hotspots for Southeast Asian Biodiversity Authors: Mark de Bruyn1,*,a, Björn Stelbrink2,*,a, Robert J. Morley3,4,5, Robert Hall3, Gary R. Carvalho1, Charles H. Cannon6,7, Gerrit van den Bergh8, Erik Meijaard9,10,11, Ian Metcalfe12,13, Luigi Boitani14, Luigi Maiorano14, Robert Shoup15, Thomas von Rintelen2 1 Environment Centre Wales, School of Biological Sciences, Bangor University, Deiniol Road, Bangor LL57 2UW, UK 2 Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der Humboldt-Universität zu Berlin, Invalidenstr. 43, 10115 Berlin, Germany 3 Southeast Asia Research Group, Department of Earth Sciences, Royal Holloway University of London, Egham Hill, Egham, TW20 0EX, UK 4 Palynova Limited, 1 Mow Fen Road, Littleport, Cambs, CB6 1PY, UK 5 Niko Asia Ltd, Plaza City View, Jl Kemang Timur 22, Jakarta 12510, Indonesia 6 Department of Biological Sciences, Texas Tech University, Texas 79409-3131, USA 7 Chinese Academy of Sciences, Xishuangbanna Tropical Botanic Garden, Yunnan 666303, P. R. China 8 Centre for Archaeological Science, University of Wollongong, Wollongong, NSW 2522, Australia 9 Borneo Futures Project, People and Nature Consulting International, Country Woods house 306, JL. WR Supratman, Pondok Ranji, Ciputat, Jakarta, 15412, Indonesia 1 10 School of Archaeology & Anthropology, Building 14, Australian National University, Canberra, ACT 0200, Australia 11 School of Biological Sciences, University of Queensland, St Lucia, QLD 4072, Australia 12 Earth Sciences, School of Environmental and Rural Science, University of New England, Armidale, NSW 2351, Australia 13 GEMOC, ARC Centre of Excellence for Core to Crust Fluid Systems, Department of Earth and Planetary Sciences, Macquarie University, NSW 2109, Australia 14 Department of Biology and Biotechnologies "Charles Darwin", Sapienza Università di Roma, viale dell'Università 32, 00185 Rome, Italy 15 Clastic Reservoir Systems, 10700 Richmond Avenue, Suite 325, Houston, Texas 77042, USA *Joint first authors aTo whom correspondence should be addressed: Mark de Bruyn, Molecular Ecology and Fisheries Genetics Laboratory, School of Biological Sciences, Bangor University, Deiniol Road, Bangor LL57 2UW, UK. Tel: +44- 1248-382344. Email: [email protected] Björn Stelbrink, Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung an der Humboldt-Universität zu Berlin, Invalidenstr. 43, 10115 Berlin, Germany. Tel: +49-30-20938430. Email: [email protected] 2 Abstract.−Tropical Southeast Asia harbors extraordinary species richness and in its entirety comprises four of the Earth’s 34 biodiversity hotspots. Here, we examine the assembly of the Southeast Asian biota through time and space. We conduct meta-analyses of geological, climatic and biological (including 61 phylogenetic) datasets to test which areas have been the sources of long-term biological diversity in SE Asia, particularly in the pre-Miocene, Miocene and Plio-Pleistocene, and whether the respective biota have been dominated by in situ diversification, immigration and/or emigration, or equilibrium dynamics. We identify Borneo and Indochina, in particular, as major ‘evolutionary hotspots’ for a diverse range of fauna and flora. While most of the region’s biodiversity is a result of both the accumulation of immigrants and in situ diversification, within-area diversification and subsequent emigration have been the predominant signals characterizing Indochina and Borneo’s biota since at least the early Miocene. In contrast, colonization events are comparatively rare from younger volcanically active emergent islands such as Java, which show increased levels of immigration events. Few dispersal events were observed across the major biogeographic barrier of Wallace’s Line. Accelerated efforts to conserve Borneo’s flora and fauna in particular, currently housing the highest levels of Southeast Asian plant and mammal species richness, are critically required. [phylogenetics; biogeography; geology; palynology; climate change; ecology] 3 Text: Since the early research (1859-1871) of Alfred Russel Wallace, Southeast (SE) Asia has held considerable interest for the study of evolutionary biology and the field of biogeography. Megadiverse SE Asia comprises both mainland and numerous archipelagos of oceanic and continental islands (Lohman et al. 2011). The fragmented nature of SE Asia’s geography is mirrored in the distribution of its extant biota and a high degree of regional and local endemism (e.g., Woodruff 2010; Wong 2011). The four biodiversity hotspots that constitute SE Asia (Indo-Burma, Sundaland, Wallacea and the Philippines; Myers et al. 2000; Woodruff 2010) are under increasing threat from the unsustainable use of native species and their habitats (Koh and Sodhi 2010; Wilcove et al. 2013). Biotic evolution in SE Asia cannot be understood without considering the region’s complex tectonic and climatic evolution (Fig. 1; Fig. S1-S4, Supplementary Information, http://datadryad.org, doi: 10.5061/dryad.67s40), dominated since the late Paleozoic by a series of ongoing continental collisions (Metcalfe 2011a). These geological events have influenced many environmental variables e.g., geomorphology, topography, atmospheric circulation, the hydrological cycle and ocean current patterns, the opening and closure of ocean gateways, and as a consequence climate and climate change (see Fig. 2 for Quaternary sea-level scenarios), either directly or indirectly (Hall 2009; Morley 2012). Present-day SE Asia is the result of more than 300 million years of ‘Colliding Worlds’ (van Oosterzee 1997) characterized by continent-continent and continent-arc collisions (see Supplementary Information for reconstructions of tectonic evolution). SE Asian continental blocks (Fig. S1-S3) were all derived from the Southern Hemisphere supercontinent Gondwana, and travelled north to progressively collide and coalesce prior to the ongoing 4 collision with the northwards-moving Australian continent (Metcalfe 2011a). By the end of the Triassic (c. 200
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