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Estimating the Age of Fire in the Cape Flora of South Africa from an Orchid Phylogeny

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Estimating the Age of Fire in the Cape Flora of South Africa from an Orchid Phylogeny Bytebier, B; Antonelli, A; Bellstedt, D U; Linder, H P (2011). Estimating the age of fire in the Cape flora of South Africa from an orchid phylogeny. Proceedings of the Royal Society of London. Series B, Biological Sciences , 278(1703):188-195. Postprint available at: http://www.zora.uzh.ch University of Zurich Posted at the Zurich Open Repository and Archive, University of Zurich. Zurich Open Repository and Archive http://www.zora.uzh.ch Originally published at: Bytebier, B; Antonelli, A; Bellstedt, D U; Linder, H P (2011). Estimating the age of fire in the Cape flora of South Winterthurerstr. 190 Africa from an orchid phylogeny. Proceedings of the Royal Society of London. Series B, Biological Sciences , CH-8057 Zurich 278(1703):188-195. http://www.zora.uzh.ch Year: 2011 Estimating the age of fire in the Cape flora of South Africa from an orchid phylogeny Bytebier, B; Antonelli, A; Bellstedt, D U; Linder, H P Bytebier, B; Antonelli, A; Bellstedt, D U; Linder, H P (2011). Estimating the age of fire in the Cape flora of South Africa from an orchid phylogeny. Proceedings of the Royal Society of London. Series B, Biological Sciences , 278(1703):188-195. Postprint available at: http://www.zora.uzh.ch Posted at the Zurich Open Repository and Archive, University of Zurich. http://www.zora.uzh.ch Originally published at: Bytebier, B; Antonelli, A; Bellstedt, D U; Linder, H P (2011). Estimating the age of fire in the Cape flora of South Africa from an orchid phylogeny. Proceedings of the Royal Society of London. Series B, Biological Sciences , 278(1703):188-195. Estimating the age of fire in the Cape flora of South Africa from an orchid phylogeny Abstract Fire may have been a crucial component in the evolution of the Cape flora of South Africa, a region characterized by outstanding levels of species richness and endemism. However, there is, to date, no critical assessment of the age of the modern fire regime in this biome. Here, we exploit the presence of two obligate post-fire flowering clades in the orchid genus Disa, in conjunction with a robust, well-sampled and dated molecular phylogeny, to estimate the age by which fire must have been present. Our results indicate that summer drought (winter rainfall), the fire regime and the fynbos vegetation are several million years older than currently suggested. Summer drought and the fynbos vegetation are estimated to date back to at least the Early Miocene (ca 19.5 Ma). The current fire regime may have been established during a period of global cooling that followed the mid-Miocene Climatic Optimum (ca 15 Ma), which led to the expansion of open habitats and increased aridification. The first appearance of Disa species in the grassland biome, as well as in the subalpine habitat, is in striking agreement with reliable geological and palaeontological evidence of the age of these ecosystems, thus corroborating the efficacy of our methods. These results change our understanding of the historical mechanisms underlying botanical evolution in southern Africa, and confirm the potential of using molecular phylogenies to date events for which other information is lacking or inconclusive. 1 Estimating the age of fire in the Cape flora of South Africa from an orchid Formatiert: Englisch (Großbritannien) 2 phylogeny 3 4 1 2,4 3 2 5 Benny Bytebier , Alexandre Antonelli , Dirk U. Bellstedt , and H. Peter Linder Formatiert: Englisch (Großbritannien) 6 7 1Bews Herbarium, School of Biological and Conservation Sciences, University of 8 KwaZulu-Natal, Private Bag X01, Scottsville 3209, Pietermaritzburg, South Africa Formatiert: Englisch (Großbritannien) 9 2 10 Insitute of Systematic Botany, University of Zurich, Zollikerstrasse 107, CH 8008, Formatiert: Englisch (Großbritannien) 11 Zurich, Switzerland 12 3 13 Department of Biochemistry, Stellenbosch University, Private Bag X1, Matieland Formatiert: Englisch (Großbritannien) 14 7602, South Africa 15 16 4Current address: Gothenburg Botanical Garden, Carl Skottsbergs gata 22A, SE 413 17 19, Göteborg, Sweden. 18 19 20 Corresponding author: 21 22 Benny Bytebier, Bews Herbarium, School of Biological and Conservation Sciences, 23 University of KwaZulu-Natal, Private Bag X01, Scottsville 3209 Pietermaritzburg, 24 South Africa. E-mail: [email protected]; Ph: +27 33 260 5145. Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 25 1 26 27 28 Key index words: Cape flora, Orchidaceae, Disa, Palaeoecology 2 29 Abstract 30 31 Fire may have been a crucial component in the evolution of the Cape flora of South 32 Africa, a region characterised by outstanding levels of species richness and endemism. Formatiert: Englisch (Großbritannien) 33 However, there is to date no critical assessment of the age of the modern fire regime 34 in this biome. Here we exploit the presence of two obligate post-fire flowering clades 35 in the orchid genus Disa, in conjunction with a robust, well-sampled, and dated 36 molecular phylogeny, to estimate the age by which fire must have been present. Our 37 results indicate that summer drought (winter rainfall), the fire regime, and the fynbos 38 vegetation are several million years older than currently suggested. Summer drought 39 and the fynbos vegetation are estimated to date back to at least the Early Miocene 40 (~23 19.5 Ma)., whereas tThe current fire regime may have been presentestablished 41 during a period of global cooling that followed by the mMid-Miocene Climatic Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 42 Optimum (~ 15 Ma), which when mean global temperatures led to the expansion of 43 open habitats and increased aridificationreached some 5º C above today’s level. The 44 first appearance of Disa species in the grassland biome, as well as in the subalpine 45 habitat, are in striking agreement with reliable geological and palaeontological 46 evidence of the age of these ecosystems, thus corroborating the efficacy of our 47 methods. These results change our understanding of the historical mechanisms 48 underlying botanical evolution in southern Africa, and confirm the potential of using 49 molecular phylogenies to date events for which other information is lacking or 50 inconclusive. 51 3 52 1. INTRODUCTION 53 54 Fire influences global ecosystem patterns and processes and must have had a 55 pronounced effect on the evolution of these biotas (Bowman et al. 2009; Simon et al. Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 56 2009). Without fire, the distribution of ecosystems around the world would be Formatiert: Englisch (Großbritannien) 57 substantially different (Bond & Keeley 2005; Bond et al. 2005). Vast areas of Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 58 flammable biomes such as Mediterranean ecosystems, savannas and humid grasslands 59 would turn into forest, particularly in Africa (Bond et al. 2005). Although fossil Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 60 charcoal appears in the geological record soon after the emergence of terrestrial plants 61 (Scott 2000; Scott & Glasspool 2006), the age of the current fire regimes is largely Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 62 unknown (Scott 2000), but is presumed to be not much older than 6-8 million years Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 63 (Ma) (Bond et al. 2005). Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 64 65 Palaeoclimatological and palaeoecological reconstructions are mostly based on plant 66 and animal fossil data (e.g. van der Hammen & Hooghiemstra 2000; Jacobs 2004; Formatiert: Englisch (Großbritannien) 67 Jacobs & Herendeen 2004). Although geochemical and geomorphological information Formatiert: Englisch (Großbritannien) 68 is also used (e.g. Diester-Haass et al. 1992; Zachos et al. 2001), this comes mainly Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 69 from marine basins and the link between marine and terrestrial realms is not always 70 clear (Bobe 2006). For southern Africa the Neogene fossil record is remarkably poor Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 71 and does not allow for confident reconstruction of the climate or vegetation (Linder Formatiert: Englisch (Großbritannien) 72 2003), nor the age of the current fire regimes (Bond et al. 2005). Other methodologies Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 73 and independent lines of evidence would thus be welcome to augment our patchy Formatiert: Englisch (Großbritannien) 74 understanding of past ecosystems. Here, we use the evolutionary history of the orchid 75 genus Disa (subfamily Orchidoideae) based on neontological data, to reconstruct past 76 climates and environments. 4 77 78 A reconstruction of the palaeoenvironments of the Cape floristic region (CFR) of 79 southern Africa is particularly interesting since it constitutes, with 9000 species in an 2 80 area of 90 000 km (Goldblatt & Manning 2000; Goldblatt et al. 2005), the second Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 81 most important centre of endemism richness in the world (Kier et al. 2009). Most of Formatiert: Englisch (Großbritannien) Formatiert: Englisch (Großbritannien) 82 this richness is found in fynbos, a shrubland ecosystem similar to chaparral in 83 California and kwongan in southwest Australia, characterised by a Mediterranean 84 climate
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