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Does Plate Tectonics Determine Distributional Acta Zootaxonomica Sinica，38 ( 4): 679 － 686 ( Oct． 2013) ISSN 1000-0739 DOES PLATE TECTONICS DETEＲMINE DISTＲIBUTIONAL PATTEＲNS OF EXTANT ANIMALS? A CASE STUDY USING THE AMPHIBIANS OF CHINA M ENG Kaibayier1 ，T AO Ye1 ，Ｒobert W ． M urphy 2，3 ，LI Shu-Q iang1* 1． Institute of Zoology，Chinese Academy of Sciences，Beijing 100101，China 2． Centre for Biodiversity ＆ Conservation Biology，Department of Natural History，Ｒoyal Ontario Museum，Toronto ON，Canada M5S 2C6 3． State Key Laboratory of Genetic Ｒesources and Evolution，Kunming Institute of Zoology，Chinese Academy of Sciences，Kunming 650223，China Abstract C hinese amphibians were selected to investigate the relationship between distributional patterns of animals and tectonic patterns，especially tectonic facies． C hina was initially divided into 294 quadrats of 2° latitude by 2° longitude． Distributional occurrences for 401 species of C hinese amphibians were summarized for each quadrate． Parsimony analysis of endemicity ( PAE) was used to classify the 294 quadrats based on the shared distributional patterns． T he analysis identified 27 areas for the extant amphibians． A comparison these areas with geological patterns identified five regions: Northeastern Ｒegion，Northwestern Ｒegion，Southeastern Ｒegion，Southwestern Ｒegion，and C entral Ｒegion． T hese five distributional patterns were closely associated with the geological evolution of C hina，and with patterns for spiders． T his association suggests that common continental patterns of speciation may be associated with geological history． Key words Geographic patterns，geological history，PAE，species distributions． 1 Introduction During the 20th century，the concept of tectonic facies ( tectonic-based physiographic structure ) was An ancient C hinese proverb states “side soil developed by Hsü ( 1991) and Huang et al． ( 2008) ． supports people ”． In English， this translates to T he concept is an effective platform for analyzing “unique features of a local environment provide formations and identifying plates． For example， important attributes to the region's inhabitants”． T his， Ｒobertson ( 1994) tested the concept in relation to the of course，is a major aspect of the evolution of species． T ethys Sea in the Eastern M editerranean area． Hsü T o a great extent， the distributions of extant and C hen ( 1999) produced a geological atlas of C hina organisms have been influenced by historical using the concept． Xiao et al． ( 2000) investigated the geography and this association has been recognized for W est Kunlun M ountains． centuries ( Hou et al．，2011) ． Abraham O rtelius first Based on tectonic facies，the Bangong-Nujiang- put continental drift forward in 1596，although his Great Southeast Asia suture zone forms the boundary concept largely laid idol until developed by Alfred between the Northern Laurentian /C athaysian realm W egener in 1912． Patterns of life formed some of the and the Southern Gondwanan realm． T hese strongest early evidence for continental drift． In the paleogeographical realms contain six geological early 1960s，the revelation of plate tectonics started a provinces and 180 map units． M ost of the map units revolution in the earth sciences ( Kious ＆ T illing， correspond to tectonic facies， although some are 1996) ． In turn，it profoundly affected biogeographical tectonic or tectono-stratigraphic units that simply do research，especially in promoting the development of not fit into a single category of facies． T he geological the theory of vicariance biogeography ( Humphries ＆ provinces that correspond to tectonic facies are as Parenti，1999) ． Nowadays，plate tectonics forms the follows: Ⅰ ) Laurentian /C athaysian Southern and backbone of biogeography ( Ｒiddle，2005 ) ． Good Southwestern M argin; Ⅱ) Northwest C hina，Inner geological data are irreplaceable for interpreting M ongolia and North C hina; Ⅲ) C entral C hina; Ⅳ) biogeographical patterns ( Heaney et al．，2005) and yet South C hina; Ⅴ) T ibet; Ⅵ) Pacific C hina ( Hsü ＆ ( all geological history is not written in stone Lindell et C hen 1999 ) ． T hese tectonic facies also have ， ; ， ; ， al． 2006 Ｒiddle et al． 2008 U pton ＆ M urphy corresponding tectonic units as assigned by W an 1997) ． * Corresponding author，E-mail: lisq@ ioz． ac． cn This research was supported by the National Natural Sciences Foundation of China ( NSFC-30970336) and by a Visiting Professorship for Senior International Scientists from the Chinese Academy of Sciences to Ｒ． W． M． Manuscript preparation was supported by the National Sciences and Engineering Ｒesearch Council of Canada ( Discovery Grant A3148 to Ｒ． W． M． ) ． Ｒeceived 20 Aug． 2013，accepted 11 Oct． 2013． 679 书 680 Acta Zootaxonomica Sinica Vol． 38 No． 4 ( 2004) ． Ｒhinella marina ( Linnaeus， 1758 ) ． Further， the Because geography often strongly affects the distribution of Polypedates colletti ( Boulenger，1890) was distribution of species，it is possible that tectonic faces not within the geographic range of analysis and the correspond to the distributions of organisms，especially distribution of Odorrana sinica ( Ahl，1927 ) remained those with relatively low dispersal abilities． T o test this uncertain． T herefore，our analyses were restricted to possibility，herein we explore the relationships between only 401 species ( T able of data can be obtained from tectonic facies，tectonic units，and the distributional author) ． patterns of C hinese amphibians． 2. 2 Data compilation 2 Materials and Methods T he study area was constrained to continental C hina and it excluded islands in the South C hina Sea． 2. 1 Study organisms C hina was divided into 294 quadrats of 2° latitude by Extant amphibians ( Lissamphibia) are a highly 2° longitude without consideration of physiographical successful group of tetrapods with diverse body plans features． Distributional data were summarized in these that differ in modes of locomotion， reproductive quadrats． Presence of a species in a quadrat was scored specializations，and life histories ( Duellman ＆ T rueb， ‘1’and absence as ‘0’． Geological data were taken 1994; Pough et al．，1998; Zardoya ＆ M eyer，2001) ． from W an ( 2004)，Hsü and C hen ( 1999)，and the For example，the typical living salamander ( O rder Editing C ommittee of A Dictionary of Earth Sciences C audata) has a slender body with a well-developed tail ( 2006) ． and proportionally paired limbs． W hereas modern caecilians ( Gymnophiona ) are completely limbless， 2. 3 Data analysis and most of them are adapted to a fossorial lifestyle in T he distributional data were subjected to a having elongated bodies， reduced eyes， and parsimony analysis of endemicity ( PAE )， which compacted skulls，extant frogs ( Anura) lack tails and encapsulated the natural patterns of biotic distributions have powerful hind limbs， a shortened， stiffened ( Ｒosen，1988; Ｒosen ＆ Smith，1988 ) ． T his was vertebral column， and an urostyle， all unique analogous to a cladistic analysis where areas were adaptations for jumping ( Zardoya ＆ M eyer，2001) ． equivalent to taxa and the occurrences or absence of Amphibian distributions are usually associated with species formed the characters． Geographic areas were given geographical units because the species do not grouped by the occurrence of shared species，and then migrate and tend to be philopatric ( M ann et al．，1991; they were superimposed onto the map． T he cladogram Poynton ＆ Boycott， 1996; Zhang， 2004 ) ． was rooted using a hypothetical area void of taxa C onsequently， amphibians are highly useful in ( Brooks ＆ van Veller， 2003; C racraft， 1991; reconstructing the biogeographical history of a region Geraads，1998; M orrone ＆ C risci，1995; M orrone ＆ ( M orrone ＆ C risci，1995; W iley，1988; Zhang， Escalante， 2002; Porzecanski ＆ C racraft， 2005; 2004 ) ． In this respect， biogeographic studies on Ｒosen，1988; Ｒosen ＆ Smith， 1988; W aggoner， amphibian species are significant ( Bock et al．，1981; 1999 ) ． Although PAE could have been used to C he et al．，2010; Hanlin et al．，2000; Zhang et al．， provide a summary of localities，areas of endemism， 2010) ． or quadrats ( C risci et al．，2003)，herein it was applied C hina is one of the richest areas in terms of to combined quadrats only． T his approach was amphibian diversity ( C hen ＆ Bi 2007) ． Having about deemed to be the most objective use of PAE． 407 species ( Fei，1999; Frost，2009 )，C hina ranks T he taxon x area data matrix was analyzed using fifth behind Brazil ( 731 species )，C olumbia ( 698 PAU P* v. 4. 0b10 for PC ( Swofford，1998 ) ． W e species)，Ecuador ( 447 species )，and Peru ( 398 conducted maximum parsimony ( M P) analyses using species) ． W ith 215 endemic species，C hina also ranks the heuristic search mode and the tree bisection- fifth behind Brazil ( 467 species )，C olumbia ( 336 reconnection ( T BＲ ) swapping algorithm． T o species)， M exico ( 233 species )， and M adagascar maximize the efficacy of searching， given the low ( 221 species)( Xie et al．，2006) ． number of areas relative to the number of taxa，we W e compiled distributional data for all species of used at least 1 000 random taxon addition replicates for frogs and salamanders based on the summaries of Fei each search． T his procedure maximized the probability ( 1999)，Fei et al． ( 2007)，Frost ( 2009)，and more that all islands of most parsimonious clusters would be specific studies，such as Bain et al． ( 2003) and Zhao et discovered ( M addison，1991; Page，1993; Ｒice et al．， al． ( 2004) ． W e used a cut-off date of Dec． 2009 for 1997) ． Bootstrapping ( Felsenstein，1985; Sanderson， compiling the data． T he following non-native， 1995 ) with
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