Acta Zootaxonomica Sinica,38 ( 4): 679 - 686 ( Oct. 2013) ISSN 1000-0739

DOES PLATE TECTONICS DETERMINE DISTRIBUTIONAL PATTERNS OF EXTANT ? A CASE STUDY USING THE 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

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( 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 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 ( 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 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 heuristic searches for 1 000 bootstrap translocated species were excluded from the analyses: replications and 200 random sequence additions was Lithobates catesbeianus ( Shaw , 1802 ), L. grylio used to estimate the confidence intervals for the ( Stejneger,1901),L. heckscheri ( W right,1924),and recovered nodes. Bootstrapping involved subsets of the O ct. 2013 M eng et al.: Plate tectonics and distributional patterns of animals 681 data because evaluation of the full dataset was an association with the geological data, and in precluded by computational constraints due to the large particular tectonic facies. T his approach allowed for number of taxa and the low ratio of informative areas possible explanations of the distributional patterns of to taxa. All areas were weighted equally. C hinese amphibians. Good geological data were T rees were viewed by DENDRO SC O PE ( Huson shown to be irreplaceable when interpreting et al.,2007 ), downloaded from http: / /www -ab. biogeographic patterns ( Heaney et al.,2005) . Indeed, informatik. uni-tuebingen. de /software /dendroscope. Flessa ( 1980 ) and Riddle ( 2005 ) stated that plate Spearman's rank correlation coefficients were used tectonics, by means of its geographical and to assess the relationship between the number of environmental consequences, influences the organic species and quadrats from the results of the PAE world. In other words,“side soil supports people”. analysis. T his non-parametric test was selected because 3. 2 Chinese amphibian regions and the data were not normally distributed ( Sokal & Rohlf, geological history 1995) . Geological evolution,including that of the plate 3 Results and Discussion elements, was a very complex process in Asia. Primarily three plates - Pacific,Eurasian,and Indian 3. 1 Trees and areas plates - formed C hina,although some micro - plates Data analysis yielded only 76 most parsimonious were also involved. T hese plates amalgamated from trees ( length = 1 158 steps,C I = 0. 3463,RI = between 23. 5 to about 0. 78 M a ( W an,2004) . W e 0. 7969 ) based on 401 amphibian species. A compared the result of the PAE with the pattern of comparison discovered that all trees were very similar. tectonic facies assuming the Archipelago M odel ( Hsü et M apping of the terminals resolved 27 geographic areas al.,1998; Hsü & C hen,1999 ) and tectonic units ( Fig. 1),which were named from A1 to A27 and the ( W an,2004 ) . T he five primary regions for the strict consensus tree was used for further analysis ( Figs C hinese amphibians, including Northeastern, 2a - g) . Northwestern, Southeastern, Southwestern and Nine areas ( A3,A4,A10,A11,A12,A15, C entral regions, roughly corresponded to the A19,A23,and A25) consisted of two quadrats each geological provinces,as detailed below . T he detailed of which had ten,seven,76,45,42,30,two,three, information for the species, genera, and families and one species,respectively. Four areas ( A2,A7, distributed in each region were provided in T able 2, A17,and A24 ) comprised three quadrats and each available online. had seven,42,18,and one species,respectively. T he Northeastern Region contains 50 quadrats in Areas A6,A8,A14,and A27 contained four quadrats two areas ( A1 and A2 ) plus some other quadrats with 61,68,56,and 11 species,respectively. Four ( Figs 1,2b ) . It mainly includes the eastern Inner areas,A18,A22,A5,and A16,were comprised of M ongolia Plateau, Daxing'anling M ountains, and five, six, seven, and ten quadrats, respectively, Northeast C hina Plain. T his region has 15 species and which had three, three, 99, and 49 species, 11 genera ( T able 1 ) . T his biotic region forms the respectively. Areas A13 and A20 were composed of 11 eastern part of the Northwest C hina,Inner M ongolia quadrats with 89 and two species,respectively. Areas and North C hina geological provinces,as defined by A9 and A26 were made up of 14 quadrats with 158 tectonic facies ( Hsü et al.,1998; Hsü & C hen,1999) . and 66 species,respectively. O ne large area,A21, T he tectonics in this region involves the Sino-Korean contained 20 quadrats with six species. T he largest Plate,Ergun Block,Harbin ( Songhuajiang-Nenjiang) area,A1,held 37 quadrats but only 15 species. No Block, Xingkai ( W andashan ) Block, and other statistically significant relationship occurred between the blocks ( W an,2004) . T he first plate belongs to Sino- size of area and the number of corresponding species Korean T ectonic Domain and the others belong to the ( rs = 0. 2872,p = 0. 1753 ); these two variables Peri-Siberian T ectonic Domain. were independent. Bootstrap values were generally T he Northwestern Region contains 52 quadrats low ,as expected given the very low ratio of taxa to including four primary areas ( A19,A20,A21,and areas. T he top three bootstrap values were 100 % A24) plus some others quadrats ( Figs 1,2e,except ( A12),64. 8 % ( A19),and 52. 1 % ( A7) . Q207) . Its range primarily encompasses the T ianshan Although PAE resolved patterns of habitation and and Altai M ountains,and the T arim,T urpan,and species associations,including the 27 areas resolved for Junggar basins. W ith only seven species and four C hinese amphibians ( Fig. 1),this description did not genera ( T able 1),this is the most species-depauperate offer a causative explanation because of the lack of area. It is comprised of two geological provinces: 1) correlated data ( García-Barros et al.,2002; Garzón- the Northwest and North C hina geological provinces; O rdua et al.,2008) . T herefore,as Fattorini ( 2002) advised,the distributional patterns were examined for 682 Acta Zootaxonomica Sinica Vol. 38 No. 4 O ct. 2013 M eng et al.: Plate tectonics and distributional patterns of animals 683

Figs 1 - 2. 1. M ap of amphibian distributional patterns. T wenty-seven areas,A1 to A27,were integrated into five regions based on geological data: Northeastern Region,Northwestern Region,Southeastern Region,Southwestern Region,and C entral Region. 2. M aximum parsimony strict consensus tree of 76 equally parsimonious trees. T he tree depicts shared amphibian species among 294 geographical quadrants in mainland C hina ( tree length 1 158 steps,C I = 0. 3463; RI = 0. 7969) . Asterisks above the lines denote bootstrap values greater than 35 % . 2a. Strict consensus tree. 2b. Northeastern Region including areas A1,A2,and some other quadrats. 2c. Southeastern Region including areas A5 - A15 and some other quadrats. 2d,f. C entral Region including areas A16, A17,A23,A24,and some other quadrats. 2e. Northwestern Region including areas A19,A20,A21,A22,and some other quadrats ( except Q207) . 2g. Southwestern Region including areas A25,A26,and A27.

2 ) the Laurentian /C athaysian Southern and C hen, 1999 ), including the Q iangtang Block, Southwestern M argin geological provinces ( Hsü et al., Gangdise ( Lhasa ) Block, Himalayan Block, and 1998; Hsü & C hen,1999) . T hese are not the only Indian Plate ( W an,2004 ) . Species richness in this matching geological provinces in the atlas of tectonic region is associated with a great environmental facies. However,the origins of the tectonic units are diversity, which, in turn, is associated with the similar to the Northeastern Region where the blocks northward movement of the Indochina Plate. and plates belong to the Peri-Siberian T ectonic Domain T he Southeastern Region has 65 quadrats and the Korean T ectonic Plate, respectively. For including 11 areas ( A5 - A15 ) plus a few quadrats example, the Altay Block, T urpan-Xingxingxia ( Figs 1,2c) . Its range mainly includes the Yangtze Block,Ili-Balchas Block,and other blocks belong to River valley and more southerly regions. T his region Peri-Siberian T ectonic Domain. T he T arim Plate,and is extremely speciose and accounts for 81 % ( 325) of Altun-Dunhuang-Alxa and central W est Kunlun blocks C hina's species and 95 % ( 59 ) of the genera of belong to the Xiyu T ectonic Domain,a transition-type amphibians ( T able 1 ) . T raditionally, C hina is domain that can be classified into Korean T ectonic partitioned into the palearctic and oriental biological Plate of the mid-Proterozoic period ( W an,2004) . realms. T heir boundary corresponds precisely to the Laurasia was formed during the late Early northern boundary of Southeastern Region, a Permian by the amalgamation of blocks from the Peri- conclusion similar to prior findings ( C hen,2004; Siberian T ectonic Domain,e. g. Altay Block,Ili- Stuart et al.,2008; Zhang,2004) . T he tectonic facies Balchas Block, Harbin ( Songhuajiang-Nenjiang ) of the South C hina geological provinces ( Hsü et al., Block,and the main body of the Sino-Korean Plate. 1998; Hsü & C hen, 1999 ) correspond to the T he region is also associated with the T arim Plate and Southeastern Region, including the North Yangtze Altun-Dunhuang-Alxa Block ( Du & T ang, 2009; Plate,C athaysian Plate,and Sanya Block ( Northeast W an,2004) . T his history predates the appearance of Indochina Plate ), among others. T hese plates or the first amphibian fossils from the Late Permian at the blocks belong to the Peri-Yangtze T ectonic Domain foot of T ian M ountains; additional fossils are assigned ( W an,2004 ) . A weakly resolved boundary occurs to the T riassic ( Du & T ang,2009) . T he Northeastern between areas A6 and A8,plus A9,and areas A7 plus and Northwestern regions share only three genera,but A11 - A14 ( Fig. 1) . Areas A15 and A10 form the no species. And the number of species in transition zone. T hese areas roughly correspond with Northwestern Region is less than in Northeastern Yangtze Plate and C athaysian Plate, respectively. Region. T he Northeastern and Northwestern regions Notwithstanding,the distributions of extant amphibian have similar tectonic units yet they do not form a single do not form distinct,exclusive patterns and there is biological region. T he transitional region between much overlap in the distributions of species. C ertainly, them,such as areas A18,A3,and A4 plus some other the distributions reflect geological events, especially quadrats,became a geographic barrier to amphibian those associated with quaternary glacial cycling,which dispersal via geological evolution after amalgamation of occurred after amalgamation of the Yangtze and the plates and blocks. T oday,this barrier consists of C athaysian plates. Q uaternary glacial and interglacial the Loess Plateau and deserts that formed by uplifting cycling changed habitats particularly in the of the Q inghai-T ibetan Plateau. Southeastern Region. During the late Q uaternary, T he Southwestern Region is comprised of 20 river courses changed directions, the continent quadrats including three areas ( A25,A26,and A27) extended eastward and southward from dozens to ( Figs 1,2g ) . It ranges mainly southward from the hundreds kilometers,sea levels rose and fell more than Q inghai-T ibetan Plateau and includes the Gangdise 100 meters, landbridge islands were ephemerally M ountains, Nyainqentanglha Range, and Yarlung reconnected to the mainland,and monsoon patterns Zangbo River. Although it has the smallest size,it changed in response to climatic shifts ( Zhuang et al., contains 68 species and 22 genera ( T able 1 ) . T his 2003) . T hese events certainly affected the distributions region is comprised of tectonic facies associated with of many species of amphibians. C onsequently,any the T ibet geological province ( Hsü et al.,1998; Hsü & 684 Acta Zootaxonomica Sinica Vol. 38 No. 4

Table 1. Area,quadrats and taxonomy of the amphibian regions ( Regions mapped in Figs 1 - 2) . Number Percent in total ( % ) Region Area Q uadrat O rder Family Genus Species Area Q uadrat O rder Family Genus Species

4 31 2 10 24 58 C entral Region 15 11 67 83 39 14

11 65 3 12 59 325 Southeastern Region 41 22 1 1 95 81

3 20 1 6 22 68 Southwestern Region 11 6 33 50 35 17

2 50 2 6 11 15 Northeastern Region 7 17 67 50 18 4

4 52 2 3 4 7 Northwestern Region 15 18 67 25 6 2

27 294 3 12 62 401 T otal pre-Q uaternary pattern may have been disrupted,such biological areas that conform to five geographical as mentioned by Sérsic et al. ( 2011) . regions ( Fig. 1): Northeastern Region,Northwestern C ontaining 31 quadrats, the C entral Region Region,Southeastern Region,Southwestern Region, encompasses four areas ( A16,A17,A22,and A23) and C entral Region. T hese five geographical regions ( Figs 1,2d,2f) . T he region mainly includes the can be viewd as representing C hinese amphibian Sichuan Basin,the southwestern Loess Plateau,and regions. T he biological pattern likely reflects geological the northeastern Q inghai-T ibetan Plateau. Although and environmental evolution. For example, this is the second smallest area,species richness ranks disconnection of the Northeastern and Northwestern second with 58 species and 24 genera ( T able 1) . T he regions is likely the result of the loss of ecological high-density diversity corresponds closely to tectonic continuity by the formation of the Loess Plateau and diversity. In terms of tectonic facies, the C entral deserts due to uplifting of the Q inghai-T ibetan Plateau. Region includes parts of all geological provinces, C omplex amphibian relationships in the Southeastern except the Pacific C hina geological province ( Hsü et Region are likely related to Q uaternary glacial cycling. al.,1998; Hsü & C hen,1999) . T he tectonic structure T he exceptional species richness in the Southwestern corresponds to the junction of Eurasian,Indochina, Region likely reflects the environment diversity that and South C hina plates ( W an,2004) . T he diversity developed when the Indochina Plate moved of extant families ( 10. 83 % of diversity in C hina) and northwards. T he C entral Region is comprised of three genera ( 24. 39 % of diversity in C hina) of amphibians primary tectonic sources and these plates are likely in this region ( T able 1) provides indirect evidence that responsible for the high diversity of amphibian families tectonic history is a driver of species diversity. and genera. 3. 3 Frogs and spiders REFERENCES M eng et al. ( 2008 ) present a PAE analysis of Bain,R. H.,Lathrop ,A.,M urphy,R. W .,O rlov,N. L. C hinese spiders. T he distribution of their C lade C and C uc,H. T . 2003. C ryptic species of a cascade ( C1 - C5 ) is virtually identical to the Southwestern from Southeast Asia: taxonomic revisions and descriptions Region of amphibians. T heir subclade C4 has an of six new species. American Museum Novitates,3 417: 1 extremely similar distribution to amphibian group A9. - 60. T he amphibian subgroup A13 corresponds with Bock, J. H., Bock, C . E. and Fritz, R. J. 1981. : subgroups C5 plus C3 of spiders. Spider clade B4 Biogeography of Illinois reptiles and amphibians a numerical analysis. American Midland Naturalist,106: 258 corresponds with the Northeastern Region amphibian - 270. group. T hese associations suggest that the two groups Brooks,D. R. and van Veller,M . G. P. 2003. C ritique of likely experienced some of the same driving forces in parsimony analysis of endemicity as a method of historical their geographical patterns. Discrepancies between the biogeography. Journal of Biogeography,30: 819 - 825. two studies may owe to completeness of sampling,the C he,J, Zhou, W -W , Hu, J-S et al. 2010. Spiny frogs occurrence of cryptic species,differing biologies,or a ( Paini) illuminate the history of the Himalayan Region and combination of these factors. A synthetic comparison Southeast Asia. Proceedings of the National Academy of Sciences, 107: 13 765 - 13 770. involving many additional groups of species is highly C hen,L 2004. T he precise biogeographic division of Palearctic desirable. and O riental Realm in the East of C hina: based on data of 4 Conclusion amphibians. Zoological Research,25 ( 5): 367 - 377. ( In C hinese) . An analysis of amphibian distributions in C hina C hen,Y-H and Bi,J-F 2007. Biogeography and hotspots of by PAE reveals that the extant taxa occupy 27 amphibian species of C hina: implications to reserve O ct. 2013 M eng et al.: Plate tectonics and distributional patterns of animals 685

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现生生物分布格局与地质板块关系的探讨

孟凯巴依尔1 陶 冶1 Robert W . M urphy 2,3 李枢强1* 1. 中国科学院动物研究所 北京 100101 2. C entre for Biodiversity & C onservation Biology,Department of Natural History,Royal O ntario M useum,T oronto O N,C anada M5S 2C6 3. 中国科学院昆明动物研究所遗传资源与进化国家重点实验室 昆明 650223

摘 要 通过比对地质板块分布格局与现生生物分布格局, 5 个区域与中国大地构造相有很大程度的重叠,同时它们与 探讨板块尤其是微板块对生物分布格局的影响。首先将中国 蜘蛛分布格局也重叠。推断这 5 个区域是中国物种分布的共 大陆进行 2° × 2°的经纬网络设定,总计 294 个样方。根据两 同格局,同时这种分布格局与地质板块的进化历史有密切关 栖类分布信息进行汇总,建立数据表。在此基础上进行特有 系,这种关系不仅仅体现在传统概念的板块之间,也体现在 性简约分析,并与地质格局进行比较。结果表明,中国两栖 微板块。 类的分布包括 27 个分布区,并可进一步汇总为 5 个区域。这

关键词 地理格局,地质历史,特有性简约分析,物种分布. 中图分类号 Q958

* 通讯作者,E-mail: lisq@ ioz. ac. cn