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1 Acta Theriologica Sinica * Q A Shape variation in mandibles of Parascaptor leucura Mammalia Talpidae based on geometric morphometrics implication for geographic diversification HE Kai 1 2 BAI Ming 3 WAN Tao 1 4 LI Quan 1 4 WANG Yingxiang 1 JIANG Xuelong 1* 1 State Key Laboratory of Genetic Resources and Evolution Kunming Institute of Zoology Chinese Academy of Sciences Kunming China 2 Institute of Eastern-Himalaya Biodiversity Research Dali University Dali China 3 Key Laboratory of Zoological Systematics and Evolution Institute of Zoology Chinese Academy of Sciences Beijing China 4 University of Chinese Academy of Sciences Beijing China Abstract Parascaptor is a monotypic genus distributed across Asia including Southwestern China Assam India Bengal and Northern Burma and Laos all of which possess extremely complex and diverse topography climate and habitat. To test whether morphological diversification has taken place in these different environments we examined the mandibles of 49 P. leucura specimens from 18 localities throughout the distribution area. Geometric morphometric analyses were used to examine the shape variations. The specimens were clustered into three groups Northeastern Yunnan group 1 Southern Hengduan Mountain group 2 and Indian groups group 3 congruent with their geographic distribution. We found important distinctions Among these different groups in the horizontal ramus shape such as the length of the lower molar row the curvature of the ventral margin and the relative depth of the horizontal ramus. Among the three groups the specimens from Northeastern Yunnan have the shortest molar row and the thinnest and most flattened horizontal ramus. Our results im- 2011CB CB KSCX2 - EW - G - 4 GREKF11-03 SYSKF * Corresponding author jiangxl@ mail. kiz. ac. cn

2 8 33 ply that the three geographic groups may represent three subspecies or even full species strongly warranting a taxonomic revision using comprehensive methods. Key words Geometric morphometrics Mandible Parascaptor leucura PCA Parascaptor Sorico- morpha Talpidae Talpini P. leucura Chakraborty et al Hutterer 2005 Molur Wang Fig. 1. Ⅰ Ⅱ Map of sample localities. Numbers refer to locality numbers in Appendix Ⅰ and Ⅱ P. leucura Blyth 1850 Sanchez-Villagra 2006 Khasi Hills Mastoid process Talpa Gill 1875 Talpa leucura Parascaptor Parascaptor Allen 1938 Miller 1940 Corbet 1978 Parascaptor cf. leucura 1984 Abe et al Hutterer Talpini Sanchez-Villagra sp et al Geometric morphometrics http / / www. conservation. org / Adams et al. 2004

3 1 9 Zelditch 2004 Mitteroecker and Gunz 2009 Zelditch cm Mitteroecker and Gunz Barrow 90 and Macleod Landmark LM Rohlf et Semi-landmark al tpsutil v1. 46 Rohlf Barrow and Macleod 2008 Dig v2. 15 Rohlf 2010a IOZ KIZ Bookstein AMNH LM1 NMNH LM2 FMNH 49 LM15 LM16 LM23 LM type I LM1 LM2 LM23 6 type Ⅱ type Ⅲ 6 49 LM1 LM2 LM15 LM23 LM24 LM26 Barrow I http / /www. gbif. org / 26 00' ' Zelditch b TPS tps- LM25 LM26 LM27 Semi-landmark Zelditch 2004 Barrow 2008 LM2 LM D300 AF-S LM16 LM23 VR105mm LM1 ~ LM27

4 /. L1 L2 L3 angle 1 Fig. 2 Mandible of Parascaptor leucura lateral. Dots are landmarks /semi-landmarks used in geometric morphometric analyses. L1 L2 L3 and angle1 are measurements used in traditional morphometric analyses PCA group 1 group 2 35 group 3 Deformation Barrow LM1 ~ LM15 LM LM15 ~ LM ~ 2. 3 Ⅱ IMP7 Sheets 2012 PCA LM1 LM23 L1 LM2-LM10-LM15 Angle 1 2 tpsdig v2. Superimposition 15 Procrustes superimposition IMP7 Sheets 2012 Co- One-way ANO- ordgen One-way ANOVA Partial warps scores 2 3 VA 2 3 One-way ANOVA Gen7a P < Thin-plate Spline TPS coefficient Robust Tests of Equality of Means One-way ANO-

5 1 11 VA SPSS v L1 angle1 LM1 ~ LM7 L2 LM23 ~ LM10 L SPSS v % / % PC1 3 PC2 3 PC1 PC PC / PC1 PC2. Fig. 3 Plots of P. leucura on mandible shape in PC1 and PC2 referred from 27 landmarks / semi-landmarks. Deformation grids show the outline shapes at both ends of PC1 and PC2 and mean specimens of each group

6 / PC1 PC2. Fig. 4 Plots of P. leucura on mandible horizontal ramus shape in PC1 and PC2 referred from 16 landmarks / semi-landmarks. Deformation grids show the outline shapes at both ends of PC1 and PC2 and mean specimens of each group 2. 3 > / < % ~ / PC1 PC2. Fig. 5 Plots of P. leucura on mandible ascending ramus shape in PC1 and PC2 referred from 12 landmarks / semi-landmarks. Deformation grids show the outline shapes at both ends of PC1 and PC2 and mean specimens of each group

7 ANOVA 3 3 P = angle1 group ± range = n = 3 group ± 3 group range = n = 26 group ± range n = 3 group ± range n = 26 group ± range = n = 20 P = ANOVA ± range n = 20 3 P < P = Fig. 6 6 a b Length of lower molar row a and curvature of the ventral margin of the horizontal ramus b in each group Ⅱ 89. 3% L1 L2 angle1 > L PC1 PC2 Fig. 7 Plot of P. leucura mandible on principal component factor 1 and 2 based on traditional morphometric analyses

8 Barrow 2008 Tribe Ihle et al Michaux et al Meloro and O Higgins Barrow Mus musculus Klingenberg et al Boell and Tautz m m Hall m Molur Ihle et al Klingenberg http / /www. conservation. org / http / /www. iucnredlist. org / 1989 Sanchez-Villagra Singh 1994 Parascaptor cf. leucura Zhang et al Zhang et al Chen et al Chen et al Liu et al Chen et al

9 BMC Evolutionary Biology and Biology. New York Cambridge University Press Eileen Westwig Darrin Lunde Lawrence Heaney 120. James F. Rohlf P. David Polly Julie Meachen- Samuels Jenny McGuire ond edition. Washington D. Abe H Shiraishi S Arai S A new mole from Uotsuri-jima the Ryukyu Islands. Journal of the Mammalogical Society of Japan Adams D C Rohlf F J Slide D E Geometric morphometrics ten years of progress following the revolution. Italian Journal of Zoology Allen G M The Mammals of China and Mongolia. Part I New York American Museum of Natural History Bai M Yang X K Application of geometric morphometrics in biological researches. Chinese Bulletin of Entomology in Chinese Barrow E Macleod N Shape variation in the mole dentary Talpidae Mammalia. Zoological Journal of the Linnean Society Blyth E Description of a new species of mole Talpa leucura. Journal of Asiatic Society of Bengal Boell L Tautz D Micro-evolutionary divergence patterns of mandible shapes in wild house mouse Mus musculus populations. Bookstein F L Morphometric Tools for Landmark Data Geometry Chakraborty S Sirinivasalu C Sirinivasalu B Pradhan M Nameer P Checklist of insectivores Mammalia Insectivora of South Asia. Zoos Print Journal Chen S D Liu S Y Liu Y He K Chen W C Zhang X Y Fan Z X Tu F Y Jia X D Yue B S Molecular phylogeny of Asiatic Short-Tailed Shrews genus Blarinella Thomas 1911 Mammalia Soricomorpha Soricidae and its taxonomic implications. Zootaxa Chen W Li Y Liu Y Liu S Yue B Complex topographic configuration in the Hengduan Mountains shaped the phylogeographic structure of Chinese white-bellied rats. Journal of Zoology 284 Chen W Liu S Liu Y Hao H Zeng B Chen S Peng H Yue B Zhang X Phylogeography of the large white-bellied rat Niviventer excelsior suggests the influence of Pleistocene Glaciations in the Hengduan Mountains. Zoological Science Corbet G B The Mammals of the Palaearctic Region A Taxonomic Review. London Cornell University Press. Gill T Synopsis of insectivorous mammals. Bulletin of the United States Geological and Geographical Survey of the Territories Hall B K Unlocking the black box between genotype and phenotype cell condensations as morphogenetic modular units. Biology and Philosophy Hutterer R Order Insectivora. In Groves C P E ed. Mammal Species of the World A Taxonomic and Geographic Reference. Sec- C. Smithsonian Institution Press. Hutterer R Order Soricomorpha. In Wilson D E Reeder D A E eds. Mammal Species of the World A Taxonomic and Geographic Reference. Third edition ed. Baltimore John Hopkins University Press. Ihle S Ravaoarimanana I Thomas M Tautz T Z D An analysis of signatures of selective sweeps in natural populations of the house mouse. Molecular Biology and Evolution Jia T Lin A Q Wang R Zhu W L Xiao C H Liu C Y Meng L H Lian X Wang Z K Pilot study of Tupaia belangeri from Yunnan Province based on morphometrics of the skulls and mandibles. Acta Theriologica Sinica in Chinese Jin C Z Qin D G Pan W S Wang Y Zhang Y Q Deng C L Zheng J J Micromammals of the Gigantopithecus fanuna from Sanhe cave Chongzuo Guangxi. Quaternary Sciences in Chinese Klingenberge C P Morphological integration and developmental modularity. Annual Review of Ecology Evolution and Systematics Klingenberg C P Mebus K Auffray J C Developmental integra-

10 16 33 tion in a complex morphological structure how distinct are the modules in the mouse mandible Evolution and Development Li B Y Geomorphologic regionalization of the Hengduan Mountainous region. Journal of Mountain Research in Chinese Li X W Li J A preliminary floristic study on the seed plants from the region of Hengduan Mountain. Acta Botanica Yunnanica in Chinese Liu W Clarke R Xing S Geometric morphometric analysis of the early Pleistocene hominin teeth from Jianshi Hubei Province China. Science China Earth Sciences in Chinese Liu Q Chen P He K Kilpatrick C W Liu S Y Yu F H Jiang X L Phylogeographic study of Apodemus ilex Rodentia Muridae in Southwest China. PLoS ONE 7 e Melorod C O Higgins P Ecological Adaptations of mandibular form in Fissiped Carnivora. Journal of Mammalian Evolution Michaux J Chevret P Renaud S Morphological diversity of Old World rats and mice Rodentia Muridae mandible in relation with phylogeny and adaptation. Journal of Zoological Systematics and Evolutionary Research Miller G S Notes on some moles from southeastern Asia. Journal of Mammalogy Mitteroecker P Gunz P Advances in geometric morphometrics. Evolutionary Biology Molur S Status of South Asian Non-volant Small Mammals Conservation Assessment Management Plan CAMP Workshop Report. Qiu Z D Li C K Hu S J Late Pleistocene micromammal fauna of Sanjiacun Kunming. Vertebrata PalAsiatica in Chinese Rohlf F. 2010a. tpsdig ed. New York SUNY at Stony Brook. Rohlf F. 2010b. tpsutil ed. New York SUNY at Stony Brook. Rohlf F J Loy A Corti M Morphometric analysis of Old World Talpidae Mammalia Insectivora using partial-warp scores. Systematic Biology Sanchez-Villagra M R Horovitz I Motokawa M A comprehensive morphological analysis of talpid moles Mammalia phylogenetic relationships. Cladistics Sheets H IMP-integrated morphometrics package. 7 ed. Buffalo NY. Singh R L India a Regional Geography. Varanasi National Geographical Society of India. Wang Y Z Zhang Z G Insectivores from Sichuan I Erinaceidae and Talpidae. Sichuan Journal of Zoology in Chinese Wang Y X A Complete Checklist of Mammal Species and Subspecies in China A Taxonomic and Geographic Reference. Beijing China Forestry Publishing House. in Chinese Xing S Zhou M Liu W Crown morphology and variation of the lower premolars of Zhoukoudian Homo erectus. Chinese Sci Bull in Chinese Yang H Zhang Z H The geometric morphometric research on the skull of Apodemus draco and Apodemus peninsulae. Sichuan Journal of Zoology in Chinese Zelditch M Geometric Morphometrics for Biologists A Primer. Academic Press. Zhang T C Comes H P Sun H Chloroplast phylogeography of Terminalia franchetii Combretaceae from the eastern Sino-Himalayan region and its correlation with historical river capture events. Molecular Phylogenetics and Evolution Zhang Y H Volis S Sun H Chloroplast phylogeny and phylogeography of Stellera chamaejasme on the Qinghai-Tibet Plateau and in adjacent regions. Molecular Phylogenetics and Evolution Zhao E M Yang D T Amphibians and Reptiles of the Hengduan Mountain Region. Beijing Science Press. in Chinese Ⅰ R. Clark

11 1 17 I Appendix I Specimens examined in this study China 1 Luquan Yunnnan N E KIZ KIZ KIZ Zhongdian Yunnan N E IOZ Dayao Yunnan N E KIZ - DY Binchuan Yunnan N E KIZ KIZ Mt. Ailao Yunnan N E KIZ KIZ Mt. Wuliang Yunnan N E KIZ Yongde Yunnan N E IOZ00024 KIZ KIZ KIZ KIZ KIZ Lancang Yunnan N E KIZ Gongshan Yunnan N E KIZ73229 KIZ73401 KIZ73407 KIZ Bijiang Yunnan N E KIZ Yinjiang Yunnan N E KIZ76274 KIZ76275 KIZ57117 KIZ76260 KIZ Gangfang Yunnan N E AMNH Arunachalpradesh Tibet N E NMNH Burma 13 Kachin Hpare N E AMNH India 15 Naga Hills Assam N E FMNH Karong Assam N E FMNH76242 FMNH Khasi Hills Assam N E FMNH76217 FMNH76219 FMNH76220 FMNH76221 FMNH76225 FMNH76227 FMNH76228 FMNH76230 FMNH76231 FMNH76232 FMNH76233 FMNH76236 FMNH76239 FMNH Lushai Hills Assam N E FMNH76244 FMNH76245 Ⅱ Appendix Ⅱ Specimen information and measurements used in PCA analyses Locality number Specimen catalog L1 L2 L3 Angle 1 Locality number Specimen L1 L2 L3 Angle 1 catalog 1 KIZ KIZ KIZ KIZ KIZ AMNH IOZ17411 * AMNH KIZ - DY NMNH KIZ FMNH KIZ FMNH KIZ FMNH KIZ FMNH76217 * KIZ FMNH IOZ00024 * FMNH76220 * KIZ FMNH KIZ FMNH76225 * KIZ FMNH KIZ FMNH76228 * KIZ FMNH76230 * KIZ * FMNH76231 * KIZ FMNH KIZ73401 * FMNH KIZ FMNH KIZ FMNH KIZ FMNH KIZ76274 * FMNH KIZ FMNH KIZ57117 * # * # * Broken specimens not used in the first and third analyses # ascending ramus deformity

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Dan Buettner / / 39 1 2015 1 Vol. 39 No. 1 January 2015 74 Population Research 80 + /60 + 90 + 90 + 0 80 100028 Measuring and Comparing Population Longevity Level across the Regions of the World Lin Bao Abstract Appropriate