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China's transboundary waters: new paradigms for water and ecological security through applied ecology
He DM*1; Wu RD1; Feng Y1; Li YG1; Ding CZ1; Wang W1; Yu DW*2,3; dmhe@ynu.edu.cn; dougwyu@gmail.com
2014
发表期刊JOURNAL OF APPLIED ECOLOGY
卷号51期号:5页码:1159-1168
合作性质其它
摘要1. China is Asia's most important upstream riparian country, sharing 110 rivers and lakes with 18 downstream countries. Consequently, China's management of transboundary water resources must consider both environmental and geopolitical risks. 

2. The major threats to and conflicts over international rivers in China revolve around biotic homogenisation due to the installation of transport links, water allocation, water pollution, alteration of natural flow patterns and disruption of fisheries due to the installation of hydropower dams, and droughts and floods exacerbated by climate change. Because these problems have an international component, they fall under China's Peaceful Rise strategy, mandating that transboundary conflicts be resolved amicably as part of the overarching goal of increasing regional economic growth with as little conflict as possible. 

3. Science-backed policy is more likely to result in long term, mutually agreeable solutions; the results of applied ecological research have already resulted in a number of mitigation measures, including setting operational thresholds to reduce the downstream impact of dams, designating protected areas along key river stretches where dams cannot be installed (one dam in a critical location has been cancelled), and the installation of terrestrial protected-area networks. 

4. Synthesis and applications. Applied ecology will continue to play an important role in the diagnosis and resolution of environmental threats to China's transboundary waters. More importantly, applied ecology can inform the development of a transboundary environmental compensation mechanism and regional consultative mechanisms that support informed, cooperative decision-making for China and its riparian neighbours.
资助者This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support.
收录类别SCI
语种英语
资助者This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support. ; This study was funded by the National Natural Science Foundation of China (No. U1202232 & No. 31260148) and the National Key Technologies R&D Program of China (No. 2011BAC09B07 & 2013BAB06B03). DY thanks Yunnan Province (20080A001),theChineseAcademy ofSciences(0902281081, KSCX2-YW-Z- 1027), the National Natural Science Foundation of China (31272327, 31170498),theMinistryofScienceandTechnologyofChina(2012FY110800), the University of East Anglia and the State Key Laboratory of Genetic Resources and Evolution at the Kunming Institute of Zoology for support.
WOS记录号WOS:000342851300005
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被引频次：29[WOS]   [WOS记录]     [WOS相关记录]
文献类型期刊论文
条目标识符http://ir.kiz.ac.cn/handle/152453/8166
专题科研部门_生态学与环境保护中心（俞维理）
通讯作者dmhe@ynu.edu.cn; dougwyu@gmail.com
作者单位1.Yunnan Key Laboratory of International Rivers and Transboundary Eco-Security, Yunnan University, 2 North Road of the Green Lake, Kunming, Yunnan 650091, China
2.Ecology, Conservation, & Environment Center (ECEC), Kunming Institute of Zoology, 32 Jiaochang East Road, Kunming, Yunnan 650223, China
3.School of Biological Sciences, University of East Anglia, Norwich Research Park, Norwich, Norfolk NR7TJ, UK
推荐引用方式
GB/T 7714		 He DM*,Wu RD,Feng Y,et al. China's transboundary waters: new paradigms for water and ecological security through applied ecology[J]. JOURNAL OF APPLIED ECOLOGY,2014,51(5):1159-1168.
APA He DM*.,Wu RD.,Feng Y.,Li YG.,Ding CZ.,...&dougwyu@gmail.com.(2014).China's transboundary waters: new paradigms for water and ecological security through applied ecology.JOURNAL OF APPLIED ECOLOGY,51(5),1159-1168.
MLA He DM*,et al."China's transboundary waters: new paradigms for water and ecological security through applied ecology".JOURNAL OF APPLIED ECOLOGY 51.5(2014):1159-1168.
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