| A comprehensive phylogeny of the genus Kurixalus (Rhacophoridae, Anura) sheds light on the geographical range evolution of frilled swamp treefrogs | |
| Lv YY1,2,3; He K2; Klaus S4; Brown RM5; Li JT*1,3,6; lijt@cib.ac.cn | |
| 2017 | |
| 发表期刊 | Molecular Phylogenetics and Evolution
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| 卷号 | **期号:**页码:Epub ahead of print |
| 摘要 | Currently, the genus Kurixalus comprises 14 species distributed in Southern, Southeast and East Asia. Because of their relatively low dispersal capability and intolerance of seawater, this group may be an ideal one for the study of terrestrial range evolution, especially that portion of its distributional range that extends into the island archipelagos of southern Asia. We assembled a large dataset of mitochondrial and nuclear genes, and estimated phylogeny by maximum likelihood and Bayesian methods, and we explored the history of each species via divergence-time estimation based on fossil-calibrations. A variety of ancestral-area reconstruction strategies were employed to estimate past changes of the species' geographical range, and to evaluate the impact of different abiotic barriers on range evolution. We found that frilled swamp treefrogs probably originated in Taiwan or South Vietnam in the Oligocene. Alternatively, the lineage leading to Kurixalus appendiculatus strongly supports a hypothesis of terrestrial connection between the Indian and Asian continents in the Oligocene. The outcome of both our divergence-time estimates and ancestral-area reconstruction suggests that the divergence between species from Indochina and Taiwan can probably be attributed to the opening of the South China Sea, approximately 33 million years ago. We could not find evidence for dispersal between mainland China and Taiwan Island. Formation of both Mekong and Red River valleys did not have any impact on Kurixalus species diversification. However, coincidence in timing of climate change and availability of plausible dispersal routes from the Oligocene to the middle Miocene, plausibly implied that Kurixalus diversification in Asia resulted from contemporaneous, climate-induced environmental upheaval (Late Oligocene Warming at 29 Ma; Mi-1 glaciation since 24.4 to 21.5 Ma; Mid-Miocene Climatic Optimum at 14 Ma), which alternatively opened and closed dispersal routes. |
| 关键词 | Kurixalus Southeast Asia Biogeography Climate Phylogeny Speciation |
| 资助者 | This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). |
| 收录类别 | SCI |
| 语种 | 英语 |
| 资助者 | This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). ; This study was supported by the National Natural Science Foundation of China (31722049, 31772434, 31372173); the Strategic Priority Research Program of the Chinese Academy of Sciences (XDPB0202); the Southeast Asia Biodiversity Research Institute, Chinese Academy of Science (Y4ZK111B01); the Sichuan Science & Technology Bureau (2016HH0083); the Fund of the State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences (GREKF15-08); the Frontier Science Key Research Programs of CAS (QYZDB-SSW-SMC058) and the Youth Innovation Promotion Association of CAS. SK was supported by the German Research Foundation (DFG KL2378/2-1). |
| 文献类型 | 期刊论文 |
| 条目标识符 | http://ir.kiz.ac.cn/handle/152453/12053 |
| 专题 | 遗传资源与进化国家重点实验室 科研部门_兽类生态与进化（蒋学龙） |
| 通讯作者 | lijt@cib.ac.cn |
| 作者单位 | 1.CAS Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization & Ecological Restoration and Biodiversity Conservation Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences No.9 Section 4, Renmin Nan Road, Chengdu 610041, Sichuan, P.R. China 2.State Key Laboratory of Genetic Resources and Evolution, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, P.R. China 3.Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences, Yezin, Nay Pyi Taw 05282, Myanmar 4.Dept. of Ecology and Evolution, J.W. Goethe-Universität, D-60438 Frankfurt/M., Germany 5.Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045-7561, USA 6.University of Chinese Academy of Sciences, Beijing, 100049, P.R. China |
| 推荐引用方式 GB/T 7714 | Lv YY,He K,Klaus S,et al. A comprehensive phylogeny of the genus Kurixalus (Rhacophoridae, Anura) sheds light on the geographical range evolution of frilled swamp treefrogs[J]. Molecular Phylogenetics and Evolution,2017,**(**):Epub ahead of print. |
| APA | Lv YY,He K,Klaus S,Brown RM,Li JT*,&lijt@cib.ac.cn.(2017).A comprehensive phylogeny of the genus Kurixalus (Rhacophoridae, Anura) sheds light on the geographical range evolution of frilled swamp treefrogs.Molecular Phylogenetics and Evolution,**(**),Epub ahead of print. |
| MLA | Lv YY,et al."A comprehensive phylogeny of the genus Kurixalus (Rhacophoridae, Anura) sheds light on the geographical range evolution of frilled swamp treefrogs".Molecular Phylogenetics and Evolution **.**(2017):Epub ahead of print. |
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