Diversity and endemism of ungulates on the Qinghai-Tibetan Plateau: Evolution and conservation

JIANG Zhigang 1,2 LI Lili 1,2,3 HU Yiming 1,2 HU Huijian 4 LI Chunwang 1,2 PING Xiaoge 1,2 LUO Zhenhua 5

(1.Key Laboratory of Animal Ecology and Conservation Biology, Institute of Zoology, Chinese Academy of Sciences, Beijing 100101)
(2.University of Chinese Academy of Sciences, Beijing 100049)
(3.Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, Yunnan 666303)
(4.Guangdong Institute of Applied Biological Resources, Guangzhou 510260)
(5.School of Life Sciences, Central China Normal University, Wuhan 430079)

【Abstract】The Qinghai-Tibetan Plateau is a unique physical geographical unit, rich in habitat types and species, and a hot spot in biodiversity and global environmental change research. Unique fauna evolves on the Qinghai-Tibetan Plateau. Among these wild animals are the endemic hoofed animals which possess unique characteristics, such as Tibetan antelope (Pantholops hodgsonii), wild yak (Bos mutus), Tibetan gazelle (Procapra picticaudata), Przewalski’s gazelle (P. przewalskii), and white-lipped deer (Przewalskium albirostris). In this study, we explored the following questions: How many ungulate species are there on the Qinghai-Tibetan Plateau? How many endemic ungulate species are there? What is the distribution pattern of these species? What is the status of their survival? What is their conservation status? We firstly identified the geographic boundaries of the Qinghai-Tibetan Plateau. On the plateau, there are 28 ungulate species, 10 of which are endemic to the plateau. The ungulates of the Qinghai-Tibetan Plateau account for 42% of China’s extant ungulate species, whereas the density of the hoofed species per unit area on the plateau is 62% higher than that of the whole country. However, the distribution of ungulate species on the Qinghai-Tibetan Plateau is uneven, and shows a pattern with low species density in the western region but high species density in the eastern part of the plateau, different from the high species density of the endemic ungulates in the hinterland of the plateau. However, the ratio of threatened ungulates on the Qinghai-Tibetan Plateau is high. Among them, 71% of the ungulates are threatened species, 54% are included in CITES Appendix I or II. The Red List Index of ungulate species on the Qinghai-Tibetan Plateau continued to decline from 1998 to 2015, and this trend would be compounded and worsened by the trend of global change, indicating that the living status of ungulates on the plateau is continuously deteriorating. Through four decades of conservation in the country, some important populations and habitats of the ungulates on the plateau are not protected by nature reserves and the newly established Three-River-Source National Park. Therefore, to achieve the win-win goal of human social and economic development and nature conservation on the plateau, we should manage the grassland ecosystems sustainably, save wildlife populations and habitats, and protect biodiversity on the plateau.

【Keywords】 Qinghai-Tibetan Plateau; evolution; fauna; global change; diversity; endemism; Pan-Third Pole; One Belt and One Road;


【Funds】 National Special Basic Research Fund (2013FY110300) Special Found for Biodiversity Conservation of Ministry of Ecology and Environment of China

Download this article


    [1]. (1) Zhao XF (2013) Study in Distribution and Conservation Situation of Gaoligong Mountain B. taxicolor taxicolor. Master’s thesis at Qinghai Normal University in Xining (in Chinese).


    Andrew TS, Xie Y (2009) A Guide to the Mammals of China. Hunan Education Press, Changsha (in Chinese).

    Bubb PJ, Butchart SHM, Collen B, Dublin H, Kapos V, Pollock C, Stuart SN, Vié JC (2009) IUCN Red List Index—Guidance for National and Regional Use. IUCN, Gland, Switzerland.

    Butchart SHM, Walpole M, Collen B, van Strien A, Scharlemann JPW, Almond REA, Baillie JEM, Bomhard B, Brown C, Bruno J, Carpenter KE, Carr GM, Chanson J, Chenery AM, Csirke J, Davidson NC, Dentener F, Foster M, Galli A, Galloway JN, Genovesi P, Gregory RD, Hockings M, Kapos V, Lamarque J-F, Leverington F, Loh J, McGeoch MA, McRae L, Minasyan A, Morcillo MH, Oldfield TEE, Pauly D, Quader S, Revenga C, Sauer JR, Skolnik B, Spear D, Stanwell-Smith D, Stuart SN, Symes A, Tierney M, Tyrrell TD, Vié J-C, Watson R (2010) Global biodiversity: Indicators of recent declines. Science, 328, 1164–1168.

    Cardillo M, Mace GM, Jones KE, Bielby J, Bininda-Emonds ORP, Sechrest W, David L, Orme C, Purvis A (2005) Multiple causes of high extinction risk in large mammal species. Science, 309, 1239–1241.

    Castell JR (2016) Bovids of the World. Princeton University Press, Princeton, New Jersey.

    Chen FH, An CB, Dong GH, Zhang DJ (2017) Human activities, environmental changes, and rise and decline of silk road civilization in Pan-Third Pole region. Bulletin of Chinese Academy of Sciences, 32, 967–975 (in Chinese with English abstract).

    Chen J, Jiang Z, Li C, Ping XG, Cui S, Tang S, Chu H, Liu B (2015) Identification of ungulates used in a traditional Chinese medicine with DNA barcoding technology. Ecology and Evolution, 5, 1818–1825.

    Chen J, Li C, Yang J, Luo Z, Tang S, Li F, Li C, Liu B, Jiang Z (2012) Isolation and characterization of cross-amplification microsatellite panels for species of Procapra (Bovidae; Antilopinae). International Journal of Molecular Sciences, 13, 8805–8818.

    Chen YY, Chen YF, Liu HZ (1996) Studies on the position of the Qinghai-Xizang Plateau region in zoogeographic divisions and its eastern demarcation line. Acta Hydrobiological Sinica, 20, 97–103 (in Chinese with English abstract).

    Cui SP, Luo X, Li CW, Hu HJ, Jiang ZG (2018) Predicting the potential distribution of white-lipped deer using the Max Ent model. Biodiversity Science, 26, 171–176 (in Chinese with English abstract).

    Deng T, Wang XM, Wang SQ, Li Q, Hou SK (2015) Evolution of the Chinese Neogene mammalian faunas and its relationship to uplift of the Tibetan Plateau. Advances in Earth Science, 30, 407–415 (in Chinese with English abstract).

    Dirzol R, Young HS, Galetti M, Ceballos G, Isaac N, Collen B (2014) Defaunation in the Anthropocene. Science, 345, 401–406.

    Duckworth JW, Mac Kinnon J (2008) Capricornis thar. The IUCN Red List of Threatened Species 2008: e.T3816A10096556. http://www.iucnredlist.org/details/3816/0 (accessed on 2018-2-13).

    Duckworth JW, Steinmetz R, Chaiyarat R (2008) Naemorhedus griseus. The IUCN Red List of Threatened Species 2008: e.T14303A4430834. http://www.iucnredlist.org/details/14303/0 (accessed on 2018-2-13).

    Feng ZJ, Cai GQ, Zheng CL (1986) Mammals of Tibet. Science Press, Beijing (in Chinese).

    Gao EH, He JK, Wang ZC, Xu Y, Tang XP, Jiang HS (2017) China’s zoogeographical regionalization based on terrestrial vertebrates. Biodiversity Science, 25, 1321–1330 (in Chinese with English abstract).

    Groves CP, Grubb P (2011) Ungulate Taxonomy. Johns Hopkins University Press, Baltimore.

    Guo SC, Liu JQ, Qi DL, Yang J, Zhao XQ (2006) Taxonomic placement and origin of yaks: Implications from analyses of mtDNA D-loop fragment sequences. Acta Theriologica Sinica, 26, 325–330 (in Chinese with English abstract).

    He J, Kreft H, Gao E, Wang Z, Jiang H (2017) Patterns and drivers of zoogeographical regions of terrestrial vertebrates in China. Journal of Biogeography, 44, 1172–1184.

    Hu J, Jiang Z, Chen J, Qiao H (2015) Niche divergence accelerates the evolution in Asian endemic Procapra gazelles. Scientific Reports, 5, 10069.

    Hu YM, Yao ZJ, Huang ZW, Tian Y, Li HB, Pu Q, Yang DD, Hu HJ (2014) Mammalian fauna and its vertical changes in Mt. Qomolangma National Nature Reserve, Tibet, China. Acta Theriologica Sinica, 34, 28–37 (in Chinese with English abstract).

    IUCN (2017) IUCN Redlist of Threatened Species. http://www.iucnredlist.org/ (accessed on 2017-03-20).

    Jiang Z, Jiang J, Wang Y, Zhang E, Zhang Y, Li L, Cai B, Luo Z, Li C, Ping X, Xie F, Cao L (2016) China’s ecosystems: Overlooked species. Science, 353, 657.

    Jiang ZG, Jiang JP, Wang YZ, Zhang E, Zhang YY, Li LL, Xie F, Cai B, Cao L, Zheng GM, Dong L, Zhang ZW, Ding P, Luo ZH, Ding CQ, Ma ZJ, Tang SH, Cao WX, Li CW, Hu HJ, Ma Y, Wu Y, Wang YX, Zhou KY, Liu SY, Chen YY, Li JT, Feng ZJ, Wang Y, Wang B, Li C, Song XL, Cai L, Zang CX, Zeng Y, Meng ZB, Fang HX, Ping XG (2016a) Red List of China’s Vertebrates. Biodiversity Science 24, 500–551 (in Chinese and in English).

    Jiang ZG, Li LL, Luo ZH, Tang SH, Li CW, Hu HJ, Ma Y, Wu Y, Wang YX, Zhou KY, Liu SY, Feng ZJ, Cai L, Zang CX, Zeng Y, Meng ZB, Ping XG, Fang HX (2016b) Evaluating the status of China’s mammals and analyzing their causes of endangerment through the red list assessment. Biodiversity Science, 24, 552–567 (in Chinese with English abstract).

    Jiang ZG, Liu SY, Wu Y, Jiang XL, Zhou KY (2017) China’s mammalian Diversity (2nd edtion). Biodiversity Science, 25, 886–895 (in Chinese with English abstract).

    Jiang ZG, Ma Y, Wu Y, Wang YX, Feng ZJ, Zhou KY, Liu SY, Luo ZH, Li CW (2015a) China’s mammalian diversity. Biodiversity Science, 23, 351–364 (in Chinese with English abstract).

    Jiang ZG, Ma Y, Wu Y, Zhou KY, Liu SY, Feng ZJ (2015b) China’s Mammals: Diversity and Geographic Distribution. Science Press, Beijing (in Chinese with English summary)

    Lei R, Jiang Z, Hu Z, Yang W (2003) Phylogenetic relationships of Chinese antelopes based on mitochondrial ribosomal RNA gene sequences. Journal of Zoology, 261, 227–237.

    Lei RH, Jiang ZG, Hu ZA, Yang WL (2004) Taxonomic status of Procapra and the classification of Chinese antelopes and gazelles. Acta Zootaxonomica Sinica, 29, 622–627 (in Chinese with English abstract).

    Leslie DM, Schaller GB (2009) Bos grunniens and Bos mutus (Artiodactyla: Bovidae). Mammalian Species, 836, 1–17.

    Li JJ, Han LX, Cao HF, Tian Y, Peng BY, Wang B, Hu HJ (2013) The fauna and vertical distribution of birds in Mount Qomolangma National Nature Reserve. Zoological Research, 34, 531–548 (in Chinese with English abstract).

    Lian X, Li X, Zhou D, Yan P (2012) Avoidance distance from Qinghai-Tibet Highway in sympatric Tibetan antelope and gazelle. Transportation Research Part D, 17, 585–587.

    Lian X, Zhang T, Cao Y, Su J, Thirgood S (2011) Road proximity and traffic flow perceived as potential predation risks: Evidence from the Tibetan antelope in the Kekexili National Nature Reserve, China. Wildlife Research, 38, 141–146.

    Liu J, Luo YH, Li DZ, Gao LM (2017) Evolution and maintenance mechanisms of plant diversity in the Qinghai-Tibet Plateau and adjacent regions: Retrospect and prospect. Biodiversity Science, 25, 163–174 (in Chinese with English abstract).

    Luo Z, Jiang Z, Tang S (2015) Impacts of climate change on distributions and diversity of ungulates on the Tibetan Plateau. Ecological Applications, 25, 24–38.

    Mallon DP, Jiang ZG (2009) Grazers on the plains: Challenges and prospects for large herbivores in Central Asia. Journal of Applied Ecology, 46, 516–519.

    Pan HJ, Yang DD, Qin HH, Zhang L, Jiang K, Hu HJ (2013) Herpetofauna of Mount Qomolangma National Nature Reserve in Tibet, China. Biodiversity Science, 21, 610–615 (in Chinese with English abstract).

    Pan QH, Wang YX, Yan K (2007) A Field Guide to Mammals of China. pp. 420. China Forestry Publishing House, Beijing (in Chinese).

    Ping XG, Li CW, Li CL, Tang SH, Fang HX, Cui SP, Chen J, Wang EG, He YB, Cai P, Zhang Y, Wu YL, Jiang ZG (2018) The distribution, population and conservation status of Przewalski’s gazelle, Procapra przewalskii. Biodiversity Science, 26, 177–184 (in Chinese with English abstract).

    Qiao HJ, Wang XY, Wang W, Luo ZH, Tang K, Huang Y, Yang SN, Cao WW, Zhao XQ, Jiang JP, Hu JH (2018) From nature reserve to national park system pilot: Changes of environmental coverage in the Three-River-Source National Park and implications for amphibian and reptile conservation. Biodiversity Science, 26, 202–209 (in Chinese with English abstract).

    Schaller GB, Rabinowitz A (2004) Species of barking deer (genus Muntiacus) in the eastern Himalayan region. Journal of the Bombay Natural History Society, 101, 442–444.

    Shackleton DM, Lovari S (1997) Classification adopted for the Caprinae survey. In:Wild Sheep and Goats and Their Relatives: Status Survey and Conservation Action Plan for Caprinae (ed. Shackleton DM), pp. 9–14. IUCN/SSC Caprinae Specialist Group, Gland, Switzerland and Cambridge, UK.

    Simpson G (1945) The principles of classification and a classification of mammals. Bulletin American Museum of Nature History, 85, 1–35.

    Sutherland WJ, Butchart SHM, Connor B, Culshaw C, Dicks LV, Dinsdale J, Doran H, Entwistle AC, Fleishman E, Gibbons DW, Jiang Z, Keim B, Le Roux X, Lickorish FA, Markillie P, Monk KA, Mortimer D, Pearce-Higgins JW, Peck LS, Pretty J, Seymour CL, Spalding MD, Tonneijck FH, Gleave RA (2018) A 2018 horizon scan of emerging issues for global conservation and biological diversity. Trends in Ecology & Evolution, 32, 47–58.

    Turghan M, Jiang Z, Groves CP, Yang J, Fang H (2013) Subspecies in Przewalski’s gazelle Procapra przewalskii and its conservation implication. Chinese Science Bulletin, 58, 1897–1905.

    Wang S (1998) China Red Data Book of Endangered Animals: Mammals. Science Press, Beijing (in Chinese with English summary).

    Wang S, Xie Y (2004) China Species Red List Vol. II: Vertebrates Part 2. Science Press, Beijing (in Chinese with English summary).

    Wang XM, Wang Y, Li Q, Tseng ZJ, Takeuchi GT, Deng T, Xie GP, Chang MM, Wang N (2015) Cenozoic vertebrate evolution and paleoenvironment in Tibetan Plateau: Progress and prospects. Gondwana Research, 27, 1335–1354.

    Wang YX (2003) A Complete Checklist of Mammal Species and Subspecies in China: A Taxonomic and Geographic Reference. China Forestry Publishing House, Beijing (in Chinese with English summary).

    Wilson DE, Mittermeier RA (2012) Handbook of the Mammals of the World, Vol. 2. Hoofed Mammals. Lynx Edicions, Barcelona.

    Wilson DE, Reeder DM (2005) Mammal Species of the World: A Taxonomic and Geographic Reference, 3rd edn. John Hopkins University Press, Baltimore, MA.

    Xia WP (1964) Album of China’s Animals: Mammals. Science Press, Beijing (in Chinese).

    Yang J, Jiang Z, Zeng Y, Turghan M, Fang H, Li C (2011) Effect of anthropogenic landscape features on population genetic differentiation of Przewalski’s gazelle: Main role of human settlement. PLoS ONE, 6, e20144.

    Yang Q, Xia L (2008) Tibetan wildlife is getting used to the railway. Nature, 452, 810–811.

    Yao TD, Chen FH, Cui P, Ma YM, Xu BQ, Zhu LP, Zhang F, Wang WC, Ai LK, Yang XX (2017) From Tibetan Plateau to Third Pole and Pan-Third Pole. Bulletin of Chinese Academy of Sciences, 32, 924–931 (in Chinese with English abstract).

    Yin B, Huai H, Zhang Y, Zhou L, Wei W (2006) Influence of the Qinghai-Tibetan railway and highway on the activities of wild animals. Acta Ecologica Sinica, 26, 3917–3923.

    You Z, Jiang Z, Li C, Mallon D (2013) Impacts of grassland fence on the behavior and distribution range of the critically endangered Przewalski’s gazelle on the Qinghai-Tibetan Plateau. Chinese Science Bulletin, 58, 2262–2268.

    Zhang F, Jiang Z, Xu A, Zeng Y, Li C (2013) Recent geological events and intrinsic behavior influence the population genetic structure of the Chiru and Tibetan gazelle on the Tibetan Plateau. PLoS ONE, 8, e60712.

    Zhang RZ (1979) Physical Geography of China: Zoogeography. Science Press, Beijing (in Chinese).

    Zhang RZ (1997) Distribution of Mammalian Species in China. China Forestry Publishing House, Beijing (in Chinese).

    Zhang YL, Li BY, Zheng D (2002) A discussion on the boundary and area of the Tibetan Plateau in China. Geographic Research, 21, 1–8 (in Chinese with English abstract).

This Article



Vol 26, No. 02, Pages 158-170

February 2018


Article Outline


  • 1 Research methods
  • 2 Results
  • 3 Discussion
  • Footnote