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Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

Dietary overlap and co-existence of sympatric wild yak, Tibetan wild ass and Tibetan antelope in Arjin Shan National Nature Reserve, Xinjiang Province, China

Jianbin Shi A D E , Feiying Lu B , Xiaowen Li A , Zihui Zhang B , Xukun Su A , Shikui Dong A , Huadong Xu C and Xiang Zhang C
+ Author Affiliations
- Author Affiliations

A School of Environment, Beijing Normal University, Beijing 100875, China.

B School of Life Sciences, Capital Normal University, Beijing 100066, China.

C Administration Bureau of Arjin Shan National Nature Reserve, Kuerle, Xinjiang 841000, China.

D Department of Geography, University of Western Ontario, London, Ontario, Canada.

E Corresponding author. Email: jbshi@bnu.edu.cn; jianbin.shi@gmail.com

Wildlife Research 43(4) 323-331 https://doi.org/10.1071/WR16045
Submitted: 11 December 2015  Accepted: 25 May 2016   Published: 4 July 2016

Abstract

Context: Wild yak (Bos grunniens), Tibetan wild ass (Equus kiang, also known as kiang) and Tibetan antelope (Pantholops hodgsoni, also known as chiru) have been sympatric on the Tibetan Plateau for numerous generations. There is concern over potential dietary competition among them, particularly in winter when their forage resources are scarcer, but little is known about their dietary interactions.

Aim: We aimed to determine diet composition and dietary overlap of the three ungulate species, and to assess the extent of potential dietary competition among them in order to enhance our understanding of mechanisms underlying their co-existence. We predicted that the three species would widen their choice of forages in winter (December) compared with summer (June–July), and that there would be diet segregation among them, particularly in winter when food resources were scarcer.

Methods: We collected fresh faecal samples of the three species and reference forage samples from the field during our transect line surveys in the summers of 2012 and 2013, and winter of 2012 in Arjin Shan National Nature Reserve (ANNR) of Xinjiang Province, China. Slides of faecal samples were prepared for microhistological analysis, and plant fragments in faecal samples were identified under microscope. Relative percentage frequency of diet plant species, diet breadth and dietary overlap were calculated.

Key results: The three species consumed similar forage plants, but varied greatly in their percentage use of the available forages. Wild yak mainly fed on sedges in summer and graminoids in winter, supplemented with forbs. Kiang consumed mainly forbs in summer and browse in winter, and chiru mainly fed on graminoids and forbs in summer. Wild yak had the widest choice of forages, and kiang had the narrowest one in summer, but both wild yak and kiang became less selective in their food choice in winter, resulting in higher diet breadth. The dietary overlaps among them, as represented by Pianka’s index, were generally low, ranging from 0.26 between chiru and kiang to 0.51 between wild yak and kiang. The dietary overlap between wild yak and kiang decreased from summer (0.51) to winter (0.35), suggesting an increased dietary divergence.

Key conclusion: Our study reveals that the three species are ‘opportunistic feeders’, able to cope with coarse forage resources. There is dietary segregation among them as indicated by low dietary overlap, suggesting risk of potential competition for diets. We suggest that competition may have shaped their dietary divergence, contributing to their co-existence in ANNR.

Implication: Although there is dietary segregation among the three species in ANNR, it is not possible to make definitive conclusions about interspecific competition from these results. Further studies are needed to investigate if there is spatial and habitat segregation among them, which may facilitate their dietary divergence and thus contribute further to their co-existence in ANNR. Continued efforts are also needed to monitor their population dynamics and status of forage resources in order to prevent overgrazing.

Additional keywords: dietary overlap, wild yak, Tibetan wild ass, Tibetan antelope, exploitation competition.


References

Bagchi, S., Mishra, C., and Bhatnagar, Y. V. (2004). Conflicts between traditional pastoralism and conservation of Himalayan ibex (Capra sibirica) in the trans-Himalayan mountains. Animal Conservation 7, 121–128.
Conflicts between traditional pastoralism and conservation of Himalayan ibex (Capra sibirica) in the trans-Himalayan mountains.Crossref | GoogleScholarGoogle Scholar |

Begon, N., Harper, J. L., and Townsend, C. R. (1996). ‘Ecology: Individuals, Populations and Communities.’ (Blackwell Science Ltd.: Oxford.)

Bell, R. H. V. (1971). A grazing ecosystem in the Serengeti. Scientific American 225, 86–93.
A grazing ecosystem in the Serengeti.Crossref | GoogleScholarGoogle Scholar |

Belovsky, G. E. (1986). Generalist herbivore foraging and its role in competitive interactions. American Zoologist 26, 51–69.
Generalist herbivore foraging and its role in competitive interactions.Crossref | GoogleScholarGoogle Scholar |

Bertolino, S., di Montezemolo, N. C., and Bassano, B. (2009). Food-niche relationships within a guild of alpine ungulates including an introduced species. Journal of Zoology 277, 63–69.
Food-niche relationships within a guild of alpine ungulates including an introduced species.Crossref | GoogleScholarGoogle Scholar |

Campos-Arceiz, A., Takatsuki, S., and Lhagvasuren, B. (2004). Food overlap between Mongolian gazelles and livestock in Omnogobi, southern Mongolia. Ecological Research 19, 455–460.
Food overlap between Mongolian gazelles and livestock in Omnogobi, southern Mongolia.Crossref | GoogleScholarGoogle Scholar |

Cao, Y. F., Su, J. P., Lian, X. M., Zhang, T. Z., and Cui, Q. H. (2008). Food habits of Tibetan antelope (Pantholops hodgsoni) in the Kekexili nature reserve. Acta Theriologica Sinica 28, 14–19.

Cao, Y. F., Zhang, T. Z., Lian, X. M., Cui, Q. H., Deng, D. D., and Su, J. P. (2009). Diet overlap among selected ungulates in Kekexili region, Qinghai province. Sichuan Journal of Zoology 28, 49–54.

Cincotta, R. P., Van Soest, P. J., Robertson, J. B., Beall, C. M., and Goldstein, M. C. (1991). Foraging ecology of livestock on the Tibetan Changtang: a comparison of three adjacent grazing areas. Arctic and Alpine Research 23, 149–161.
Foraging ecology of livestock on the Tibetan Changtang: a comparison of three adjacent grazing areas.Crossref | GoogleScholarGoogle Scholar |

Connell, J. H. (1980). Diversity and the coevolution of competitors, or the ghost of the past. Oikos 35, 131–138.
Diversity and the coevolution of competitors, or the ghost of the past.Crossref | GoogleScholarGoogle Scholar |

Daniel, A. D. (1991). Evaluation of microscope technique to determine large herbivore diets. M.Sc. Thesis, New Mexico State University, Las Cruces.

Eltringham, S. K. (1979). ‘The Ecology and Conservation of Large African Mammals.’ (Macmillan Press: London.)

Gordon, I., and Illius, A. W. (1989). Resource partitioning by ungulates on the Isle of Rhum. Oecologia 79, 383–389.
Resource partitioning by ungulates on the Isle of Rhum.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BC3sfnsFShsg%3D%3D&md5=7c21d68d541e7a00cbe72fccabf615bcCAS | 23921404PubMed |

Harris, R. B., and Miller, D. J. (1995). Overlap in summer habitats and diets of Tibetan Plateau ungulates. Mammalia 59, 197–212.
Overlap in summer habitats and diets of Tibetan Plateau ungulates.Crossref | GoogleScholarGoogle Scholar |

Henley, S. R., and Ward, D. (2006). An evaluation of diet quality in two desert ungulates exposed to hyper-arid conditions. African Journal of Range & Forage Science 23, 185–190.
An evaluation of diet quality in two desert ungulates exposed to hyper-arid conditions.Crossref | GoogleScholarGoogle Scholar |

Hofmann, R. R. (1989). Evolutionary step of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system. Oecologia 78, 443–457.
Evolutionary step of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system.Crossref | GoogleScholarGoogle Scholar |

Holechek, J. L. (1982). Sample preparation techniques for microhistological analysis. Journal of Range Management 35, 267–268.
Sample preparation techniques for microhistological analysis.Crossref | GoogleScholarGoogle Scholar |

Holechek, J. L., and Gross, B. D. (1982). Evaluation of different calculation procedures for microhistological analysis. Journal of Range Management 35, 721–723.
Evaluation of different calculation procedures for microhistological analysis.Crossref | GoogleScholarGoogle Scholar |

Holechek, J. L., Vavra, M., and Pieper, R. D. (1982). Botanical composition determination of range herbivore diets: a review. Journal of Range Management 35, 309–315.
Botanical composition determination of range herbivore diets: a review.Crossref | GoogleScholarGoogle Scholar |

Jarman, P. J. (1974). The social organization of antelopes in relation to their ecology. Behaviour 48, 215–267.
The social organization of antelopes in relation to their ecology.Crossref | GoogleScholarGoogle Scholar |

Jenkins, K. J., and Wright, R. G. (1988). Resource partitioning and competition among Cervids in the northern rocky mountains. Journal of Applied Ecology 25, 11–24.
Resource partitioning and competition among Cervids in the northern rocky mountains.Crossref | GoogleScholarGoogle Scholar |

Kasworm, W. F., Irby, L. R., and Ihsle, H. B. (1984). Diets of ungulates using winter ranges in north central Montana. Journal of Range Management 37, 67–71.
Diets of ungulates using winter ranges in north central Montana.Crossref | GoogleScholarGoogle Scholar |

Latham, J. (1999). Interspecific interactions of ungulates in European forests: an overview. Forest Ecology and Management 120, 13–21.
Interspecific interactions of ungulates in European forests: an overview.Crossref | GoogleScholarGoogle Scholar |

Leslie, D. M., Vavra, M., Starkey, E. E., and Slater, R. C. (1983). Correcting for differential digestibility in microhistological analyses involving common coastal forages of the Pacific northwest. Journal of Range Management 36, 730–732.
Correcting for differential digestibility in microhistological analyses involving common coastal forages of the Pacific northwest.Crossref | GoogleScholarGoogle Scholar |

Li, W. D., Zhang, X., and Zhang, H. B. (2013). ‘Comprehensive Scientific Investigation of Arjinshan National Nature Reserve in Xinjiang.’ (Xinjiang People’s Press House.)

Lu, F. Y., Shi, J. B., Zhang, Z. H., Dong, S. K., and Li, X. W. (2015). Survey and distribution of sympatric wild yak, Tibetan wild ass and Tibetan antelope in Arjin Shan National Nature Reserve of Xinjiang, China. Journal of Beijing Normal University (Natural Science) 51, 374–381.

Macandza, V. A., Owen-Smith, N., and Cain, J. W. (2012). Habitat and resource partitioning between abundant and relatively rare grazing ungulates. Journal of Zoology 287, 175–185.
Habitat and resource partitioning between abundant and relatively rare grazing ungulates.Crossref | GoogleScholarGoogle Scholar |

Macarthur, R., and Levins, R. (1967). The limiting similarity, convergence, and divergence of coexisting species. American Naturalist 101, 377–385.
The limiting similarity, convergence, and divergence of coexisting species.Crossref | GoogleScholarGoogle Scholar |

Marchand, P., Redjadj, C., Garel, M., Cugnasse, J.-M., Maillard, D., and Loison, A. (2013). Are Mouflon Ovis gmelini musimon really grazers? A review of variation in diet composition. Mammal Review 43, 275–291.
Are Mouflon Ovis gmelini musimon really grazers? A review of variation in diet composition.Crossref | GoogleScholarGoogle Scholar |

Mukhtar, H. K., and Hansen, R. M. (1983). Fiber effects on microhistological analysis. Journal of Range Management 36, 477–478.
Fiber effects on microhistological analysis.Crossref | GoogleScholarGoogle Scholar |

Mysterud, A. (2000). Diet overlap among ruminants in Fennoscandia. Oecologia 124, 130–137.
Diet overlap among ruminants in Fennoscandia.Crossref | GoogleScholarGoogle Scholar |

Namgail, T. (2006). Winter habitat partitioning between Asiatic ibex and blue sheep in Ladakh, Northern India. Journal of Mountain Ecology 8, 7–13.

Nel, J. A. J. (1978). Habitat heterogeneity and changes in small mammal community structure and resource utilization in the southern Kalahari ecology and taxonomy of African small mammals. In ‘Bulletin of Carnegie Museum of Natural History 6’. (Ed. D. A. Schlitter.) (Carnegie Museum of Natural History: Pittsburgh, PA.)

Noor, A., Habib, B., and Kumar, S. (2013). Habitat selection and niche segregation between chital and nilgai in Keoladeo national park, India. European Journal Zoological Research 2, 1–9.

Pianka, E. R. (1973). The structure of lizard communities. Annual Review of Ecology and Systematics 4, 53–74.
The structure of lizard communities.Crossref | GoogleScholarGoogle Scholar |

Prins, H. H. T., and Olff, H. (1998). Species richness of African grazer assemblages: towards a functional explanation. In ‘Dynamics of Tropical Communities’. (Eds D. M. Newbery, H. H. T. Prins and N. D. Brown.) pp. 449–490. (Blackwell: Oxford.)

Prins, H. H. T., de Boer, W. F., van Oeveren, H., Correia, A., Mafuca, J., and Olf, H. (2006). Co-existence and niche segregation of three small bovid species. African Journal of Ecology 44, 186–198.
Co-existence and niche segregation of three small bovid species.Crossref | GoogleScholarGoogle Scholar |

Putman, R. J. (1996). ‘Competition and Resource Partitioning in Temperate Ungulate Assemblies.’ (Chapman & Hall: London.)

Rivals, F., Solounias, N., and Schaller, G. B. (2011). Diet of Mongolian gazelles and Tibetan antelopes from steppe habitats using premaxillary shape, tooth mesowear and microwear analyses. Mammalian Biology – Zeitschrift fur Saugetierkunde 76, 358–364.

Schaller, G. B. (1998). ‘Wildlife of the Tibetan Steppe.’ (University of Chicago Press: Chicago, IL.)

Schaller, G. B., and Gu, B. Y. (1994). Ungulates in the northwest Tibet. Research & Exploration 10, 265–293.

Shrader, A. M., Owen-Smith, N., and Ogutu, J. O. (2006). How a mega-grazer copes with the dry season: food and nutrient intake rates by white rhinoceros in the wild. Functional Ecology 20, 376–384.
How a mega-grazer copes with the dry season: food and nutrient intake rates by white rhinoceros in the wild.Crossref | GoogleScholarGoogle Scholar |

Shrestha, R., Wegge, P., and Koirala, R. A. (2005). Summer diets of wild and domestic ungulates in Nepal Himalaya. Journal of Zoology 266, 111–119.
Summer diets of wild and domestic ungulates in Nepal Himalaya.Crossref | GoogleScholarGoogle Scholar |

Sietses, D. J., Faupin, G., de Boer, W. F., de Jong, C. B., Henkens, R., Usukhjargal, D., and Batbaatar, T. (2009). Resource partitioning between large herbivores in Hustai national park, Mongolia. Mammalian Biology 74, 381–393.
Resource partitioning between large herbivores in Hustai national park, Mongolia.Crossref | GoogleScholarGoogle Scholar |

Smith, C., Valdez, R., Holechek, J. L., Zwank, P. J., and Cardenas, M. (1998). Diets of native and non-native ungulates in South central New Mexico. The Southwestern Naturalist 43, 163–169.

Sparks, D. R., and Malechek, J. C. (1968). Estimating percentage dry weight in diets using a micro-scopic technique. Journal of Range Management 21, 264–265.
Estimating percentage dry weight in diets using a micro-scopic technique.Crossref | GoogleScholarGoogle Scholar |

Stace, C. A. (1965). Cuticular studies as an aid to plant taxonomy. Bulletin of British Museum of Natural History 4, 1–78.

Stewart, D. R. M. (1967). Analysis of plant epidermis in faeces, a technique for studying the food preferences of grazing herbivores. Journal of Applied Ecology 4, 83–111.
Analysis of plant epidermis in faeces, a technique for studying the food preferences of grazing herbivores.Crossref | GoogleScholarGoogle Scholar |

Stewart, K. M., Bowyer, R. T., Kie, J. G., and Dick, B. L. (2003). Niche partitioning among mule deer, elk, and cattle. Ecoscience 10, 297–302.

Vila, A. R., Borrelli, L., and Martínez, L. (2009). Dietary overlap between Huemul and livestock in Los Alerces national park, Argentina. The Journal of Wildlife Management 73, 368–373.
Dietary overlap between Huemul and livestock in Los Alerces national park, Argentina.Crossref | GoogleScholarGoogle Scholar |

Voeten, M. M., and Prins, H. H. T. (1999). Resource partitioning between sympatric domestic herbivores in the Tarangire region of Tanzania. Oecologia 120, 287–294.

Walter, G. (1991). What is resource partitioning? Journal of Theoretical Biology 150, 137–143.
What is resource partitioning?Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaK3Mzms12nsQ%3D%3D&md5=488d545fb3a160e510dc4dcf4ab30d3cCAS | 1890851PubMed |

Yin, B. F., Huai, H. Y., Zhang, Y. L., Zhou, L., and Wei, W. H. (2007). Trophic niches of Pantholops hodgsoni, Procapra picticaudata and Equus kiang in Kekexili region. Chinese Journal of Ecology 18, 766–770.
| 1:CAS:528:DC%2BD2sXlslOhs7k%3D&md5=650c076b294a46be1234e2caebb7627fCAS |