Anti-predator strategies of blue sheep (naur) under varied predator compositions: a comparison of snow leopard-inhabited valleys with and without wolves in Nepal
Kamal Thapa A * and Santosh Rayamajhi AA
Handling Editor: Alexandra Carthey
Abstract
In Nepal, naur are usually the staple wild prey for the snow leopard, a solitary stalker hunter, and in some cases, for the wolf who hunts in a pack. We assumed that naur would adapt their anti-predatory responses to the presence of chasing and ambushing predators in the Manang Valley, where there are snow leopards and wolves, and in the Nar Phu valley, an area where there is only the snow leopard.
The aim of this study was to determine if there were differences in anti-predator strategies (vigilance, habitat selection and escape terrain) of naur in two valleys over two seasons, spring and autumn.
In spring 2019, we conducted a reconnaissance survey on the status of the naur and its habitat in the Manang and Nar Phu valleys of the Annapurna Conservation Area, Nepal. In spring and autumn 2020 and 2021, we observed 360 focal naur individuals (180 individuals in each valley), using the vigilance behaviour methodology to examine the behaviour of the naur.
There was little difference in the size of the naur groups between the Manang and Nar Phu valleys. The naur were twice as vigilant in Manang (15%), where there are snow leopards and wolves, as they were in Nar Phu (9%), with only snow leopards. The distance from the naur to escape cover was significantly shorter in Manang than in Nar Phu valley. Naur used significantly more rolling terrain in Nar Phu than in Manang.
The return of wolves to the Manang valley may have resulted in an increase in the level of naur vigilance. Most likely, the wolves in Manang have already had an effect on the female-to-young-ratio, and this effect will possibly have important consequences for the naur population, as well as at the ecosystem level in the future. Other key determining factors, such as the climate crisis and changes in local resources, could have a significant impact on the naur population, indicating the need for more research.
The findings of this study would provide valuable baseline information for the design of a science-based conservation strategy for conservation managers and scientists on naur, snow leopards and wolves.
Keywords: Annapurna conservation area, antipredator behavior, blue sheep (Naur), predation, prey predator traits, snow leopard, trade-off, wolf.
References
Ale SB, Brown JS (2007) The contingencies of group size and vigilance. Evolutionary Ecology Research 9(8), 1263-1276.
| Google Scholar |
Ale SB, Brown JS (2009) Prey behavior leads to predator: a case study of the Himalayan tahr and the snow leopard in Sagarmatha (Mt. Everest) National Park, Nepal. Israel Journal of Ecology and Evolution 55(4), 315-327.
| Crossref | Google Scholar |
Bailey I, Myatt JP, Wilson AM (2013) Group hunting within the Carnivora: physiological, cognitive and environmental influences on strategy and cooperation. Behavioral Ecology and Sociobiology 67(1), 1-17.
| Crossref | Google Scholar |
Beauchamp G (2016) Function and structure of vigilance in a gregarious species exposed to threats from predators and conspecifics. Animal Behaviour 116, 195-201.
| Crossref | Google Scholar |
Berger J (1978) Group size, foraging, and antipredator ploys: an analysis of bighorn sheep decisions. Behavioral Ecology and Sociobiology 4, 91-99.
| Crossref | Google Scholar |
Brown JS (1988) Patch use as an indicator of habitat preference, predation risk, and competition. Behavioral Ecology and Sociobiology 22(1), 37-47.
| Crossref | Google Scholar |
Brown JS, Alkon PU (1990) Testing values of crested porcupine habitats by experimental food patches. Oecologia 83(4), 512-518.
| Crossref | Google Scholar | PubMed |
Brown JS, Morgan RA, Dow BD (1992) Patch use under predation risk: II. A test with fox squirrels, Sciurus niger. Annales Zoologici Fennici 29, 311-318 Available at https://www.jstor.org/stable/23735634.
| Google Scholar |
Brown JS, Laundré JW, Gurung M (1999) The ecology of fear: optimal foraging, game theory, and trophic interactions. Journal of Mammalogy 80(2), 385-399.
| Crossref | Google Scholar |
Chetri M, Odden M, Devineau O, Wegge P (2019) Patterns of livestock depredation by snow leopards and other large carnivores in the Central Himalayas, Nepal. Global Ecology and Conservation 17, e00536.
| Crossref | Google Scholar |
Elgar MA (1989) Predator vigilance and group size in mammals and birds: a critical review of the empirical evidence. Biological Reviews 64(1), 13-33.
| Crossref | Google Scholar | PubMed |
Endler JA (1978) A predator’s view of animal color patterns. In ‘Evolutionary biology’. (Eds MK Hecht, WC Steere, B Wallace) pp. 319–364. (Springer: Boston, MA) doi:10.1007/978-1-4615-6956-5_5
Filla M, Lama RP, Ghale TR, Signer J, Filla T, Aryal RR, Heurich M, Waltert M, Balkenhol N, Khorozyan I (2021) In the shadows of snow leopards and the Himalayas: density and habitat selection of blue sheep in Manang, Nepal. Ecology and Evolution 11(1), 108-122.
| Crossref | Google Scholar | PubMed |
Fowler CW (1999) Management of multi-species fisheries: from overfishing to sustainability. ICES Journal of Marine Science 56(6), 927-932.
| Crossref | Google Scholar |
Fox JL, Chundawat RS (1988) Observations of snow leopard stalking, killing, and feeding behavior. Mammalia (Paris) 52(1), 137-140.
| Google Scholar |
Fox JL, Sinha SP, Chundawat RS, Das PK (1991) Status of the snow leopard Panthera uncia in Northwest India. Biological Conservation 55(3), 283-298.
| Crossref | Google Scholar |
Frid A (1997) Vigilance by female Dall’s sheep: interactions between predation risk factors. Animal Behaviour 53(4), 799-808.
| Crossref | Google Scholar |
Hamilton WD (1971) Geometry for the selfish herd. Journal of Theoretical Biology 31(2), 295-311.
| Crossref | Google Scholar | PubMed |
Hogstad O (1988) Advantages of social foraging of Willow Tits Parus montanus. Ibis 130(2), 275-283.
| Crossref | Google Scholar |
Hugie DM, Dill LM (1994) Fish and game: a game theoretic approach to habitat selection by predators and prey. Journal of Fish Biology 45, 151-169.
| Crossref | Google Scholar |
Hunter LTB, Skinner JD (1998) Vigilance behaviour in African ungulates: the role of predation pressure. Behaviour 135(2), 195-211.
| Crossref | Google Scholar |
Jarman PJ (1974) The social organisation of antelope in relation to their ecology. Behaviour 48(1–4), 215-267.
| Crossref | Google Scholar |
Kachel S, Bayrakcısmith R, Kubanychbekov Z, Kulenbekov R, McCarthy T, Weckworth B, Wirsing A (2023) Ungulate spatiotemporal responses to contrasting predation risk from wolves and snow leopards. Journal of Animal Ecology 92, 142-157.
| Crossref | Google Scholar |
Kotler BP, Holt RD (1989) Predation and competition: the interaction of two types of species interactions. Oikos 54, 256-260.
| Crossref | Google Scholar |
Kotler BP, Gross JE, Mitchell WA (1994) Applying patch use to assess aspects of foraging behavior in Nubian ibex. The Journal of Wildlife Management 58, 299-307.
| Crossref | Google Scholar |
Lama RP, Ghale TR, Regmi GR, Suwal MK, Lama T (2017) Return of the wolf to the Nyesyang Valley, Manang District, Nepal. Canid Biology and Conservation 20(7), 28-31.
| Google Scholar |
Laundré JW, Hernández L, Altendorf KB (2001) Wolves, elk, and bison: reestablishing the “landscape of fear” in Yellowstone National Park, U.S.A. Canadian Journal of Zoology 79(8), 1401-1409.
| Crossref | Google Scholar |
Lima SL (1995) Back to the basics of anti-predatory vigilance: the group-size effect. Animal Behaviour 49(1), 11-20.
| Crossref | Google Scholar |
Lima SL, Dill LM (1990) Behavioral decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology 68(4), 619-640.
| Crossref | Google Scholar |
Lipetz VE, Bekoff M (1982) Group size and vigilance in pronghorns. Zeitschrift für Tierpsychologie 58(3), 203-216.
| Crossref | Google Scholar |
Lovari S, Boesi R, Minder I, Mucci N, Randi E, Dematteis A, Ale SB (2009) Restoring a keystone predator may endanger a prey species in a human-altered ecosystem: the return of the snow leopard to Sagarmatha National Park. Animal Conservation 12(6), 559-570.
| Crossref | Google Scholar |
Michelena P, Deneubourg J-L (2011) How group size affects vigilance dynamics and time allocation patterns: the key role of imitation and tempo. PLoS ONE 6(4), e18631.
| Crossref | Google Scholar | PubMed |
Oli MK (1994) Snow leopards and blue sheep in Nepal: densities and predator: prey ratio. Journal of Mammalogy 75(4), 998-1004.
| Crossref | Google Scholar |
Oli MK (1997) Winter home range of snow leopards in Nepal. Mammalia-Paris 61, 355-360.
| Google Scholar |
Oli MK, Rogers ME (1996) Seasonal pattern in group size and population composition of blue sheep in Manang, Nepal. The Journal of Wildlife Management 60, 797-801.
| Crossref | Google Scholar |
Oli MK, Taylor IR, Rogers DME (1993) Diet of the snow leopard (Panthera uncia) in the Annapurna Conservation Area, Nepal. Journal of Zoology 231(3), 365-370.
| Crossref | Google Scholar |
Pays O, Renaud P-C, Loisel P, Petit M, Gerard J-F, Jarman PJ (2007) Prey synchronize their vigilant behaviour with other group members. Proceedings of the Royal Society B: Biological Sciences 274(1615), 1287-1291.
| Crossref | Google Scholar |
Risenhoover KL, Bailey JA (1985) Foraging ecology of mountain sheep: implications for habitat management. The Journal of Wildlife Management 49, 797-804.
| Crossref | Google Scholar |
Schaller GB (1973) On the behaviour of blue sheep (Pseudois nayaur). Journal of Bombay Natural History Society 69(3), 523-537.
| Google Scholar |
Shrestha R, Wegge P (2006) Determining the composition of herbivore diets in the Trans-Himalayan rangelands: a comparison of field methods. Rangeland Ecology & Management 59(5), 512-518.
| Crossref | Google Scholar |
Sih A (1980) Optimal foraging: partial consumption of prey. The American Naturalist 116(2), 281-290.
| Crossref | Google Scholar |
Smith CC (2007) Independent effects of male and female density on sexual harassment, female fitness, and male competition for mates in the western mosquitofish Gambusia affinis. Behavioral Ecology and Sociobiology 61, 1349-1358.
| Crossref | Google Scholar |
Thapa K, Jackson R, Gurung L, Acharya HB, Gurung RK (2021) Applying the double observer methodology for assessing blue sheep population size in Nar Phu valley, Annapurna Conservation Area, Nepal. Wildlife Biology 2021(4), wlb.00877.
| Crossref | Google Scholar |
Underwood R (1982) Vigilance behaviour in grazing African antelopes. Behaviour 79(2–4), 81-107.
| Crossref | Google Scholar |
Wang X, Hoffmann RS (1987) Pseudois nayaur and Pseudois schaeferi. Mammalian Species [278] Issue 1-6.
| Crossref | Google Scholar |
Wegge P (1979) Aspects of the population ecology of blue sheep in Nepal. Journal of Asian Ecology 1, 10-20.
| Google Scholar |
Wegge P, Shrestha R, Flagstad Ø (2012) Snow leopard Panthera uncia predation on livestock and wild prey in a mountain valley in northern Nepal: implications for conservation management. Wildlife Biology 18(2), 131-141.
| Crossref | Google Scholar |
Xia C, Xu W, Yang W, Blank D, Qiao J, Liu W (2011) Seasonal and sexual variation in vigilance behavior of goitered gazelle (Gazella subgutturosa) in western China. Journal of Ethology 29(3), 443-451.
| Crossref | Google Scholar |
Xiao L, Hua F, Knops JMH, Zhao X, Mishra C, Lovari S, Alexander JS, Weckworth B, Lu Z (2022) Spatial separation of prey from livestock facilitates coexistence of a specialized large carnivore with human land use. Animal Conservation 25, 638-647.
| Crossref | Google Scholar |