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Activity budgets and behavioural synchrony in a wild equid living in a fission-fusion social system

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Variation in activity budgets among individuals of different age-sex classes and reproductive status may lead to decreases in behavioural synchrony (i.e., individuals performing the same behaviour at the same time in the same group) in social species. Here, we assessed the costs of behavioural synchrony in terms of time allocated to feeding behaviour among individuals of different age-sex classes and reproductive status in the kiang (Equus kiang), a poorly known wild equid that inhabits the Tibetan Plateau. Our study was conducted in Eastern Ladakh (India), during summer and fall. Our results showed that groups were highly synchronized, and that individuals in groups were particularly synchronized when feeding. Despite a slight sexual dimorphism, males and females had similar activity budgets. Males in groups, however, spent less time feeding than solitary males, and females in groups with foals spent less time feeding and more time standing than females in groups without foals. We suggest that group formation in males and the presence of foals for females incur behavioural costs by lowering their time spent feeding. Because these costs occur at a predictable time of the year, it could be profitable for adult kiangs not to form permanent groups year-round. Individuals with divergent needs might benefit from the loose social system observed in kiangs, which could be a key feature of their adaptation to a highly seasonal environment.

Affiliations: 1: Département de biologie et Centre d’études nordiques, Université Laval, Québec, QC, Canada G1V 0A6

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1. Altmann J. (1974). "Observational study of behavior: sampling methods". — Behaviour Vol 49: 227-267. [Crossref]
2. Berger J. (1983). "Induced abortion and social factors in wild horses". — Nature Vol 303: 59-61. [Crossref]
3. Berger J. (1986). Wild horses of the Great Basin: social competition and population size. — The University of Chicago Press, Chicago, IL.
4. Burnham K.P., Anderson D.R. (2002). Model selection and multimodel inference: a practical information-theoretic approach. — Springer, New York, NY.
5. Conradt L. (1998). "Could asynchrony in activity between the sexes cause intersexual social segregation in ruminants?" — Proc. Roy. Soc. Lond. B: Biol. Sci. Vol 265: 1359-1363. [Crossref]
6. Conradt L., Roper T.J. (2000). "Activity synchrony and social cohesion: a fission-fusion model". — Proc. Roy. Soc. Lond. B: Biol. Sci. Vol 267: 2213-2218. [Crossref]
7. Côté S.D., Schaefer J.A., Messier F. (1997). "Time budget and synchrony of activities in muskoxen: the influence of sex, age, and season". — Can. J. Zool. Vol 75: 1628-1635. [Crossref]
8. Denzau G., Denzau H. (1999). Wildesel. — Thorbecke, Stuttgart.
9. Duncan P. (1982). "Foal killing by stallions". — Appl. Anim. Ethol. Vol 8: 567-570. [Crossref]
10. Duncan P. (1992). Horses and grasses: the nutritional ecology of equids and their impact on the camargue. — Springer, New York, NY.
11. Fischhoff I.R., Sundaresan S.R., Cordingley J., Larkin H.M., Sellier M.J., Rubenstein D.I. (2007). "Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii". — Anim. Behav. Vol 73: 825-831. [Crossref]
12. Fox J.L., Nurbu C., Chundawat R.S. (1991). "The mountain ungulates of Ladakh, India". — Biol. Conserv. Vol 58: 167-190. [Crossref]
13. Giraldeau L.-A., Caraco T. (2000). Social foraging theory. — Princeton University Press, Princeton, NJ.
14. Hamel S., Côté S.D. (2008). "Trade-offs in activity budget in an alpine ungulate: contrasting lactating and nonlactating females". — Anim. Behav. Vol 75: 217-227. [Crossref]
15. Hauschildt V., Gerken M. (2015). "Temporal stability of social structure and behavioural synchronization in Shetland pony mares (Equus caballus) kept on pasture". — Acta Agr. Scand. A-An. Vol 65: 33-41.
16. Jarman P.J. (1974). "The social organization of antelope in relation to their ecology". — Behaviour Vol 48: 216-267. [Crossref]
17. Klingel H. (1977). "Observations on social organization and behaviour of African and Asiatic wild asses (Equus africanus and Equus hemionus)". — Z. Tierpsychol. Vol 44: 323-331. [Crossref]
18. Littell R.C., Milliken G.A., Stroup W.W., Wolfinger R.D., Schabenberger O. (2006). SAS for mixed models. — SAS Institute, Cary, NC.
19. Meldrum G.E., Ruckstuhl K.E. (2009). "Mixed-sex group formation by bighorn sheep in winter: trading costs of synchrony for benefits of group living". — Anim. Behav. Vol 77: 919-929. [Crossref]
20. Michelena P., Noël S., Gautrais J., Gerard J.-F., Deneubourg J.-L., Bon R. (2006). "Sexual dimorphism, activity budget and synchrony in groups of sheep". — Oecologia Vol 148: 170-180. [Crossref]
21. Moehlman P.D. (1998). "Feral asses (Equus africanus): intraspecific variation in social organization in arid and mesic habitats". — Appl. Anim. Behav. Sci. Vol 60: 171-195. [Crossref]
22. Moorcroft W., Trebeck G. (1841). Travels in the Provinces of Hindustan and the Panjab; in Ladakh and Kashmir; in Peshawar, Kabul, Kunduz, and Bokhara; from 1819 to 1825. — John Murray, London.
23. Neuhaus P., Ruckstuhl K.E. (2002). "The link between sexual dimorphism, activity budgets, and group cohesion: the case of the plain zebra (Equus Burchelli)". — Can. J. Zool. Vol 80: 1437-1441. [Crossref]
24. Noy-Meir I. (1973). "Desert ecosystems: environment and producers". — Annu. Rev. Ecol. Syst. Vol 4: 25-51. [Crossref]
25. Noy-Meir I. (1974). "Desert ecosystems: higher trophic levels". — Annu. Rev. Ecol. Syst. Vol 5: 195-214. [Crossref]
26. Pérez-Barbería F.J., Robertson E., Soriguer R., Aldezabal A., Mendizabal M., Pérez-Fernández E. (2007). "Why do polygynous ungulates segregate in space? Testing the activity-budget hypothesis in Soay sheep". — Ecol. Monogr. Vol 77: 631-647. [Crossref]
27. Pfister O. (2004). Birds and mammals of Ladakh. — Oxford University Press, New Delhi.
28. Putman R.J. (1986). Grazing in temperate ecosystems: large herbivores and the ecology of the New Forest. — Timber Press, Portland, OR. [Crossref]
29. Rands S.A., Cowlishaw G., Pettifor R.A., Rowcliffe J.M., Johnstone R.A. (2008). "The emergence of leaders and followers in foraging pairs when the qualities of individuals differ". — BMC Evol. Biol. Vol 8: 51. [Crossref]
30. Rawat G.S., Adhikari B.S. (2005). "Floristics and distribution of plant communities across moisture and topographic gradients in Tso Kar basin, Changthang Plateau, Eastern Ladakh". — Arct. Antarct. Alp. Res. Vol 17: 539-544. [Crossref]
31. Rook A.J., Penning P.D. (1991). "Synchronisation of eating, ruminating and idling activity by grazing sheep". — Appl. Anim. Behav. Sci. Vol 32: 157-166. [Crossref]
32. Rubenstein D.I. (1986). "Ecology and sociality in horses and zebras". — In: Ecological aspects of social evolution ( Rubenstein D.I., Wrangham R.W., eds). Princeton University Press, Princeton, NJ, p.  282-302.
33. Rubenstein D.I. (1989). "Life history and social organization in arid adapted ungulates". — J. Arid Environ. Vol 17: 145-156.
34. Rubenstein D.I. (1994). "The ecology of female social behaviour in horses, zebras, and asses". — In: Animal societies: individuals, interactions, and organization ( Jarman P., Rossiter A., eds). Kyoto University Press, Kyoto, p.  13-28.
35. Ruckstuhl K.E. (1998). "Foraging behaviour and sexual segregation in bighorn sheep". — Anim. Behav. Vol 56: 99-106. [Crossref]
36. SAS Institute (2003). SAS/stat user’s guide. — SAS Institute, Cary, NC.
37. Schaller G.B. (1998). Wildlife of the Tibetan steppe. — The University of Chicago Press, Chicago, IL.
38. St-Louis A. (2010). Écologie du kiang au Ladakh: sélection des ressources, approvisionnement et comportement social d’un équidé méconnu du Plateau Tibétain. — Ph.D. Thesis, Université Laval, Laval.
39. St-Louis A., Côté S.D. (2009). "Equus kiang (Perisodactyla: Equidae)". — Mamm. Species Vol 835: 1-11. [Crossref]
40. St-Louis A., Côté S.D. (2012). "Foraging behaviour at several temporal scales in a wild alpine equid". — Oecologia Vol 169: 167-176. [Crossref]
41. St-Louis A., Côté S.D. (2014). "Resource selection in a high-altitude rangeland equid, the kiang (Equus kiang): influence of forage abundance and quality at multiple spatial scales". — Can. J. Zool. Vol 92: 239-249. [Crossref]
42. Sundaresan S.R., Fischhoff I.R., Rubenstein D.I. (2007). "Male harassment influences female movements and associations in Grevy’s zebra (Equus grevyi)". — Behav. Ecol. Vol 18: 860-865. [Crossref]
43. van Dierendonck M.C., Sigurjónsdóttir H., Colenbrander B., Thorhallsdóttir A.G. (2004). "Differences in social behaviour between late pregnant, post-partum and barren mares in a herd of Icelandic horses". — Appl. Anim. Behav. Sci. Vol 89: 283-297. [Crossref]
44. Walker A.B.D., Parker K.L., Gillingham M.P. (2006). "Behaviour, habitat associations, and intrasexual differences of female Stone’s sheep". — Can. J. Zool. Vol 84: 1187-1201. [Crossref]
45. Ward D. (2006). "Long-term effects of herbivory on plant diversity and functional types in arid ecosystems". — In: Large herbivore ecology, ecosystem dynamics and conservation ( Danell K., Bergström R., Duncan P., Pastor J., eds). Cambridge University Press, New York, NY, p.  142-169. [Crossref]
46. Xu R. (2003). "Measuring explained variation in linear mixed effects models". — Stat. Med. Vol 22: 3527-3541. [Crossref]
47. Zar J.H. (1984). Biostatistical analysis, 2nd edn.Prentice-Hall, Englewood Cliffs.

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