Accessible Unlicensed Requires Authentication Published by De Gruyter November 8, 2014

Ecological drivers of group size in female Alpine chamois, Rupicapra rupicapra

Roberta Chirichella, Andrea Mustoni and Marco Apollonio
From the journal Mammalia

Abstract

In large mammalian herbivores, an increase in herd size not only reduces predation risk but also energy intake. As a consequence, the size of the groups made up by herbivores is often assumed to be the outcome of a trade-off depending on local predation risk and food availability. We studied Alpine chamois (Rupicapra rupicapra) group size in a 3660 ha wide area (central-Eastern Alps, Italy) where mouflon (Ovis gmelinii) were also present. We walked seven hiking trails (spanning a total of 33.9 km) in a high-elevation environment every 15 days from May to October in 2007–2009, and located all the groups of free-ranging ungulates within our study site. We analyzed the size variation in female groups with kids of Alpine chamois in relation to (i) forage availability (as estimated using the normalized difference vegetation index); (ii) distance from safe areas; (iii) presence of potential competitor species (i.e., mouflon); and (iv) environmental variables (elevation, slope exposure). We found that forage availability was one of the most important drivers of group size patterns. Larger groups were found in meadows with higher productivity, whereas smaller groups were located in lower quality meadows in sheltered areas, indicating how the female group size may be the outcome of both resource availability and predation risk avoidance. Interestingly, proximity of a mouflon group was able to modify the size of Alpine chamois female groups, especially if the nearest mouflon group was concurrently close and inclusive of a higher percentage of rams.


Corresponding author: Roberta Chirichella, Department of Science for Nature and Environmental Resources, University of Sassari, via Muroni 25, I-07100, Sassari, Italy, e-mail:

Acknowledgments

We are grateful to the Trento Province and the Adamello Brenta Nature Park. We are indebted to the “Associazione Cacciatori Trentini” (Trento provincial hunting association) and its game keepers who provided logistic support. Finally, we thank all the students and the field assistants for help in the field. Constructive comments from Stéphane Aulagnier, Jean-François Gerard, and an anonymous referee greatly improved this manuscript. G. Falk revised the English version of this article. The study complies with all relevant national, regional and provincial Italian laws and with the ethical standards. MA and RC were supported by the Italian Ministry of Education, University and Research (PRIN 2010–2011, 20108 TZKHC).

References

Aublet, J.F., M. Festa-Bianchet, D. Bergeron and B. Bassano. 2009. Temperature constraints on foraging behaviour of male Alpine ibex (Capra ibex) in summer. Oecologia 159: 237–247. Search in Google Scholar

Barrette, C. 1991. The size of axis deer fluid groups in Wilpattu National Park, Sri Lanka. Mammalia 55: 207–220. Search in Google Scholar

Beauchamp, G. 2003. Group-size effects on vigilance: a search for mechanisms. Behav. Process. 63: 111–121. Search in Google Scholar

Beauchamp, G. 2007. Vigilance in a selfish herd. Anim. Behav. 73: 445–451. Search in Google Scholar

Bertolino, S. 2003. Herd defensive behaviour of chamois, Rupicapra rupicapra, in response to predation on the young by a golden eagle, Aquila chrysaetos. Z. Jagdwiss. 49: 233–236. Search in Google Scholar

Bertolino, S., N.C. Montezemolo and B. Bassano. 2009. Food-niche relationships within a guild of alpine ungulates including an introduced species. J. Zool. 277: 63–69. Search in Google Scholar

Bögel, R. and G. Härer. 2002. Reactions of chamois to human disturbance in Berchtesgaden National Park. Pirineos 157: 65–80. Search in Google Scholar

Burnham, K.P., D.R. Anderson and K.P. Huyvaert. 2011. AIC model selection and multimodel inference in behavioural ecology: some background, observations, and comparisons. Behav. Ecol. Sociobiol. 65: 23–35. Search in Google Scholar

Byers, J.A. 1997. American pronghorn. Social adaptations and the ghosts of predators past. University of Chicago Press, Chicago, IL. pp. 318. Search in Google Scholar

Cederlund, G. 1989. Activity patterns in moose and roe deer in a north boreal forest. Ecography 12: 39–45. Search in Google Scholar

Chapman, N.G., K. Claydon, M. Claydon, P.G. Forde and S. Harris. 1993. Sympatric populations of muntjac (Muntiacus reevesi) and roe deer (Capreolus capreolus): a comparative analysis of their ranging behaviour, social organization and activity. J. Zool. 229: 623–640. Search in Google Scholar

Childress, M.J. and M.A. Lung. 2003. Predation risk, gender and the group size effect: does elk vigilance depend upon the behaviour of conspecifics? Anim. Behav. 66: 389–398. Search in Google Scholar

Chirichella, R., S. Ciuti and M. Apollonio. 2013. Effects of livestock and non-native mouflon on use of high-elevation pastures by Alpine chamois. Mammal. Biol. 78: 344–350. Search in Google Scholar

Ciuti, S., A. Pipia, F. Ghiandai, S. Grignolio and M. Apollonio. 2008. The key role of lamb presence in affecting flight response in Sardinian mouflon (Ovis orientalis musimon). Behav. Process. 77: 408–412. Search in Google Scholar

Clutton-Brock, T.H., S.D. Albon and F.E. Guinness. 1989. Fitness costs of gestation and lactation in wild mammals. Nature 337: 260–262. Search in Google Scholar

Cresswell, W. 1994. Flocking is an effective anti-predation strategy in redshanks, Tringa totanus. Anim. Behav. 47: 433–442. Search in Google Scholar

Crête, M. and R. Courtois. 1997. Limiting factors might obscure population regulation of moose (Cervidae: Alces alces) in unproductive boreal forests. J. Zool. 242: 765–781. Search in Google Scholar

Dehn, M.M. 1990. Vigilance for predator: detection and dilution effects. Behav. Ecol. Sociobiol. 26: 337–342. Search in Google Scholar

Ferrario, G., G. Scherini, G. Tosi and S. Toso. 1985. Distribuzione, consistenza e abitudini alimentari dell’Aquila reale (Aquila chrysaetos) nelle Alpi Centrali. In: (M. Fasola, ed.) Atti III Convegno di Ornitolodia. Salice Terme, Pavia, Italy. pp. 263–264. Search in Google Scholar

Festa-Bianchet, M. 1988a. Seasonal range selection in bighorn sheep: conflicts between forage quality, forage quantity, and predator avoidance. Oecologia 75: 580–586. Search in Google Scholar

Festa-Bianchet, M. 1988b. Nursing behaviour of bighorn sheep: correlates of ewe age, parasitism, lamb age, birthdate and sex. Anim. Behav. 36: 1445–1454. Search in Google Scholar

Festa-Bianchet, M. and S. Côté. 2008. Mountain goats: ecology, behavior, and conservation of an Alpine ungulate. Island Press, Washington, DC, USA. pp. 265. Search in Google Scholar

Festa-Bianchet, M. and J.T. Jorgenson. 1998. Selfish mothers: reproductive expenditure and resource availability in bighorn ewes. Behav. Ecol. 9: 144–150. Search in Google Scholar

Forsyth, D.M. 2000. Habitat selection and coexistence of the Alpine chamois (Rupicapra rupicapra) and Himalayan thar (Hemitragus jemlahicus) in the eastern Southern Alps, New Zealand. J. Zool. 252: 215–225. Search in Google Scholar

Forsyth, D.M. and G.J. Hickling. 1998. Increasing Himalayan thar and decreasing chamois densities in the eastern Southern Alps, New Zealand: evidence for interspecific competition. Oecologia 113: 377–382. Search in Google Scholar

Frid, A. 1997. Vigilance by female Dall’s sheep: interactions between predation risk factors. Anim. Behav. 53: 799–808. Search in Google Scholar

Fritz, H. and M. De Garine-Wichatitsky. 1996. Foraging in a social antelope: effects of group size on foraging choices and resource perception in impala. J. Anim. Ecol. 65: 736–742. Search in Google Scholar

Gaillard, J.M., J.M. Boutin, D. Delorme, G. Van Laere, P. Duncan and J.D. Lebreton. 1997. Early survival in roe deer: causes and consequences of cohort variation in two contrasted populations. Oecologia 112: 502–513. Search in Google Scholar

Gaillard, J.M., M. Festa-Bianchet and N.G. Yoccoz. 1998. Population dynamics of large herbivores: variable recruitment with constant adult survival. Trends Ecol. Evol. 13: 58–63. Search in Google Scholar

Gaillard, J.M., M. Festa-Bianchet, N.G. Yoccoz, A. Loison and C. Toïgo. 2000. Temporal variation in fitness components and population dynamics of large herbivores. Annu. Rev. Ecol. Syst. 31: 367–393. Search in Google Scholar

García-González, G. and P. Cuartas. 1996. Trophic utilization of a montane/subalpine forest by chamois (Rupicapra pyrenaica) in the central Pyrenees. For. Ecol. Manage. 88: 15–23. Search in Google Scholar

Gazzola, A., I. Bertelli, E. Avanzinelli, A. Tolosano, P. Bertotto and M. Apollonio. 2005. Predation by wolves (Canis lupus) on wild and domestic ungulates of the western Alps, Italy. J. Zool. 266: 205–213. Search in Google Scholar

Geist, V. 1971. Mountain sheep. University of Chicago Press, Chicago, IL. pp. 381. Search in Google Scholar

Gerard, J.F. and C. Richard-Hansen. 1992. Social affinities as the basis of the social organization of a Pyrenean chamois (Rupicapra pyrenaica) population in an open mountain range. Behav. Process. 28: 111–122. Search in Google Scholar

Gerard, J.F., E. Bideau, M.L. Maublanc, P. Loisel and C. Marchal. 2002. Herd size in large herbivores: encoded in the individual or emergent? Biol. Bull. 202: 275–282. Search in Google Scholar

Glutz Von Blotzheim, U.N., K.M. Bauer and E. Bezzel. 1971. Handbuch der Vögel Mitteleuropas IV. Falconiformes. Akademische Verlagsgesellschaft. Frankfurt am Main, Germany. pp. 943. Search in Google Scholar

Gonzalez, G. 1984. Ecoéthologie du mouflon et de l’isard dans le Massif du Carlit (Pyrénées-Orientales). Thèse Doctorat Université Paul Sabatier, Toulouse, France. pp. 313. Search in Google Scholar

Gonzalez, G. 1986. Compétition interspécifique chez les ongulés sauvages: l’isard et le mouflon dans le massif du Carlit (Pyrénées-Orientales). Bull. Mens. Off. Natl. Chasse 100: 35–39. Search in Google Scholar

Green, R.A. and G.D. Bear. 1990. Seasonal cycles and daily activity patterns of Rocky Mountain elk. J. Wildl. Manage. 54: 272–279. Search in Google Scholar

Griffith, B., D.C. Douglas, N.E. Walsh, D.D. Yound, T.R. McCabe, D.E. Russell, R.G. White, R.D. Cameron and K.R. Whitten. 2002. The Porcupine caribou herd. In: (D.C. Douglas, P.E. Reynolds and B. Rhode, eds.) Arctic refuge coastal plain terrestrial wildlife research summaries. US Geological Survey, Biological Resources Division, Biological Science Report USGS/BRD/BSR-2002-0001. pp. 8–37. Search in Google Scholar

Grignolio, S., I. Rossi, E. Bertolotto, B. Bassano and M. Apollonio. 2007. Influence of the kid on space use and habitat selection of female Alpine ibex. J. Wildl. Manage. 71: 713–719. Search in Google Scholar

Hadjisterkotis, E. 1990. The breeding ecology of Cyprus mouflon (Ovis gmellin ophion). Officer of the Game and Fauna Service. Ministry of Interior, Nicosia, Cyprus. pp. 18–19. Search in Google Scholar

Haller, H. 1996. Der Steinadler in Graubiinden. Langfristige Untersuchungen zur Populationsökologie von Aquila chrysaetos im Zentrum der Alpen. Orn. Beob. Beiheft. 9: 1–167. Search in Google Scholar

Haller, H. and P. Sackl. 1997. Golden eagle Aquila chrysaetos. In: (W.J.M. Hagemeijer and M.J. Blair, eds.) The EBCC Atlas of European breeding birds, their distribution and abundance. Poyser, London, UK. pp. 170–171. Search in Google Scholar

Hamel, S. and S.D. Côté. 2009. Maternal defensive behaviour of mountain goats against predation by golden eagles. West. N. Am. Nat. 69: 115–118. Search in Google Scholar

Hartnett, D.C., A.A. Steuter and K.R. Hickman. 1997. Comparative ecology of native and introduced ungulates. In: (F.L. Knopf and F.B. Samson, eds.) Ecology and conservation of great plains vertebrates. Springer, Berlin-Heidelberg-New York, Germany-NY. pp. 72–101. Search in Google Scholar

Herrero, J., I. Garin, R. Garcia-González and A. Aldezabal. 2002. Grouping patterns in a forest dwelling population of Pyrenean chamois. Pirineos 157: 89–101. Search in Google Scholar

Hunter, L.T.B. and J.D. Skinner. 1998. Vigilance behaviour in Africa ungulates: the role of predation pressure. Behaviour 135: 195–211. Search in Google Scholar

Isvaran, K. 2007. Intraspecific variation in group size in the blackbuck antelope: the roles of habitat structure and forage at different spatial scales. Oecologia 154: 435–444. Search in Google Scholar

Jarman, P.J. 1974. The social organization of antelope in relation to their ecology. Behaviour 48: 215–267. Search in Google Scholar

Kie, J.G. 1999. Optimal foraging and risk of predation: effects on behaviour and social structure in ungulates. J. Mammal. 80: 1114–1129. Search in Google Scholar

Knaus, W. and W. Schröder. 1983. Das Gamswind. Paul Parey, Berlin, Germany. pp. 232. Search in Google Scholar

Krämer, A. 1969. Soziale Organisation und Sozialverhalten einer Gemes-population der Alpen. Z. Tierpsychol. 26: 889–964. Search in Google Scholar

Latham, J., B.W. Staines and M.L. Gorman. 1999. Comparative feeding ecology of red (Cervus elaphus) and roe deer (Capreolus capreolus) in Scottish plantation forests. J. Zool. 247: 409–418. Search in Google Scholar

LeBlanc, M., M. Festa-Bianchet and J.T. Jorgenson. 2001. Sexual size dimorphism in bighorn sheep (Ovis canadensis): effects of population density. Can. J. Zool. 79: 1661–1670. Search in Google Scholar

Lingle, S., S.M. Pellis and W.F. Wilson. 2005. Interspecific variation in antipredator behaviour leads to differential vulnerability of mule deer and white-tailed deer fawns early in life. J. Anim. Ecol. 74: 1140–1149. Search in Google Scholar

Machlis, L., P.W.D. Dodd and J.C. Fentress. 1985. The pooling fallacy: problems arising when individuals contribute more than one observation to the data set. Z. Tierpsychol. 68: 201–214. Search in Google Scholar

Magee, L. 1990. R2 measures based on Wald and likelihood ratio joint significance tests. Am. Stat. 44: 250–253. Search in Google Scholar

Manor, R. and D. Saltz. 2003. Impact of human nuisance disturbance on vigilance and group size of a social ungulate. Ecol. Appl. 13: 1830–1834. Search in Google Scholar

McNamara, J.M. and A.I. Houston. 1987. Starvation and predation as factors limiting population-size. Ecology 68: 1515–1519. Search in Google Scholar

Mishra, C., S.E. Wieren, P. Ketner, I.M.A. Heitkönig and H.H.T. Prins. 2004. Competition between domestic livestock and wild bharal Pseudois nayaur in the Indian Trans Himalaya. J. Appl. Ecol. 41: 344–454. Search in Google Scholar

Molvar, E.M. and R.T. Bowyer. 1994. Costs and benefits of group living in a recently social ungulate: the Alaskan moose. J. Mammal. 75: 621–630. Search in Google Scholar

Nebel, D., A. Duquesne and J. Geoffroy. 1996. L’Aigle royal Aquila chrysaetos dans la Réserve Domaniale du Mont Vallier (Ariège) 1987-1995. Alauda 64: 179–186. Search in Google Scholar

Pedrini, P., M. Caldonazzi and S. Zanghellini. 2005. Atlante degli Uccelli nidificanti e svernanti in provincia di Trento. Museo Tridentino di Scienze Naturali, Trento. Studi Trentini di Scienze Naturali. Acta Biol. 80: 154–157. Search in Google Scholar

Pépin, D. and J.F. Gerard. 2008. Group dynamics and local population density dependence of group size in the Pyrenean chamois, Rupicapra pyrenaica. Anim. Behav. 75: 361–369. Search in Google Scholar

Pérez-Barbería, F.J., L. Robles and C. Nores. 1996. Horn growth pattern in Cantabrian chamois Rupicapra pyrenaica parva: Influence of sex, location and phenology. Acta Theriol. 41: 83–92. Search in Google Scholar

Pettorelli, N., J.O. Vik, A. Mysterud, J.M. Gaillard, C.J. Tucker and N.C. Stenseth. 2005. Using the satellite-derived NDVI to assess ecological responses to environmental change. Trends Ecol. Evol. 20: 503–510. Search in Google Scholar

Pipia, A., S. Ciuti, S. Grignolio, S. Luchetti, R. Madau and M. Apollonio. 2009. Effect of predation risk on grouping pattern and whistling behaviour in a wild mouflon Ovis aries population. Acta Theriol. 54: 77–86. Search in Google Scholar

Pulliam, H.R. 1973. On the advantages of floking. J. Theor. Biol. 38: 419–422. Search in Google Scholar

R Development Core Team. 2011. R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. Vienna, Austria. ISBN 3-900051-07-0. http://www.R-project.org/. Search in Google Scholar

Raithel, J.D., M.J. Kauffman and D.H. Pletscher. 2007. Impact of spatial and temporal variation in calf survival on the growth of elk populations. J. Wildl. Manage. 71: 795–803. Search in Google Scholar

Reed, B.C., J.F. Brown, D. VanderZee, T.R. Loveland, J.W. Merchant and D.O. Ohlen 1994. Measuring phenological variability from satellite imagery. J. Veg. Sci. 5: 703–714. Search in Google Scholar

Richard-Hansen, C. and R. Campan. 1992. Social environment of isard kids, Rupicapra pyrenaica p., during their ontogeny. Z. Säugetierk. 57: 351–363. Search in Google Scholar

Ruckstuhl, K.E. and P. Ingold. 1999. Aspects of mother-kid behaviour in Alpine chamois, Rupicapra rupicapra. Z. Säugetierk. 64: 76–84. Search in Google Scholar

Schröder, W. 1971. Untersuchungen zur Ökologie des Gamswildes (Rupicapra rupicapra) in einen Vorkommen del Alpen. Z. Jagdwiss. 17: 114–232. Search in Google Scholar

Shankar Raman, T.R. 1997. Factors influencing seasonal and monthly changes in the group size of chital or axis deer in southern India. J. Biosci. 22: 203–218. Search in Google Scholar

Sokal R.R. and F.J. Rohlf. 1995. Biometry: the principles and practice of statistics in biological research, 3rd ed. Freeman, New York, NY. pp. 887. Search in Google Scholar

Symonds, M.R.E. and A. Mousalli. 2011. A brief guide to model selection, multimodel inference and model averaging in behavioural ecology using Akaike’s information criterion. Behav. Ecol. Sociobiol. 65: 13–21. Search in Google Scholar

Vázquez, D.P. 2002. Multiple effects of introduced mammalian herbivores in a temperated forest. Biol. Invasions 4: 175–191. Search in Google Scholar

Zweifel-Schielly, B., M. Kreurer, K.C. Ewald and W. Suter. 2008. Habitat selection by an Alpine ungulate: the significance of forage characteristics varies with scale and season. Ecography 32: 103–113. Search in Google Scholar

Received: 2014-1-27
Accepted: 2014-10-7
Published Online: 2014-11-8
Published in Print: 2015-11-1

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