Social arenas in the open habitat: the social role of waterholes for saiga antelope



mineral lick, open landscape, social behaviour, socialization, ungulate, waterhole


In wild animals, specific locations may function as ‘social arenas’ playing a significant role in various aspects of intraspecific interactions.  The emergence of such social arenas is assumed to be particularly important under conditions of low visibility and density of individuals typical for forest dwelling species.  In the forest, open areas with a high probability of meeting conspecifics such as waterholes and mineral licks can favor socialization.  The paucity of studies on the open-habitat species limits our understanding of whether forest habitat is an essential condition for the emergence of social arenas.  Here we address this issue by investigating whether resource hotspots function as social arenas in a species adapted to open habitat.  The social behaviour in groups of saiga antelopes (Saiga tatarica tatarica) was recorded at large permanent waterholes (serving also as mineral licks) and away from them in the steppe environment.  The frequency and variety of social interactions between the group mates were compared with a special effort made to minimize the potential impact of other factors besides the location.  Also the focal individuals, interacting or not interacting with conspecifics at the waterholes, were traced continuously to explore the duration of their visit at the waterhole area. Finally, we recorded the number of individuals in the groups entering and leaving the waterhole.  Active social interactions between group members occurred more often and were more varied near the waterholes as compared to the areas away from them.  The participation in social interactions influenced the duration of individual waterhole visits.  Males that interacted with others shortly after arrival stayed significantly longer compared to those who did not.  Females whose calves engaged in social interactions with age-mates spent more time at the waterholes than females whose calves did not.  The comparison of mean group sizes showed that saigas left waterhole areas in larger groups than they entered them.  Our findings, together with previous studies, suggest that resource hotspots can function as social arenas in mammals regardless of the habitat type (open or closed).  The emergence of specific locations having particular social significance may be especially important for highly dispersed species with large home ranges.  The results of the study emphasize the significance of large permanent waterholes and mineral licks used by many individuals year after year for the social behaviour of saiga antelopes.

Author Biographies

Andrey Giljov, Saint Petersburg State University

PhD, Associate Professor at the Vertebrate Zoology Department

Karina Karenina, Saint Petersburg State University

PhD, Senior Researcher at the Vertebrate Zoology Department


Bekenov, A.B., Grachev, I.A., Milnerâ€Gulland, E.J. 1998. The ecology and management of the saiga antelope in Kazakhstan. Mammal Review 28:1-52.

Blank, D., Li, Y. 2022. Antelope adaptations to counteract overheating and water deficit in arid environments. Journal of Arid Land, 14:1069-1085.

Brightsmith, D.J., Villalobos, E.M. 2011. Parrot behavior at a Peruvian clay lick. The Wilson Journal of Ornithology 123:595-602.

Bro-Jørgensen, J. 2003. The significance of hotspots to lekking topi antelopes (Damaliscus lunatus). Behavioral Ecology and Sociobiology 53:324-331.

Clayton, L., MacDonald, D.W. 1999. Social organization of the babirusa (Babyrousa babyrussa) and their use of salt licks in Sulawesi, Indonesia. Journal of Mammalogy 80:1147-1157.

Couturier, S., Barrette, C. 1988. The behavior of moose at natural mineral springs in Quebec. Canadian Journal of Zoology 66:522-528.

Egbert, A.L., Stokes, A.W., Egbert, A.L. 1976. The social behaviour of brown bears on an Alaskan salmon stream. Bears: Their Biology and Management 41-56.

Fishlock, V., Lee, P.C. 2013. Forest elephants: fission–fusion and social arenas. Animal Behaviour 85:357-363.

Fishlock, V., Lee, P.C., Breuer, T. 2008. Quantifying forest elephant social structure in Central African bai environments. Pachyderm 44:19-28.

Fraser, D., Hristienko, H. 1981. Activity of moose and white-tailed deer at mineral springs. Canadian Journal of Zoology 59:1991-2000.

Gallucci, M. 2019. GAMLj: General analyses for linear models. [jamovi module]. Retrieved from

Gilev, A., Karenina, K. 2015. The significance of artesian wells for saigas within the Stepnoi Sanctuary, Astrakhan region. Saiga News 20:15–17.

Giljov, A., Malashichev, Y., Karenina, K. 2019. What do wild saiga antelopes tell us about the relative roles of the two brain hemispheres in social interactions? Animal Cognition 22:635–643.

Hardenberg, A., Bassano, B., Peracino, A., Lovari, S. 2000. Male Alpine chamois occupy territories at hotspots before the mating season. Ethology 106:617-630.

Hii, N. 2017. Asian elephants’ social structure and mineral lick usage in a Malaysian rainforest using camera traps. M.Sc. Thesis. The University of Nottingham Malaysia Campus, Selangor, Malaysia.

Karimova, T.Y., Lushchekina, A.A. 2018. Features of the spatial distribution and ethological structure of saiga population within the “Stepnoy†sanctuary (Astrakhan oblast). Ecosystems: Ecology and Dynamics 2:73-91.

Karimova, T.Y., Lushchekina, A.A., Neronov, V.M., Pyurvenova, N.Y., Arylov, Y.N. 2020. Biological features of the Northwest Pre-Caspian saiga population at different sizes. Arid Ecosystems 10: 298-304.

King, A., Behie, A.M., Hon, N., Rawson, B.M. 2016. Patterns of salt lick use by mammals and birds in northeastern Cambodia. Cambodian Journal of Natural History, 1:40-50.

Klaus, G., Klaus-Hugi, C., Schmid, B. 1998. Geophagy by large mammals at natural salt licks in the rainforest of Dzanga National Park, Central African Republic. Journal of Tropical Ecology 14:829-839.

Klausâ€Hügi, C., Klaus, G., and Schmid, B. 2000. Movement patterns and home range of the bongo (Tragelaphus eurycerus) in the rain forest of the Dzanga National Park, Central African Republic. African Journal of Ecology 38:53-61.

Melletti, M., Penteriani, V., Boitani, L. 2007. Habitat preferences of the secretive forest buffalo (Syncerus caffer nanus) in Central Africa. Journal of Zoology 271:178-186.

Merte, C.E., Goodwin, T.E., Schulte, B.A. 2010. Male and female developmental differences in chemosensory investigations by African elephants (Loxodonta africana) approaching waterholes. Behavioral Ecology and Sociobiology 64:401-408.

Milner-Gulland, E.J., Bukreeva, O.M., Coulson, T., Lushchekina, A.A., Kholodova, M.V., Bekenov, A.B., Grachev, I.A. 2003. Reproductive collapse in saiga antelope harems. Nature 422:135-135.

Neronov, V.M., Lushchekina, A.A., Karimova, T.Y., Arylova, N.Y. 2012. Population dynamics of a key steppe species in a changing world: the critically endangered saiga antelope. Pp. 335-356 in Eurasian steppes. Ecological problems and livelihoods in a changing world. (M. J. Werger, M. A. Van Staalduinen, eds.) Springer, Dordrecht, Netherlands.

Ruggiero, R.G., Fay, J.M. 1994. Utilization of termitarium soils by elephants and its ecological implications. African Journal of Ecology 32:222-232.

Sokolov, V.E., Zhirnov, L.V. 1998. The Saiga: Phylogeny, Systematics, Ecology, Conservation and Use. Russian Academy of Sciences, Moscow, Russia.

Turkalo, A., Fay, J.M. 1995. Studying forest elephants by direct observation: preliminary results from the Dzanga clearing, Central African Republic. Pachyderm 20:45-54.

Turkalo, A.K., Wrege, P.H., Wittemyer, G. 2013. Long-term monitoring of Dzanga Bai forest elephants: forest clearing use patterns. PloS One 8:e85154.

van Overveld, T., et al. 2020. Seasonal grouping dynamics in a territorial vulture: ecological drivers and social consequences. Behavioral Ecology and Sociobiology 74:28.

van Strien, N.J. 1985. The Sumatran Rhinoceros - Dicerorhinus sumatrensis Fischer, 1814 - in the Gunung Leuser National Park, Sumatra, Indonesia; its Distribution, Ecology and Conservation. Ph.D. thesis. Wageningen, the Netherlands.

Weir, J.S. 1969. Chemical properties and occurrence on Kalahari sand of salt licks created by elephants. Journal of Zoology 158:293-310.


Additional Files