Activity pattern plasticity in ungulates serves as an evolutionary adaptation to optimize fitness in inconsistent environments. Given that time is a limited and valuable resource for foraging wildlife species, provisioning and attraction may affect the activity pattern plasticity and reduce complexities of time partitioning for different activities by impala in closed environments. We assessed activity budgets of free-ranging impala social groups in a closed environment. Social group type had an influence on the activity budgets of impala except for foraging and moving activity states. Both the harem and bachelor groups spent more than 30% of their daily time foraging. Bachelor groups spent more time exhibiting vigilance tendencies than the harem groups. Season influenced the activity budgets of social groups other than vigilance and foraging activity states. Foraging time was highly correlated with vigilance, resting, and grooming. We concluded that provisioning and attraction may have reduced the influence of seasonality on the proportion of time spent on different activity states by impala social groups. There is a need to establish long-term socioecological, physiological, and reproductive consequences of provisioning and habituation on impala under closed environments. 1. Introduction Impalas (Aepyceros melampus melampus, Lichtenstein, 1812) are regarded as the most common, widely distributed, and abundant medium-sized antelope species throughout southern and east Africa [1, 2]. Classified as intermediate feeders, impalas are adapted to browsing and grazing, thus making them successful inhabitants of the savanna ecosystems [3, 4]. Favoured for game farming as well as hunting, the subspecies has been widely introduced to privately owned land and game reserves in Zimbabwe, South Africa, and Namibia [1]. For that reason, impalas are extremely important to the game ranching and conservation sector of southern Africa. In natural ecosystems, time is a valuable limited resource for all animals, and its partitioning might be influenced by sociality and as such may constrain sociality of free-ranging individuals [5]. Nakayama et al. [6] assert that the allocation of time for multiple activities has significant effects on the survival of wildlife species. Consequently, individuals adapt to environmental changes, such as food availability and temperature, by adjusting the amount of time spent in different behavioural activities [7]. The seasonality of activity budgets might be highly flexible in response to seasonal fluctuations in food supply and
References
[1]
R. East, Antelope Specialist Group Report, IUCN, Gland, Switzerland, 1998.
[2]
E. D. Lorenzen, P. Arctander, and H. R. Siegismund, “Regional genetic structuring and evolutionary history of the impala Aepyceros melampus,” Journal of Heredity, vol. 97, no. 2, pp. 119–132, 2006.
[3]
J. D. Skinner and R. H. N. Smithers, Mammals of the Southern African Sub-Region, University of Pretoria, Pretoria, South Africa, 1990.
[4]
J. Kingdom, Field Guide To African Mammals, Academic Press, London, UK, 1997.
[5]
K. A. Pollard and D. T. Blumstein, “Time allocation and the evolution of group size,” Animal Behaviour, vol. 76, no. 5, pp. 1683–1699, 2008.
[6]
Y. Nakayama, S. Matsuoka, and Y. Watanuki, “Feeding rates and energy deficits of juvenile and adult Japanese monkeys in a cool temperate area with snow coverage,” Ecological Research, vol. 14, no. 3, pp. 291–301, 1999.
[7]
M. F. Jaman and M. A. Huffman, “Enclosure environment affects the activity budgets of captive Japanese macaques (Macaca fuscata),” American Journal of Primatology, vol. 70, no. 12, pp. 1133–1144, 2008.
[8]
N. Vasey, “Activity budgets and activity rhythms in red ruffed lemurs (Varecia rubra) on the Masoala Peninsula, Madagascar: seasonality and reproductive energetics,” American Journal of Primatology, vol. 66, no. 1, pp. 23–44, 2005.
[9]
L. G. Rapaport, “Provisioning in wild golden lion tamarins (Leontopithecus rosalia): benefits to omnivorous young,” Behavioral Ecology, vol. 17, no. 2, pp. 212–221, 2006.
[10]
J. W. K. Parr and J. W. Duckworth, “Notes on diet, habituation & sociality of yellow-throated Marten (Martes flavigula),” Small Carnivore Conservation, vol. 36, pp. 27–29, 2007.
[11]
J. Knight, “Making wildlife viewable: habituation & attraction,” Society and Animals, vol. 17, no. 2, pp. 167–184, 2009.
[12]
V. Geist, “Wildlife habituation: advances in understanding and management application,” Human-Wildlife Interactions, vol. 5, pp. 9–12, 2011.
[13]
M. B. Orams, “Feeding wildlife as a tourism attraction: a review of issues and impacts,” Tourism Management, vol. 23, no. 3, pp. 281–293, 2002.
[14]
R. B. Gill, “Build an experience and they will come: managing the biology of wildlife viewing for benefi ts to people and wildlife,” in Wildlife Viewing: A Management Handbook, M. J. Manfredo, Ed., pp. 218–253, Oregon State University Press, Corvallis, Ore, USA, 2002.
[15]
O. Pays, P. Blanchard, M. Valeix et al., “Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna,” Oecologia, vol. 169, pp. 419–430, 2011.
[16]
D. Fortin, M. S. Boyce, E. H. Merrill, and J. M. Fryxell, “Foraging costs of vigilance in large mammalian herbivores,” Oikos, vol. 107, no. 1, pp. 172–180, 2004.
[17]
J. L. Koprowski and M. C. Corse, “Time budgets, activity periods, and behavior of Mexican fox squirrels,” Journal of Mammalogy, vol. 86, no. 5, pp. 947–952, 2005.
[18]
E. Donadio and S. W. Buskirk, “Flight behavior in guanacos and vicu？as in areas with and without poaching in western Argentina,” Biological Conservation, vol. 127, no. 2, pp. 139–145, 2006.
[19]
D. W. S. Challender, N. V. Thai, M. Jones, and L. May, “Time-budgets and activity patterns of captive Sunda pangolins (Manis javanica),” Zoo Biology, vol. 31, no. 2, pp. 206–218, 2012.
[20]
R. Bulton, The Makabusi Historical Background, Mukuvisi Woodlands Association, Quick Print Publishers, Harare, Zimbabwe, 1995.
[21]
M. G. Dyck and R. K. Baydack, “Vigilance behaviour of polar bears (Ursus maritimus) in the context of wildlife-viewing activities at Churchill, Manitoba, Canada,” Biological Conservation, vol. 116, no. 3, pp. 343–350, 2004.
[22]
J. Altmann, “Observational study of behavior: sampling methods,” Behaviour, vol. 49, no. 3-4, pp. 227–267, 1974.
[23]
H. F. Xu and E. D. Zhang, Wildlife Conservation and Management Principles and Techniques, East China Normal University Press, Shanghai, China, 1998.
[24]
T. Namgail, J. L. Fox, and Y. V. Bhatnagar, “Habitat shift and time budget of the Tibetan argali: the influence of livestock grazing,” Ecological Research, vol. 22, no. 1, pp. 25–31, 2007.
[25]
P. Martin and P. Bateson, Measuring Behaviour: An Introductory Guide, Cambridge University Press, 2nd edition, 1993.
[26]
K. E. Ruckstuhl, “Foraging behaviour and sexual segregation in bighorn sheep,” Animal Behaviour, vol. 56, no. 1, pp. 99–106, 1998.
[27]
B. Shorrocks and A. Cokayne, “Vigilance and group size in impala (Aepyceros melampus Lichtenstein): a study in Nairobi National Park, Kenya,” African Journal of Ecology, vol. 43, no. 2, pp. 91–96, 2005.
[28]
Z. Li, Z. Jiang, and G. Beauchamp, “Vigilance in Przewalski's gazelle: effects of sex, predation risk and group size,” Journal of Zoology, vol. 277, no. 4, pp. 302–308, 2009.
[29]
C. Xia, W. Xu, W. Yang, D. Blank, J. Qiao, and W. Liu, “Seasonal and sexual variation in vigilance behavior of goitered gazelle (Gazella subgutturosa) in western China,” Journal of Ethology, vol. 29, no. 3, pp. 443–451, 2011.
[30]
J. Burger, C. Sallna, and M. Gochfeld, “Factors affecting vigilance in springbok: importance of vegetative cover, location in herd, and herd size,” Acta Ethologica, vol. 2, no. 2, pp. 97–104, 2000.
[31]
Z. Li and Z. Jiang, “Group size effect on vigilance: evidence from Tibetan gazelle in Upper Buha River, Qinghai-Tibet Plateau,” Behavioural Processes, vol. 78, no. 1, pp. 25–28, 2008.
[32]
J. Shi, D. Li, and W. Xiao, “Influences of sex, group size, and spatial position on vigilance behavior of Przewalski's gazelles,” Acta Theriologica, vol. 56, pp. 73–79, 2010.
[33]
Y. Di Blanco and B. T. Hirsch, “Determinants of vigilance behavior in the ring-tailed coati (Nasua nasua): the importance of within-group spatial position,” Behavioral Ecology and Sociobiology, vol. 61, no. 2, pp. 173–182, 2006.
[34]
J. Burger and M. Gochfeld, “Vigilance in African mammals: differences among mothers, other females, and males,” Behaviour, vol. 131, no. 3-4, pp. 153–169, 1994.
[35]
R. J. Steidl and R. G. Anthony, “Experimental effects of human activity on breeding bald eagles,” Ecological Applications, vol. 10, no. 1, pp. 258–268, 2000.
[36]
A. Frid and L. M. Dill, “Human-caused disturbance stimuli as a form of predation risk,” Conservation Ecology, vol. 6, no. 1, p. 11, 2002.
[37]
S. L. Lima and L. M. Dill, “Behavioral decisions made under the risk of predation: a review and prospectus,” Canadian Journal of Zoology, vol. 68, no. 4, pp. 619–640, 1990.
[38]
S. L. Lima, “Stress and decision making under the risk of predation: recent developments from behavioural, reproductive and ecological perspectives,” Advances in the Study of Behavior, vol. 27, pp. 215–290, 1998.
[39]
C. M. Papouchis, F. J. Singer, and W. B. Sloan, “Responses of desert bighorn sheep to increased human recreation,” Journal of Wildlife Management, vol. 65, no. 3, pp. 573–582, 2001.
[40]
X. Lian, T. Zhang, Y. Cao, J. Su, and S. Thirgood, “Group size effects on foraging and vigilance in migratory Tibetan antelope,” Behavioural Processes, vol. 76, no. 3, pp. 192–197, 2007.
[41]
M. S. Mooring and B. L. Hart, “Costs of allogrooming in impala: distraction from vigilance,” Animal Behaviour, vol. 49, no. 5, pp. 1414–1416, 1995.
[42]
A. G. McAdam and D. L. Kramer, “Vigilance as a benefit of intermittent locomotion in small mammals,” Animal Behaviour, vol. 55, no. 1, pp. 109–117, 1998.
[43]
A. Treves, “Theory and method in studies of vigilance and aggregation,” Animal Behaviour, vol. 60, no. 6, pp. 711–722, 2000.
[44]
C. M. Bealle, “The behavioural ecology of disturbance responses,” International Journal of Comparative Psychology, vol. 20, pp. 111–120, 2007.
[45]
D. Whittaker and R. L. Knight, “Understanding wildlife responses to humans,” Wildlife Society Bulletin, vol. 26, no. 2, pp. 312–317, 1998.
[46]
J. W. Nyahongo, “Flight initiation distances of five herbivores to approaches by vehicles in the Serengeti National Park, Tanzania,” African Journal of Ecology, vol. 46, no. 2, pp. 227–229, 2008.
[47]
D. Newsome and K. Rodger, “To feed or not to feed: a contentious issue in wildlife tourism,” Australian Zoologist, vol. 34, pp. 255–270, 2008.
[48]
W. McGrew, The Cultured Chimpanzee: Reflections on Cultural Primatology, Cambridge University Press, Cambridge, UK, 2004.
[49]
J. Knight, “Feeding Mr. Monkey: cross-species food “exchange” in Japanese monkey parks,” in Animals in Person: Cultural Perspectives on Human-Animal Intimacies, J. Knight, Ed., pp. 231–253, Berg, Oxford, UK, 2005.
[50]
Q. K. Zhao and Z. Y. Deng, “Dramatic consequences of food handouts to Macaca thibethana at Mount Emei, China,” Folia Primatologica, vol. 58, pp. 24–31, 1992.
[51]
S. Periquet, L. Todd-Jones, M. Valeix et al., “Influence of immediate predation risk by lions on the vigilance of prey of different body size,” Behavioural Ecology, vol. 23, pp. 970–976, 2012.
[52]
T. Wronski, “Feeding ecology and foraging behaviour of impala Aepyceros melampus in Lake Mburo National Park, Uganda,” African Journal of Ecology, vol. 40, no. 3, pp. 205–211, 2002.
[53]
P. Blanchard and H. Fritz, “Seasonal variation in rumination parameters of free-ranging impalas Aepyceros melampus,” Wildlife Biology, vol. 14, no. 3, pp. 372–378, 2008.
[54]
P. Blanchard, R. Sabatier, and H. Fritz, “Within-group spatial position and vigilance: a role also for competition? The case of impalas (Aepyceros melampus) with a controlled food supply,” Behavioral Ecology and Sociobiology, vol. 62, no. 12, pp. 1863–1868, 2008.
[55]
K. M. Dunham, “The foraging behaviour of impala Aepyceros melampus,” South African Journal of Wildlife Research, vol. 12, pp. 36–40, 1982.
[56]
H. Robbel and G. Child, “Notes on the 1969 Rut in the Moremi,” Botswana Notes & Records, vol. 2, pp. 95–97, 1970.
[57]
R. A. Norberg, “An ecologigical theory on foraging time and energetic a choice of optional food searching method,” Journal of Animal Ecology, vol. 46, pp. 511–529, 1977.
[58]
P. E. Komers, F. Messier, and C. C. Gates, “Search or relax: the case of bachelor wood bison,” Behavioral Ecology and Sociobiology, vol. 31, no. 3, pp. 192–203, 1992.
[59]
J. T. Du Toit and C. A. Yetman, “Effects of body size on the diurnal activity budgets of African browsing ruminants,” Oecologia, vol. 143, no. 2, pp. 317–325, 2005.
[60]
C. Vanpé, N. Morellet, P. Kjellander, M. Goulard, O. Liberg, and A. J. M. Hewison, “Access to mates in a territorial ungulate is determined by the size of a male's territory, but not by its habitat quality,” Journal of Animal Ecology, vol. 78, no. 1, pp. 42–51, 2009.
[61]
A. S. Bridges, M. R. Vaughan, and S. Klenzendorf, “Seasonal variation in American black bear Ursus americanus activity patterns: quantification via remote photography,” Wildlife Biology, vol. 10, no. 4, pp. 277–284, 2004.
[62]
K. Taira and E. T. Rolls, “Receiving grooming as a reinforcer for the monkey,” Physiology and Behavior, vol. 59, no. 6, pp. 1189–1192, 1996.
[63]
C. Lazaro-Perea, M. F. De Arruda, and C. T. Snowdon, “Grooming as a reward? Social function of grooming between females in cooperatively breeding marmosets,” Animal Behaviour, vol. 67, no. 4, pp. 627–636, 2004.
[64]
B. M. Md-Zain, N. A. Sha'ari, M. Mohd-Zaki et al., “A comprehensive population survey and daily activity budget on long-tailed macaques of Universiti Kebangsaan Malaysia,” Journal of Biological Sciences, vol. 10, no. 7, pp. 608–615, 2010.
[65]
M. Y. Akinyi, J. Tung, M. Jeneby, N. B. Patel, J. Altmann, and S. C. Alberts, “Role of grooming in reducing tick load in wild baboons (Papio cynocephalus),” Animal Behaviour, vol. 85, pp. 559–568, 2013.
[66]
J. Lehmann, A. H. Korstjens, and R. I. M. Dunbar, “Group size, grooming and social cohesion in primates,” Animal Behaviour, vol. 74, no. 6, pp. 1617–1629, 2007.
[67]
G. Schino, F. Di Giuseppe, and E. Visalberghi, “The time frame of partner choice in the grooming reciprocation of Cebus apella,” Ethology, vol. 115, no. 1, pp. 70–76, 2009.
