Shifter and resident red deer: intrapopulation and intersexual behavioural diversities in a predator-free area
A. Bocci A , I. Angelini A , P. Brambilla A , A. Monaco A B and S. Lovari A C
+ Author Affiliations
- Author Affiliations
A University of Siena, Department of Life Sciences, Research Unit of Behavioural Ecology, Ethology and Wildlife Management, via P. A. Mattioli, 4-53100 Siena, Italy.
B Present address: Latium Regional Park Agency, Latium Region, Via del Pescaccio 96, 00166 Rome, Italy.
C Corresponding author. Email: lovari@unisi.it
Wildlife Research 39(7) 573-582 https://doi.org/10.1071/WR12037
Submitted: 24 February 2012 Accepted: 22 July 2012 Published: 7 September 2012
Abstract
Context: Population management of meso-large game species; knowledge of seasonal movements and sexual segregation.
Aims: We predicted (1) a similar mixed spatial tactic in the two sexes (shifters and residents); (2) that reproductive success could be a key factor eliciting a mixed spatial tactic in hinds; and (3) that hunting activities may affect the rutting season, influencing rutting site location.
Methods: We studied the spatial behaviour of 41 red deer on the Italian north-eastern Alps, from December 2003 to October 2007, by VHF radio-tracking.
Key results: For both sexes, two main spatial tactics were described, including residents, remaining in the same area all year long, and shifters, using non-overlapping seasonal home ranges. Most deer showed consistent spatial behaviour. In all, 87% of stags and 49% of hinds were shifters. Sexual segregation was greater in winter than in summer. All hinds wintered in the same valley, with residents in the qualitatively poorer, but protected, area. Resident hinds showed a variable yearly reproductive success, whereas usually all shifters had a calf at heel. Rutting areas of hinds overlapped with their winter or summer ranges, whereas stags occupied up to three different areas yearly. Deer distribution determined a different harvesting pressure on the two sexes.
Conclusions: Although shifters and residents were found in both sexes, differences in spatial behaviour of stags and hinds were consistent for seasonal occupancy, habitat selection and number of seasonal home ranges. For hinds, a mixed spatial tactic, seasonality and hunting activities could be key factors of reproductive success.
Implications: Hunting could affect the proportions of shifter and resident hinds at different levels, with the latter benefiting from the protection accorded. This unequally distributed selective harvesting may alter the local social structure and population dynamics in the long term. Shifter behaviour, seasonal movements and sexual segregation may bias population estimates, if counts are conducted when individuals make seasonal movements. Furthermore, the size of a hunting unit should be function of the extent of local movements of deer.
Additional keywords: behaviour, habitat use, radio-telemetry, spatial ecology, wildlife management.
References
Aebischer, N. J., Robertson, P. A., and Kenward, R. E. (1993). Compositional analysis of habitat use from animal radio-tracking data. Ecology 74, 1313–1325.| Compositional analysis of habitat use from animal radio-tracking data.Crossref | GoogleScholarGoogle Scholar |
Albon, S. D., and Langvatn, R. (1992). Plant phaenology and the benefits of migration in a temperate ungulate. Oikos 65, 502–513.
| Plant phaenology and the benefits of migration in a temperate ungulate.Crossref | GoogleScholarGoogle Scholar |
Andersen, R. (1991). Habitat deterioration and the migratory behaviour of moose (Alces alces L.) in Norway. Journal of Applied Ecology 28, 102–108.
| Habitat deterioration and the migratory behaviour of moose (Alces alces L.) in Norway.Crossref | GoogleScholarGoogle Scholar |
Baker, R. R. (1978). ‘The Evolutionary Ecology of Animal Migration.’ (Holmes and Meier Publishers: New York.)
Berger, J. (2004). The last mile: how to sustain long-distance migration in mammals. Conservation Biology 18, 320–331.
| The last mile: how to sustain long-distance migration in mammals.Crossref | GoogleScholarGoogle Scholar |
Bocci, A., Monaco, A., Brambilla, P., Angelini, I., and Lovari, S. (2010a). Alternative strategies of space use of female red deer in a mountainous habitat. Annales Zoologici Fennici 47, 57–66.
Bocci, A., Canavese, G., and Lovari, S. (2010b). Even mortality patterns of the two sexes in a polygynous, near-monomorphic species: is there a flaw? Journal of Zoology 280, 379–386.
| Even mortality patterns of the two sexes in a polygynous, near-monomorphic species: is there a flaw?Crossref | GoogleScholarGoogle Scholar |
Bocci, A., Telford, M., and Laiolo, P. (). Determinants of the acoustic behaviour of red deer during breeding in a wild alpine population, and implication for species survey. Ethology Ecology and Evolution , .
Boitani, L., Lovari, S., and Vigna Taglianti, A. (2003). ‘Fauna d’Italia. Mammalia III. Carnivora–Arctiodactyla.’ (Calderini: Bologna, Italy.)
Bonenfant, C., Loe, L. E., Mysterud, A., Langvatn, R., Stenseth, N. C., Gaillard, J.-M., and Klein, F. (2004). Multiple causes of sexual segregation in red deer: enlightenments from breeding phenology at high and low latitude. Proceedings of the Royal Society of London 271, 883–892.
| Multiple causes of sexual segregation in red deer: enlightenments from breeding phenology at high and low latitude.Crossref | GoogleScholarGoogle Scholar |
Brugnoli, A., Cetto, E., Partel, P., and Zugliani, G. (1998). Censimento di cervo (Cervus elaphus hippelaphus Erxleben, 1977) al bramito nel Parco Naturale Paneveggio-Pale di San Martino. Technical report. Ente Parco Paneveggio-Pale di San Martino, Trento, Italy.
Byers, J. A., Wiseman, P. A., Jones, L., and Roffe, T. J. (2005). A large cost of female mate sampling in Pronghorn. American Naturalist 166, 661–668.
| A large cost of female mate sampling in Pronghorn.Crossref | GoogleScholarGoogle Scholar |
Carranza, J. (1995). Female attraction by males versus sites in territorial rutting red deer. Animal Behaviour 50, 445–453.
| Female attraction by males versus sites in territorial rutting red deer.Crossref | GoogleScholarGoogle Scholar |
Clutton-Brock T. H. 1989
Clutton-Brock, T. H., Guinness, F. E., and Albon, S. D. (1982). ‘Red Deer: Behaviour and Ecology of Two Sexes.’ (The University of Chicago Press: Chicago, IL.)
Clutton-Brock, T. H., Iason, G. R., and Guinness, F. E. (1987). Sexual segregation and density related changes in habitat use in male and female red deer (Cervus elaphus). Journal of Zoology 211, 275–289.
| Sexual segregation and density related changes in habitat use in male and female red deer (Cervus elaphus).Crossref | GoogleScholarGoogle Scholar |
Clutton-Brock, T. H., Coulson, T. N., Milner-Gulland, E. J., Thompson, D., and Armstrong, H. M. (2002). Sex differences in emigration and mortality affect optimal management of deer populations. Nature 415, 633–637.
| Sex differences in emigration and mortality affect optimal management of deer populations.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD38XhsVCjsbo%3D&md5=5e4a6394de636a7c6d492a4e5f30e319CAS |
Coltman, D. W., O’Donoghue, P., Jorgenson, J. T., Hogg, J. T., Strobeck, C., and Festa-Bianchet, M. (2003). Undesirable evolutionary consequences of trophy hunting. Nature 426, 655–658.
| Undesirable evolutionary consequences of trophy hunting.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BD3sXps1Cks74%3D&md5=f338de03ab210907ca85ee8be1548521CAS |
Conradt, L., Clutton-Brock, T. H., and Guinness, F. E. (2000). Sex differences in weather sensitivity can cause habitat segregation: red deer as an example. Animal Behaviour 59, 1049–1060.
| Sex differences in weather sensitivity can cause habitat segregation: red deer as an example.Crossref | GoogleScholarGoogle Scholar |
Cook, J. G., Johnson, B. K., Cook, R. C., Riggs, R. A., Delcurto, T., Bryant, L. D., and Irwin, L. L. (2004). Effects of summer-autumn nutrition and parturition date on reproduction and survival of elk. Wildlife Monographs 155, 1–61.
Del Giudice, G. D., Riggs, M. R., Joly, P., and Pan, W. (2002). Winter severity, survival and cause-specific mortality of female white-tailed deer in north central Minnesota. The Journal of Wildlife Management 66, 698–717.
| Winter severity, survival and cause-specific mortality of female white-tailed deer in north central Minnesota.Crossref | GoogleScholarGoogle Scholar |
Douglas, M. J. W. (1971). Behaviour responses of red deer and chamois to cessation of hunting. New Zealand Journal of Science 14, 507–518.
Ford, R. G. (1983). Home range in a patchy environment: optimal foraging predictions. American Zoologist 23, 325–326.
Forsyth, D. M. (1999). Long-term harvesting and male migration in a New Zealand population of Himalayan tahr Hemitragus jemlahicus. Journal of Applied Ecology 36, 351–362.
| Long-term harvesting and male migration in a New Zealand population of Himalayan tahr Hemitragus jemlahicus.Crossref | GoogleScholarGoogle Scholar |
Forsyth, D. M., Allen, R. B., Marburg, A. E., MacKenzie, D. I., and Douglas, M. J. W. (2010). Population dynamics and resource use of red deer following release from harvesting in New Zealand. New Zealand Journal of Ecology 34, 277–287.
Fryxell, J. M., and Sinclair, A. R. E. (1988). Causes and consequences of migration by large herbivores. Trends in Ecology & Evolution 3, 237–241.
| Causes and consequences of migration by large herbivores.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BC3M7gvFehtQ%3D%3D&md5=47e3b456769caa8e18fc6a9961be0f73CAS |
Gafta, D., and Pedrotti, F. (1998). Fitoclima del Trentino – Alto Adige [phytoclimate of Trentino-Alto Adige]. Studi Trentini di Scienze Naturali. Acta Biologica 73, 55–111.
Gaillard, J.-M., Festa-Bianchet, M., Yoccoz, Y., Loison, A., and Toigo, C. (2000). Temporal variation in fitness components and population dynamics of large herbivores. Annual Review of Ecology Evolution and Systematics 31, 367–393.
| Temporal variation in fitness components and population dynamics of large herbivores.Crossref | GoogleScholarGoogle Scholar |
Geist, V. (1982). Adaptive behavioral strategies. In ‘Elk of North America’. (Eds J. W. Thomas and D. E. Toweill.) pp. 219–277. (Wildlife Management Institute Book: Stackpole, Harrisburg, PA.)
Geist, V. (1999). Adaptive strategies in American mountain sheep. In ‘Mountain Sheep of North America’. (Eds R. Valdez and P. R. Krausman.) pp. 193–208. (University of Arizona Press: Tucson, AZ.)
Geist, V. (2002). Adaptive behavioral strategies. In ‘North American Elk’. (Eds D. E. Toweill and J. W. Thomas.) pp. 389–433. (Smithsonian Institution Press: Washington, DC.)
Georgii, B. (1980). Home range patterns of female red deer (Cervus elaphus L.) in the Alps. Oecologia 47, 278–285.
| Home range patterns of female red deer (Cervus elaphus L.) in the Alps.Crossref | GoogleScholarGoogle Scholar |
Georgii, B., and Schröder, W. (1983). Home range and activity patterns of male red deer (Cervus elaphus L.) in the Alps. Oecologia 58, 238–248.
| Home range and activity patterns of male red deer (Cervus elaphus L.) in the Alps.Crossref | GoogleScholarGoogle Scholar |
Guinness, F. E., Clutton-Brock, T. H., and Albon, S. D. (1978). Factors affecting calf mortality in red deer (Cervus elaphus). Journal of Animal Ecology 47, 817–832.
| Factors affecting calf mortality in red deer (Cervus elaphus).Crossref | GoogleScholarGoogle Scholar |
Hald, A. (1967). ‘Statistical Theory with Engineering Applications.’ (Wiley: New York.)
Hooge, P. N., and Eichenlaub, B. (1997). ‘Animal Movement Extension to Arcview. Ver. 1.1.’ (Alaska Biological Science Center, US Geological Survey: Anchorage, AK.)
Iason, G. R., Duck, C. D., and Clutton-Brock, T. H. (1986). Grazing and reproductive success of red deer: the effect of local enrichment by gull colonies. Journal of Animal Ecology 56, 989–999.
Irwin, L. L. (2002). Migration. In ‘North American Elk’. (Eds D. E. Toweill and J. W. Thomas.) pp. 493–513. (Smithsonian Institution Press: Washington, DC.)
Jarnemo, A. (2008). Seasonal migration of male red deer (Cervus elaphus) in southern Sweden and consequences for management. European Journal of Wildlife Research 54, 327–333.
| Seasonal migration of male red deer (Cervus elaphus) in southern Sweden and consequences for management.Crossref | GoogleScholarGoogle Scholar |
Jenness, J. (2004). ‘Weighted Mean of Point (weightmean.avx) Extension for ArcView 3.x, v. 1.2c.’ (Jenness Enterprises: Flagstaff.)
Johnson, D. H. (1980). The Comparison of usage and availability measurements for evaluating resource preference. Ecology 61, 65–71.
| The Comparison of usage and availability measurements for evaluating resource preference.Crossref | GoogleScholarGoogle Scholar |
Knight, R. R. (1970). The Sun River elk herd. Wildlife Monographs 23, 3–66.
Kruuk, L. E. B., Clutton-Brock, T. H., Rose, K. E., and Guinness, F. E. (1999). Early determinants of lifetime reproductive success differ between the sexes in red deer. Proceedings. Biological Sciences 266, 1655–1661.
| Early determinants of lifetime reproductive success differ between the sexes in red deer.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaK1MvjtVyhtg%3D%3D&md5=9f022d4f9b765ad19f512c1c886b78b7CAS |
Loison, A., Solberg, E.-J., Yoccoz, E. J., and Langvatn, R. (2004). Sex differences in the interplay of cohort and mother quality on body mass of red deer calves. Ecology 85, 1992–2002.
| Sex differences in the interplay of cohort and mother quality on body mass of red deer calves.Crossref | GoogleScholarGoogle Scholar |
Lovari, S., Sacconi, F., and Trivellini, G. (2006). Do alternative strategies of space use occur in male Alpine chamois? Ethology Ecology and Evolution 18, 221–231.
| Do alternative strategies of space use occur in male Alpine chamois?Crossref | GoogleScholarGoogle Scholar |
Lowe, V. P. W. (1967). Teeth as indicators of age with special reference to red deer (Cervus elaphus) of known age from Rhum. Journal of Zoology 152, 137–153.
| Teeth as indicators of age with special reference to red deer (Cervus elaphus) of known age from Rhum.Crossref | GoogleScholarGoogle Scholar |
Luccarini, S., Mauri, L., Ciuti, S., Lamberti, P., and Apollonio, M. (2006). Red deer (Cervus elaphus) spatial use in the Italian Alps: home range patterns, seasonal migrations, and effects of snow and winter feeding. Ethology Ecology and Evolution 18, 127–145.
| Red deer (Cervus elaphus) spatial use in the Italian Alps: home range patterns, seasonal migrations, and effects of snow and winter feeding.Crossref | GoogleScholarGoogle Scholar |
Magurran, A. E. (1988). ‘Ecological Diversity and its Measurement.’ (Croom Helm: London.)
Main, M. B. (2008). Reconciling competing ecological explanations for sexual segregation in ungulates Ecology 89, 693–704.
Martinez, J. G., Carranza, J., Fernàndez-Garcìa, J. L., and Sànchez-Prieto, C. B. (2002). Genetic variation of red deer populations under hunting exploitation in southwestern Spain. The Journal of Wildlife Management 66, 1273–1282.
McCullough, D. R. (1996). Spatially structured populations and harvest theory. The Journal of Wildlife Management 60, 1–9.
| Spatially structured populations and harvest theory.Crossref | GoogleScholarGoogle Scholar |
McNab, B. K. (1963). Bioenergetics and determination of home range size. American Naturalist 97, 133–140.
| Bioenergetics and determination of home range size.Crossref | GoogleScholarGoogle Scholar |
Milner, J. M., Nilsen, E. B., and Andreassen, H. P. (2007). Demographic side effects of selective hunting in ungulates and carnivores. Conservation Biology 21, 36–47.
| Demographic side effects of selective hunting in ungulates and carnivores.Crossref | GoogleScholarGoogle Scholar |
Minta, S. C. (1993). Sexual differences in spatio-temporal interaction among badgers. Oecologia 96, 402–409.
| Sexual differences in spatio-temporal interaction among badgers.Crossref | GoogleScholarGoogle Scholar |
Mysterud, A., Langvatn, R., Yoccoz, N. G., and Stenseth, N. C. (2001). Plant phenology, migration and geographical variation in body weight of a large herbivore: the effect of a variable topography. Journal of Animal Ecology 70, 915–923.
| Plant phenology, migration and geographical variation in body weight of a large herbivore: the effect of a variable topography.Crossref | GoogleScholarGoogle Scholar |
Nicholson, M. C., Bowyer, R. T., and Kie, J. G. (1997). Habitat selection and survival of mule deer: tradeoffs associated with migration. Journal of Mammalogy 78, 483–504.
| Habitat selection and survival of mule deer: tradeoffs associated with migration.Crossref | GoogleScholarGoogle Scholar |
Parker, K. L., Robbins, C. T., and Hanley, T. A. (1984). Energy expenditures for locomotion by mule deer and elk. The Journal of Wildlife Management 48, 474–488.
| Energy expenditures for locomotion by mule deer and elk.Crossref | GoogleScholarGoogle Scholar |
Parker, K. L., Gillingham, M. P., Hanley, T. A., and Robbins, C. T. (1996). Foraging efficiency: energy expenditure versus energy gain in free-ranging black-tailed deer. Canadian Journal of Zoology 74, 442–450.
| Foraging efficiency: energy expenditure versus energy gain in free-ranging black-tailed deer.Crossref | GoogleScholarGoogle Scholar |
Pérez-Barbería, F. J., and Gordon, I. J. (2000). Differences in body mass and oral morphology between the sexes in the Artiodactyla: evolutionary relationships with sexual segregation. Evolutionary Ecology Research 2, 667–684.
Pérez-Barbería, F. J., Gordon, I. J., and Pagel, M. (2002). The origins of sexual dimorphism in body size in Ungulates. Evolution 56, 1276–1285.
Rudd, W. J., Ward, A. L., and Irwin, L. L. (1983). Do split hunting seasons influence elk migrations from Yellowstone National Park? Wildlife Society Bulletin 11, 328–331.
Rudge, A. B. J. (1984). ‘The Capture and Handling of Deer.’ (Nature Conservancy Council, Wildlife Advisory Branch: Peterborough, UK.)
Sánchez-Prieto, C., Carranza, J., and Pulido, F. J. (2004). Reproductive behavior in female Iberian red deer: effects of aggregation and dispersion of food. Journal of Mammalogy 85, 761–767.
| Reproductive behavior in female Iberian red deer: effects of aggregation and dispersion of food.Crossref | GoogleScholarGoogle Scholar |
Siegel, S., and Castellan, N. J. (1988). ‘Nonparametric Statistics for the Behavioural Sciences.’ (McGraw-Hill: New York.)
Springer, J. T. (1979). Some sources of bias and sampling error in radio triangulation. The Journal of Wildlife Management 43, 926–935.
| Some sources of bias and sampling error in radio triangulation.Crossref | GoogleScholarGoogle Scholar |
Staines, B. W., Crisp, J. M., and Parish, T. (1982). Differences in food quality in red deer (Cervus elaphus L.) stags and hinds in winter. Journal of Applied Ecology 19, 65–67.
| Differences in food quality in red deer (Cervus elaphus L.) stags and hinds in winter.Crossref | GoogleScholarGoogle Scholar |
Taillon, J., Sauvé, D. G., and Côté, S. (2006). The effect of decreasing winter diet quality on foraging behaviour and life-history traits of white-tailed deer fawns. The Journal of Wildlife Management 70, 1445–1454.
| The effect of decreasing winter diet quality on foraging behaviour and life-history traits of white-tailed deer fawns.Crossref | GoogleScholarGoogle Scholar |
Trivers, R. L. (1972). Parental investment and sexual selection. In ‘Sexual Selection and the Descent of Man’. (Ed. B. Campbell.) pp. 136–179. (Aldine Press: Chicago, IL.)
Watson, A., and Staines, B. W. (1978). Differences in the quality of wintering areas used by male and female red deer (Cervus elaphus) in Aberdeenshire. Journal of Zoology 186, 544–550.
Worton, B. J. (1989). Kernel methods for estimating the utilization distribution in home range studies. Ecology 70, 164–168.
| Kernel methods for estimating the utilization distribution in home range studies.Crossref | GoogleScholarGoogle Scholar |
