Clan Groups: Mixed sex aggregation outside of the rut?
Paul Rattray

Paul is a project manager with   Abstract
CVC Network...
3. Mixed sex aggregation in red deer
Given these interrelationships affecting aggregation, the primary question to be
expl...
clan groups. Based on these observations, a “clan group” is a group of red deer
consisting of a mature stag and a small nu...
Jesser, Peter. (2005) “Deer Pest Status Review Series – Land Protection Deer Family cervidae in
        Queensland”, Depar...
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Clan Groups: Mixed sex aggregation outside of the rut?

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Based on anecdotal evidence, this paper proposes the existence of mixed sex groups that congregate outside of the mating season consisting of a mature stag and a small number of hinds, described as a “clan group”. Clan groups may help further explain mixed sex aggregation, predator avoidance and migration behaviours in red deer.

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Transcript of "Clan Groups: Mixed sex aggregation outside of the rut?"

  1. 1. Clan Groups: Mixed sex aggregation outside of the rut? Paul Rattray Paul is a project manager with Abstract CVC Network, a global media Research into red deer behaviour suggests that mature red deer aggregate into mixed sex groups organisation and an avid red during the mating season or rut and same sex groups outside of the rut. The only exception is linear deer hunter and researcher. groups usually consisting of a mature female hind leading its immature young. Based on anecdotal evidence, this paper proposes the existence of another mixed sex group that congregates outside of the mating season consisting of a mature stag and a small number of hinds, described as a “clan group”. Clan groups may help further explain mixed sex aggregation, predator avoidance and migration behaviours in red deer. Keywords Red deer, clan group, sexual segregation, ranging patterns, migration behaviours 1. Red deer aggregation/segregation Research into the aggregating behaviour of red deer identifies mixed sex groups congregating during the mating season then separating into sexually segregated groups during the rest of the year (Bonenfant, Loe, Mysterud, Langvatn, Gaillard & Klein, 2004). While the mechanisms causing aggregation and segregation in red deer are still debated, researchers explain such behaviour as relating to differences in a) body size, b) activity budgets and 3) risk strategies (Bonenfant, Loe, Mysterud, Langvatn, Stenseth, Gaillard & Klein, 2004) Differences in body size or sexual dimorphism relates to selection resulting from competition with one another for access to the other sex, genetic preferences and nutritious food (Catchpole, Morgan, Clutton-Brock & Coulson, 2004). The main prediction of the activity budget hypothesis is that animals with similar body sizes, energy needs and reproductive strategies aggregate socially and therefore segregate from animals with different body size and energy needs (Ruckstuhl & Neuhaus, 2000). Risk strategies, particularly in successful predator avoidance differ markedly between red hinds and stags, with studies showing females to be far more effective at evading predators than males (Mathisen, Landa, Andersen & Fox, 2003). 2. Factors causing aggregation The interrelatedness of aggregation/segregation theories come to light when applied in the field. Smietana’s 2005 study of wolf predation selectivity on red deer in the Bieszczady Mountains of Poland found predation on male deer to be higher due to the segregation of social units (hind-calf and stag groups), except during the rutting season, and the low fat reserves of males from midwinter due to a less nutritious diet than females. Jiang, Liu, Zeng, Han and Hu’s (2000) study of Pierre David’s deer in China also showed comparable predator avoidance strategies. However these authors hypothesised that segregation may result naturalistically from sexual repulsion to the opposite sex as a reproductive strategy. Similar predator avoidance strategies were identified by Bonenfant et al (2004) in their study of red deer in France and Norway. Interestingly, in these two latter studies, red deer appear to have behaved mechanistically, since large predators such as wolves are extinct in France and China and rare in Norway. A synthesis of long-term research studies into red deer aggregation and segregation conclude that numerous factors relating to body size, activity budgets and risk strategies combine to influence such behaviours (Johnson, Wisdom & Cook, 2005, Neuhaus & Ruckstuhl, 2004). Thus, while each factor is plausible none is likely to operate independently of the other factors.
  2. 2. 3. Mixed sex aggregation in red deer Given these interrelationships affecting aggregation, the primary question to be explored in this paper is why mixed sex aggregation in a clan group consisting of a mature stag and a small number of hinds would continue outside of the rut and what the benefits to such a grouping might be. Long-term studies into the ranging patterns of red deer consistently show distinctively different ranging patterns between mature males and females (Clutton-Brock, Coulson, Milner- Gulland, Thomson & Armstrong, 2002). As a rule, such findings show that mature males leave their maternal birth areas and move to new areas (Clutton- Brock, Pemberton, Kruuk & Coulson, 2008). Mature females, on the other hand, are found to “heft” or bond to their birth area for life (Pottie, 2005). Evidence of exceptions to these findings occurs when red deer ranges are expanding into new areas or shifting between habitats (Szemethy, Mátrai, Bíró & Katona, 2003, Biro, Szemethy, Katona, Heltai & Peto, 2006). Furthermore, findings from studies of individual variations in migratory and exploratory movements and habitat use by adult red deer show similar movement patterns for both sexes (Luccarini, Mauri, Ciuti, Lamberti & Apollonio 2006, Kamler, Jedrzejewska & Jedrzejewski, 2007, Pepin, Adrados, Janeau, Joachim & Mann, 2008). It is worth noting, however, that these findings also demonstrated site fidelity in that individual red deer found outside the limits of their familiar area eventually returned to these familiar areas. 4. Range establishment of feral red deer One of the limitations with research into red deer aggregation and segregation is that virtually all studies have been conducted in areas of Europe where red deer are native to an area and have long-established ranges. Thus a pertinent question to our study is whether such aggregation dynamics change when red deer colonise new areas, such as Australia and New Zealand, as feral animals. Where red deer populations are expanding it is clear that stags and hinds do move in to colonise new areas. The logic for this thinking is obvious. Otherwise red deer would not have colonised the vast areas that they have as feral, non- native animals in Australia, New Zealand and other parts of the world. Interestingly there are few empirical studies about how colonisation of or migration to new territories actually occurs. Davidson and Kean (circa 1950), scientific officers of New Zealand’s Forestry Service, studied the establishment of red deer in the Tararua Mountains of New Zealand’s north island. They noted that the pattern of red deer establishment tends to be that of stags moving out into new territories followed by hinds. Exactly how this process of range establishment occurs between sexes was not further defined. Similar studies of red deer in Southern Queensland, where most feral red deer reside in Australia, also fail to adequately explore range establishment and group composition as a primary factor in red deer management (McGhie, & Watson, 1995, Jesser, 2005). These limitations are particularly pertinent as red deer are now classified by the Queensland Government (2009) as Class 3 pests in Queensland because they have, or could have, an adverse economic, environmental or social impact. It is somewhat surprising that red deer are acknowledged as being pests due to them colonising new areas of the state, yet there are no specific studies as to how red deer colonise such areas. 5. Clan groups It is the contention of this paper, based on anecdotal evidence from a number of areas hunted by its author, that feral red deer migrating to new areas may utilise
  3. 3. clan groups. Based on these observations, a “clan group” is a group of red deer consisting of a mature stag and a small number of hinds that aggregate together outside of the rut. There are three main sites in Southeast Queensland, Australia where the author has observed red deer in what appear to be clan groups: 1) Woolooga (2007), 2) Wonga (2008) and 3) Widgee (2009). According to local land holders, Sites 1 and 3 have only been colonised by red deer during the last 15 years, mostly likely from Site 2, which has had a longer history of colonisation, possibly 30 years or so. It is important to note however that this information is only based on anecdotal reports and personal observations. 6. Conclusion In conclusion, to ascertain the existence of clan groups as mixed sex groups of adult red deer that remain together for extended periods outside of the rut and the degree to which clan groups may play a role in red deer migration to new territories, requires further scientific research. Long-term radiotelemetry (radio- tracking) of the ranging behaviour of feral, male and female red deer similar to studies by Szemethy et al (2003), Luccarini et al (2006) and Kamler et al (2007) are recommended. Further research, utilising direct observations of red deer ranging behaviour by involving local landholders and hunters along the lines of Mysterud, Meisingset, Veiberg, Langvatn, Solberg, Loe and Stenseth (2007) in Sweden and Janiszewski, Niczyporuk and Hanzal (2007) in Poland who monitored live and harvested red deer numbers may also shed light on clan group activity. Another helpful area of research could measure the foraging ecology of red deer using organic matter digestibility (OMD) and composition of the diet of a target population of red deer males and females (Bugalho, Milne & Racey 2001). For southern Queensland, where feral dogs and native dingoes abound, a study of predation patterns, similar to Smietana’s (2005) wolf study and would also assist in determining whether predators influence clan group behaviour. Combining these research methodologies to study a population of feral male and female red deer could shed further light on clan groups in particular and provide valuable information to policy makers, landholders and hunters as to how feral red deer migrate to and colonise new territories. 7. References Biro, Z., Szemethy, L. Katona, K., Heltai, M. & Peto, Z. (2006), “Seasonal distribution of red deer (Cervus elaphus) in a forest-agriculture habitat in Hungary”, Mammalia (2006): 70–75. Bonenfant, C. Loe, L. E., Mysterud, A., Langvatn, R. Stenseth, N. C., Gaillard, J. M. & Klein, F. (2004) “Multiple causes of sexual segregation in European red deer: enlightenments from varying breeding phenology at high and low latitude”, PROCEEDINGS ROYAL SOCIETY LONDON, 271, 883-892. Bugalho, M. N., Milne, J. A. & Racey, P. A. (2001) “The foraging ecology of red deer (Cervus elaphus) in a Mediterranean environment: is a larger body size advantageous?” JOURNAL OF ZOOLOGY, 255(3), November 2001 pp 285-289. Catchpole E. A., Fan Y., Morgan B. J. T., Clutton-Brock T.H. & Coulson T. (2004) “Sexual dimorphism, survival and dispersal in red deer”, JOURNAL OF AGRICULTURAL, BIOLOGICAL & ENVIRONMENTAL STATISTICS, Vol 9, No 1, 1 March 2004 , pp. 1-26(26). Clutton-Brock, T. H., Coulson, T. N., Milner-Gulland, E. J., Thomson, D. & Armstrong, H. M. (2002). “Sex differences in emigration and mortality affect optimal management of deer populations,” NATURE, Vol 415(7), February 2002, www.nature.com. Clutton-Brock, T. H., Pemberton, J. Kruuk, L. & Coulson, T. (2008), "Red Deer Research on the Isle of Rum - Research Findings," http://www.zoo.cam.ac.uk/zoostaff/larg/pages/Rumresearch.html, Downloaded: Monday 7 July 2008. Davidson, M. M. & Kean, R. I. (circa 1950) “Establishment of Red Deer Range in Tararua Mountains”, NEW ZEALAND FORESTRY SERVICE, Auckland, pp. 293-324. Janiszewski, P., Niczyporuk, M. & Hanzal, V. (2007) “Quality of the Red Deer (Cervus Elaphus) Harvested in Hunting Grounds of the Bialowieza Primeval Forest”, ACTA ZOOLOGICA LITUANICA, 2007, Vol. 17, No. 3, pp. 228-233.
  4. 4. Jesser, Peter. (2005) “Deer Pest Status Review Series – Land Protection Deer Family cervidae in Queensland”, Department of Natural Resources and Mines, Queensland Government: Brisbane. Jiang. Z., Liu, B., Zeng, Y., Han, G & Hu, H. (2000). “Attracted by the same sex, or repelled by the opposite sex?—Sexual segregation in Pere David’s deer”, CHINESE SCIENCE BULLETIN, 45(6), March, 2000. Johnson, B. K, M. J. Wisdom & J. G. Cook (2005) “Issues of Elk Productivity for Research and Management”, pages 81-93 in Wisdom, M. J., technical editor, The Starkey Project: a synthesis of long-term studies of elk and mule deer, reprinted from the 2004 Transactions of the North American Wildlife and Natural Resources Conference, Alliance Communications Group, Lawrence, Kansas, USA. Kamler, J. F., Jedrzejewska, B. & Jedrzejewski, W. (2007), “Factors affecting daily ranges of red deer Cervus elaphus in Bialowieza Primeval Forest, Poland”, ACTA THERIOLOGICA 52(2), April 2007, 113-118(6). Kamler, J. F., Jêdrzejewska, B. & Mioecicki, S. (2004), Red deer – a tale of two deer, “Essays on Mammals Białowieza Forest,”Bogumi Jêdrzejewska and Jan Marek Wójcik (Eds.) Mammal Research Institute, Polish Academy of Sciences, Poland: Białowieza, pp 51-58. Luccarini, S., Mauri, L. Ciuti, S., Lamberti, P. & 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 & EVOLUTION, 18 127-145. Mathisen, J. H., Landa, A., Andersen, R. & Fox, J. L. (2003) “Sex-specific differences in reindeer calf behaviour and predation vulnerability”, BEHAVIORAL ECOLOGY Vol. 14 (1), pp. 10-15. McGhie, C. J. & Watson, S. (1995) Queensland wild deer and their role in sustainable wildlife management, in “Conservation through sustainable use of wildlife”, Ed. by G. C. Grigg, P. T. Hale and D. Lunney, Brisbane, Queensland Australia: Centre for Conservation Biology, The University of Queensland. Mysterud, A., Meisingset, E.L., Veiberg, V., Langvatn, R., Solberg, E.J., Loe, L.E. & Stenseth, N.C. (2007) “Monitoring population size of red deer Cervus elaphus: an evaluation of two types of census data from Norway”, WILDLIFE BIOLOGY, 13(3) 285-298. Neuhaus, P. & Ruckstuhl, K. E. (2004) “A Critique: Can the Activity Budget Hypothesis Explain Sexual Segregation in Desert Bighorn Sheep?” BEHAVIOUR 141, 513-520. Pepin, D., Adrados, C., Janeau, G., Joachim, J. & Mann, C. (2008) “Individual variation in migratory and exploratory movements and habitat use by adult red deer (Cervus elaphus L.) in a mountainous temperate forest”, ECOLOGICAL RESOURCES, 23 1005-1013. Pottie, S. (2005) “Species Profile - Red Deer”, CALEDONIA WILD! Spring 2000, Trees for Life, The Park, Findhorn Bay, Forres IV36 3TZ, Scotland, UK. Queensland Government (2009), “Queensland Land Protection (Pest and Stock Route Management) Amendment Regulation (No. 1) 2009, Subordinate Legislation 2009 No. 50 made under the Land Protection (Pest and Stock Route Management) Act 2002. Ruckstuhl, K. E. & Neuhaus, P. (2000) “Sexual Segregation in Ungulates: A New Approach”, BEHAVIOUR 137, 361-377. Smietana, W. (2005) “Selectivity of wolf predation on red deer in the Bieszczady Mountains, Poland”, ACTA THERIOLOGICA, 50(0): 1–12. Szemethy, L., Mátrai, K., Bíró, Z. & Katona, K. (2003), “Seasonal home range shift of red deer in a forest-agriculture area in southern Hungary.” ACTA THERIOLOGICA, 48(4), October 2003, pp. 547-556. (Mysterud, Meisingset, Veiberg, Langvatn, Solberg, Loe & Stenseth, 2007).

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