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.

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. 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. 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.
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