My first assignment for my behaviour course was to study the behaviour of an animal in captivity, and of course I chose a relative to the dog. It is a very limited study, but as a fist attempt at a bit of scientific writing I am quite pleased with it. So I am posting the whole thing here, perhaps it will be of some interest.
Pack cohesion
behaviour in the Dhole (Cuon alpinus)
in captivity
Introduction
The dhole (Cuon alpinus), or
Asiatic Wild Dog, is the only extant species of the Cuon genus of the Canidae
family. Widespread in Central, Southern and Eastern Asia in prehistoric times,
it is currently found mainly on the Indian subcontinent and in smaller numbers across
South-East Asia (Durbin et al. 2004).
The species is found in a range of habitats from tropical forest to high
mountainous regions (Maisch 2010). However, the dhole is now on the IUCN Red
List of Threatened Species, because loss of habitat and prey animals to human
activity, and persecution by humans in the past (IUCN 2011, Fox 1984).
The dwindling numbers of dholes in the last century, and the fact that
dholes are very wary of humans, means that the study of their behaviour in the
wild has been difficult (Johnsingh 1982, Cohen 1982, Fox 1984). Early problems
with breeding in captivity has also limited the number of observations of dhole
behaviour in captivity (Cohen 1982, Maisch 2010). However, in the last decade
the numbers of captive dholes has increased, as have studies of their behaviour
(Maisch 2010).
Dholes are medium-sized canids (12-20kg) with a red or brown, thick coat
and a darker bushy tail (Durbin et al. 2004).
Taxonomically the dhole has been placed together with the African wild dog (Lycaon pictus) and the South American bush
dog (Speothos venaticus) due to the lesser
number of post-carnassial molars found in these species, distinguishing them
from the genus Canis (including the wolf and the
domestic dog) (Davidar 1975, Durbin et
al. 2004). This classification has been questioned, however, attributing
these similarities in dentition to convergent evolution due to diet (Durbin et al. 2004). Recent genetic studies
confirm that the dhole is in fact more closely related to the genus Canis than previously believed (Durbin et al. 2004, Graphodatsky et al. 2008, Zhang and Chen 2010).
The dhole is a highly social animal, living in packs of up to 40
animals, although the average pack size is between 8-12 (Davidar 1975, Johnsingh
1982, Fox 1984). Dholes are highly co-operative, undertaking both hunting and
breeding as a group (Kleiman 1967, Venkatamaran et al. 1995, Fox 1984). They use a wide range of vocalizations for individual
recognition and pack co-ordination (Johnsingh 1982, Volodin et al. 2001, Durbin et al. 2004, Volodina et al. 2006).
The dhole displays a notably wide range of behaviours relating to pack
cohesion and hierarchy (Cohen 1982, Johnsingh 1982). From early studies onward
the predominance of socio-positive and submissive behaviours in establishing
and maintaining dhole pack hierarchy has been noted (Kleiman 1967, Davidar
1975, Cohen 1982, Johnsingh 1982, Fox 1984). This has been more recently contrasted
to the more fractious, dominance driven dynamics of the wolf pack (Durbin et al. 2004, Maisch 2010).
As the hegemony of interpreting the behaviour of the domestic dog (Canis familiaris) as an evolution from
the rigid pack structure of the wolf (Canis
lupus) is being reappraised (Koler-Matznick 2002, Bradshaw 2011), studies
of the behaviour of the dhole are ever more pertinent to an understanding of the
evolution of the Canidae. Bradshaw
has suggested that the behaviour of the common ancestor of both the domestic
dog and the wolf would have been different to that which has evolved in the
current living wolf, which is not easily domesticated (Bradshaw 2011). Indeed,
the dhole has been mentioned along other species of wild dogs as a more likely behavioural
analogue for an early ancestor of the Canis
genus (Koler-Matznick 2002). An ancestor animal with a predisposition to
cementing social relationships with submissive or playful interaction would
have not only been more easily domesticated, but would explain the relative
placidity and lack of aggression in the domestic dog as opposed to the wolf.
The aim of this study was to obeserve the social interaction of dholes
in captivity, to test the hypothesis that this interaction relies heavily on
socio-positive and co-operative behaviour rather than socio-negative and
antagonistic behaviour.
Animals and Study Area
The animals studied were in captivity in Howletts Wild Animal Park in
Kent, United Kingdom. The group studied consisted of twelve females, ranging in
age from one to eight years, including two sibling yearlings. All dholes had
been bred at Howletts.
The animals were housed in a roughly square enclosure measuring
approximately 50m x 50m. The enclosure consisted of level grassy terrain,
enriched with a number of trees and bushes, a small pond, two wooden platforms,
a group of concrete pipes, and a wooden shelter. Within the enclosure there was
a smaller fenced-off area of 20m x 20m, to which the doles had free access via
a small opening. See Figure 1.
 |
| Figure 1: Dhole enclosure |
Methods
The animals were observed from the accessible northwest and northeast
sides of the enclosure, primarily from the small viewing area at the western
corner.
Individual dholes are notoriously difficult to distinguish (Fox 1984,
Johnsingh 1982), and the animals were thus studied as a group. Thirty minutes
of initial ad libitum observation was undertaken between 3.30pm and 4.00pm on
21 November 2011, noting the full range of behaviours seen. The weather was
misty but dry, and visibility was fairly good on this day.
An ethogram was devised following these observations in conjunction
with previous studies of dhole behaviour (see Appendix I: Ethogram). From of this
ethogram, eleven categories of behaviour were chosen for the main scan sampling
study. All data gathering on vocalizations was abandoned, since it was not
possible to ascertain how many animals were producing sounds at any one time.
In addition, categories that were deemed less defined and/or more difficult to
observe reliably, such as sniffing and moving away from conspecifics were
dropped. During the main study it also became clear that the category of
approaching a conspecific was too difficult to observe in a number of
individuals at the same time, and it was effectively abandoned.
Data was then gathered for this selection of behaviours using the scan
sampling method, recording the number of animals performing each behaviour at 5
minute intervals. The data gathering session lasted two hours between 1.00pm and
3.00pm on the 24 November 2011. The weather went from overcast to sunny spells,
with good visibility.
There are several limitations to this study, notably the fact that the
group of dholes consisted exclusively of females. An all-male group of a similar
size exists at Howletts, however, so there is scope for a comparative study
across the sexes.
The visibility of the female dhole group was sometimes partial, due to
the amount of vegetation in the enclosure, and the limited area and elevation
from which observations could be made. Also, the observer was clearly visible
to the dholes, and they were aware of her presence, acknowledging it with
curiosity and sometimes fear and aggression. These facts, as well as the
relatively small enclosure available to the captive dholes, who have a range in
the wild of 40km squared (Johnsingh 1982), has to be kept in mind when
considering the results of this study.
In addition, the preliminary observations were made on a day on which
the dholes had been fed in the morning. The dholes are fed every other day, and
the scan sampling was thus undertaken on a day when the dholes were not fed. It
has to be assumed that this will have had some impact on the animals’
behaviour, although the keepers at Howletts suggested that their behaviour was
fairly uniform over the feeding and non-feeding days (pers. comm.).
Results
The scan sample data can be found in Appendix II. The results are
represented graphically in Chart 1. Note that behaviours which
were not observed at all during the scan sampling were excluded from the chart.
It has to be noted that from minute 75 onwards, the active period, at
several sample points not all dholes were visible to the observer.
During the first hour of observation the group was resting and
immobile the vast majority of the time. There was one case of an animal
defecating away from but near its sleeping site, and there were some movement
to rearrange resting positions in the other animals. On the whole, the animals
remained in the same places, in two groups of three, one group of two and four
solitary animals. The group of two appeared to be the two yearlings, judging by
their size, and were partially hidden in the wooden shelter.
Two of the four solitary animals appeared to have look-out or guarding
roles. Guarding in the dhole pack has indeed been reported by previous
observers (Johnsingh 1982, Fox 1984). They were placed on the extremes of the
area occupied by the resting pack, and showed more movement and alertness than
the other animals. However, as they were also intermittently resting they were
not counted as guarding on the scan sampling data chart. The resting positions
of the animals from 0-60 minutes are recorded in Figure 2.
 |
| Figure 2: Resting positions of Dholes |
At the interval 65-70 minutes into the study, one of the yearlings
emerged from the shelter. After a few minutes of stretching and looking around
it rushed towards Group 1, three adults, making the characteristic repeated yipping
sounds of the dholes. It engaged in begging behaviour towards one of the larger
animals. This raised the whole group into intense activity, and multiple cases
of begging behaviour took place. At any time in this interval there were three
or four groups of two or three animals engaged in begging behaviour directed
towards one individual in each group.
From this time and for the rest of the study, 70-120 minutes, the
dholes remained relatively active. Apart from multiple instances of begging,
guarding and patrolling recorded, and the frequent reorganization of small
groups of sitting and lying dholes, the animals engaged in some brief chase
games and play fighting. During this period of activity the dholes frequently
vocalized, making a repeated yipping sound. It appeared to be made by several
if not all individuals, at intervals of a few seconds, lasting from ten seconds
to several minutes at a time.
There was an instance of communal defecation. Initially three to four
individuals defecated on the same spot, in the interval 80-85 minutes into the
study. The place was later repeatedly visited, sniffed and defecated on by
other individuals throughout the rest of the observation period.
Apart from one staring match observed during the preliminary study, no
fights or threat behaviour was observed among the dholes, although some begging
behaviour was met with brief inhibited biting. In contrast, the dholes growled
at the observer on a couple of occasions. This was usually performed by a
single animal, in one case reminiscent to the “grumble on hind legs” decribed
by Volodin et al. 2005 (See ethogram
in Appendix II). Towards the end of the observation time, between 110-115
minutes, almost the whole group approached the fence in front of the observer,
apparently led by a one of the larger animals. The group was yipping excitedly
and two of the larger animals reared up, growled and bit at the fence. After a
few minutes the group seemed to lose interest and carried on with their other
activities.
Discussion
All behaviour observed was of a neutral or socio-positive kind. Two
characteristics of the behavioural data gathered stand out:
1) Approach and interaction between dholes was mainly performed
through begging behaviour, which, as there was no food present at the day of
the main observation, must be interpreted as a submissive greeting and social
bonding ritual (Maisch 2005). In the active period, begging was frequent (6 out
of 11 sample points) and undertaken by a significant proportion (10-25%) of the
individuals observed. In addition to this some play was observed, although not
sampled.
Also, the over-marking of feaces by the group was observed,
effectively producing a communal latrine. It has been suggested that this
behaviour serves an intra-group communicative function, rather than being a territorial
marking (Cohen 1982, Johnsingh 1982, Durbin et
al. 2004). However, it has to be noted that the latrine was placed on the
edge of the dhole enclosure, bordering that of the African wild dogs (see
Figure 1 and 2). There may therefore also have been a territorial element to
this behaviour in this case.
2) Throughout the observation period a significant proportion of the
dholes were engaged in behaviours undertaken in a group, either passively
sitting or lying together, or actively interacting in the begging ritual. On
average, these behaviours together accounted for 55% of individuals at any one
time. If patrolling is included as a group behaviour, the average rises to 62%
of individuals at any one time.
The yipping noise made by many dholes while patrolling has been
interpreted as a way of coordinating the group’s movement (Johnsingh 1982, Fox
1884, Durbin et al. 2004, Volodina et al. 2006, Maisch 2010), suggesting that
patrolling can be seen as a group behaviour, although the data gathered by this
study is not sufficient to distinguish between group and solitary patrolling.
It was also notable that throughout the initial period of inactivity, the
dholes were resting in well-defined groups, as seen in Figure 2. It was clear that
Group 1 (see Figure 3.), which included the largest animals, as well as Group
2, were composed of higher ranking animals. These two groups occupied the
highest points in the enclosure, the two wooden platforms. Indeed, these
animals seemed to be among the ones mostly at the receiving end of begging
behaviour, although as mentioned, the reliable distinction of individuals was
not possible. The smaller group of two yearlings that were hiding in the shelter,
on the other hand, appeared to be the instigators of much begging.
 |
| Figure 3: Group 1 consisting of three large adult dholes on Platform 1. |
Thus, even the sleeping formation of the dhole pack seems to be related to the social relationships within the group, and must be seen as a behaviour that positively aids social cohesion.
The prolonged period of inactivity at the beginning of the observed period may be explained by two factors. 1) The dholes had no food in the enclosure, the presence of which had been a reason for some activity seen during preliminary observations. 2) Dholes are often, although not exclusively, crepuscular (Johnsingh 1982, Durbin et al. 2004), and the period of activity seemed to correlate with approaching dusk. Indeed the preliminary observations, in which the dholes were more active, were undertaken later in the day.
The prolonged period of inactivity at the beginning of the observed period may be explained by two factors. 1) The dholes had no food in the enclosure, the presence of which had been a reason for some activity seen during preliminary observations. 2) Dholes are often, although not exclusively, crepuscular (Johnsingh 1982, Durbin et al. 2004), and the period of activity seemed to correlate with approaching dusk. Indeed the preliminary observations, in which the dholes were more active, were undertaken later in the day.
The data gathered in this brief study does seem to indicate a mainly
socio-positive and co-operative behavioural pattern in the dhole pack. This
result correlates with earlier observations of the pack-cohesion of the dhole
(Kleiman 1967, Cohen 1982, Johnsingh 1982, Fox 1984). The predominance of
submissive displays and play behaviour in the Cuon alpinus is particularly interesting in the context of the Canidae family and the evolution of the Canis genus. The majority of canids are far
less socially co-operative (Kleiman 1967, Fox 1976, Bradshaw 2011) than the Canis genus and its precursors. In
addition, the placidity and predisposition to submissive social bonding of the
domestic dog contrasts with the dominance displays of the current wolf. The
similarities between the socio-cohesive behaviour of the dhole and the domestic
dog (Davidar 1975) suggests and interesting avenue of investigation of the Cuon alpinus as a more apt behavioural
analogue of the ancestors of the Canis genus,
and precursors to the domestic dog.
References
Bradshaw, J. (2011) In Defense
of the Dog: Why dogs need our understanding London: Penguin.
Cohen, J. A. (1982) ‘A Note on the Behaviour of captive
dholes (Cuon alpinus).’ Journal of the Bombay Natural History
Society, 62: 146-148.
Durbin, L.S.,
Venkataraman, A., Hedges, S., and Duckworth, W. (2004) ‘Dhole.’ In
Sillero-Zubiri, C., Hoffmann, M., and Macdonald, D. W. (eds.) Canids: Foxes, Wolves, Jackals and Dogs -
2004 Status Survey and Conservation Action Plan. IUCN/SSC Canid Specialist
Group, 210-219. Available at: http://www.canids.org/cap/index.htm#On-line%20Copy%20of%20the%20Action%20Plan
(Accessed 17 November 2011).
Davidar, E. R. C.
(1975) ‘Ecology and the behaviour of the Dhole or Indian Wild Dog (Cuon alpinus).’ In Fox, M. W. (ed.) The Wild Canids: Their Systematics,
Behavioural Ecology and Evolution Wenatchee: Dogwise Publishing.
Fox, M. W. (1984) The Whistling Hunters: Field Studies of the
Asiatic Wild Dog (Cuon alpinus)
Albany: Universiy of New York Press.
Graphodatsky, A. S., Perelman, P. L.,
Sokolovskaya, N. V., Beklemisheva, V. R., et al. (2008) ‘Phylogenomics of the
dog and fox family (Canidae, Carnivora) revealed by chromosome
painting.’ Chromosome Research 16(1), 129-143.
Karanth, K. U., and Sunquist, M. E.
(1995) ‘Prey selection by tiger, leopard and dhole in tropical forests.’ Journal of Animal Ecology 64(4),
439-450.
Karanth, K. U., and Sunquist, M. E.
(2000) ‘Behavioural correlates of predation by tiger (Panthera tigris), leopard (Panthera
pardus) and dhole (Cuon alpinus)
in Nagarahole, India.’ Journal of Zoology
250(2), 255-265.
Kleiman, D.G. (1967) ‘Some Aspects of
Social Behavior in the Canidae.’ American Zoologist 7(2), 365-372.
Koler-Matznick, J. (2002) ‘The origins
of the dog revisited.’ Antrhozoos 15(2),
98-118.
Johnsingh, A. J. T.
(1982) ‘Reproductive and social behaviour of the Dhole, Cuon Alpinus (Canidae).’ Journal of Zoology, 198, 443-463.
Maisch, H. (2005) Ist das Fortpflanzungssystem bei Rothunden
Cuon alpinus die Ursache oder eine Konsequenz des Rudellebens? Dissertation
at Fachbereich Biologie/Chemie der Universität Osnabrück. Available at: http://repositorium.uni-osnabrueck.de/handle/urn:nbn:de:gbv:700-2006041112
(Accessed: 17 November 2011).
Maisch, H. (2010) ‘The influence of husbandry and pack management
on Dhole Cuon alpinus reproduction.’ International Zoo Yearbook 44, 149-164.
Venkataraman, A. B. Arumugam, R., and Sukumar, R., (1995) ‘The
foraging ecology of dhole (Cuon alpinus)
in Mudumalai Sanctuary, southern India.’ Journal
of Zoology 237(4), 543-561.
Volodin, I. A., Volodina, E. V., and Isaeva, I. V. (2001) ‘Vocal
repertoire in the dhole Cuon alpinus
(Carnivora, Canidae) in captivity.’ Entomological Review 81(1), S161-S166.
Volodina, E. V., Volodin, I. A., Isaeva, I. V. and Unck, C. (2006)
‘Biphonation may function to enhance individual recognition in the dhole, Cuon alpinus.’ Ethology 112(8), 815-825.
Wang, S. W. and Macdonald, D. W. (2009) ‘Feeding habits and niche
partitioning in a predator guild composed of tigers, leopards and dholes in a
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of Zoology 277(4), 275-283.
Zhang, H. & Chen, L. (2010) ‘The
complete mitochondrial genome of dhole Cuon
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Appendix I: Ethogram
Sources:
Observation ad libitum of all-female
group of twelve individuals aged 1-8 years, captive in Howletts Widlife Park,
3.30pm -4.00pm on 21 November 2011.
Behaviour observed by Volodin 2001, and
Maisch 2005.
Behaviour selected to be studied by scan
sampling in the main data gathering session of this study are indicated in
bold.
Vocalisations:
Yip
|
Short squeaking noise.
Responses: yips / no response.
|
Grumble
|
Longer growling noise.
Responses: quick retreat / no response.
|
Grumble on hind legs
|
Longer growling noise, standing on hind legs
facing the “threat”.
|
Movement:
Guarding
|
Individual sitting or standing near perimeter of
enclosure, paying attention to the outside.
|
Patrolling
|
Moving at a moderate
pace, without a specific goal. Alert and attentive to other dholes. Often
accompanied by repeated “yip” vocalization.
|
Approaching other dhole
|
Walking or running
towards another dhole and coming within 1 meter of other individual.
|
Moving away from other dhole
|
Purposefully walking or running away from other
individual, increasing distance to over 5 feet. Not a chase game.
|
Sniffing ground
|
Stationary or walking, nose to the ground.
|
Sniffing conspecific
|
Stationary or walking, nose close to conspecific.
|
Sitting or lying down –
solitary
|
Animal sitting or lying
further than 1 meter away from other animal.
|
Positive social
interactions:
Sitting or lying down –
group
|
Animal sitting or lying
within 1 meter of other animal.
|
Begging
|
Approach to conspecific,
with head held low, ears lying back. Tail wagging and/or coiled on the side.
Contact made with snout, often licking conspecific’s muzzle.
May be accompanied by
pawing, circling of the conspecific, and whining.
Responses: snarling, inhibited
muzzle biting, reciprocating, playing, giving food (if food is around),
denying food, moving away, passive.
|
Chase game
|
Enticing chase by
approaching and then quickly running away from conspecific, which follows.
Roles may be swapped. May lead to play fight.
|
Play fight
|
Enticed by play bowing,
nudging or muzzle nipping. Involves one individual playing submissive, head
held low, rolling over, and other individual standing over or jumping over,
nudging, bumping and playfully biting neck of “submissive” animal. Roles
frequently reversed. May be combined with chase game.
|
Negative social
interactions:
Staring match
|
Both parties head held
low, ears back, eyes almost closed. Stare at each other and try to wait each
other out. May be accompanied by growling.
|
Wrestling fight
|
Standing on hind legs,
trying to press opponent to the ground. The loser runs away. Growling and
“miii” sounds may be made.
|
Threat
|
No physical contact.
Tail held horizontal, u-shaped or up. Hackles raised. Ears and eyes pointing
at opponent.
Back may be arched, legs
may be stiff and pushing into ground.
|
Appendix II: Scan Sample Data and Calculations
