Table Of ContentSTATUS, ECOLOGY AND CONSERVATION OF THE INDIAN WOLF
CAMS LUPUS PALLIPES SYKES
(With two text-figures and one plate)
Yadvendradev Jhala1
Key words: Food habits, human - wolfconflict, threats, distribution
Two wolfsubspecies were believed to inhabit the Indian subcontinent, the Tibetan wolf(Canis
lupus chanco) whose range extends from the trans-Himalaya into Tibet and China, and the Indian
wolf Canis lupuspallipes) that ranges over much ofpeninsular India. Recent genetic data shows
(
three extant wolflineages in the Indian subcontinent. Two ofthese, the Himalayan and peninsular
lineages, are ancient and unique to the Subcontinent, while wolves from Kashmir belong to the
widespread wolf-dog clade. The Indian wolf (peninsular clade) inhabits semi-arid agro-pastoral
landscapes and scrub forests. It is considered endangered, with its numbers ranging between 2,000-
3,000 individuals. Undisturbed habitat patches of5-1 5 sq. km that offer good cover and water are
critical for successful breeding in established packs. Wolf territories range between 150 and
300 sq. km and are a function of prey and denning habitat availability. Indian wolves whelp in
December-January in an underground den excavated by the alpha pair. The mean litter size is 4.8
pups. Juvenile wolves can disperse from their natal pack after the age of8 months. The majority of
wolf populations in India survive outside protected areas, and subsist primarily on livestock. In
some areas, wolves have been reported to attack children. The common factors in such areas are
high human population, poverty, too few orheavilyguarded livestock, and poorwild prey availability.
Human-wolfconflicts are ofserious magnitude over much ofthe wolf’s range in India. Wolves are
persecuted by smoking pups in their dens, sometimes by shooting and recently by poisoning.
Majorthreats to the continued survival ofwolves in Indiaare persecution by poisoning, and loss of
denning habitat to intensive agriculture, development, and industry.
Introduction much smaller in comparison to other subspecies
of wolves, except C.l. arabs that is found in the
Canis lupus once had the largest natural Arabian peninsula.
range ofany land mammal, besides Homosapiens Molecular genetic data from wolfand dog
(Sheldon 1992), and had successfully colonised populations from around the world suggests that
much ofthe Northern hemisphere (Mech 1970). they belong to a closely related wolf- dog clade
Out ofthe 32 odd subspecies of wolves that are (Vila etal. 1997). Genetic analysis ofmitochondrial
DNA
currently recognised (Mech 1974), two are (control region and cytochrome b) ofwolves
believed to occur in the Indian subcontinent. from the Indian subcontinent has shown that there
Canis lupus chanco or the Tibetan wolfis found are three extant lineages, two of them very
in the trans-Himalayan region and its range different from the wolf- dog clade. Wolves from
extends into Tibet, China, Manchuria and peninsular India considered to be C.l. pallipes
Mongolia. The Indian wolf C.l. pallipes ranges may have diverged from the wolf- dog clade about
over much of peninsular India and the same 500.000 years ago and are different from the
subspecies is believed to occur in Iran and Israel pallipes found in the Middle East. Wolves from
(Mendelssohn 1982, Shahi 1982). C.l pallipes is Himachal Pradesh to eastern Nepal (considered
to be C.l. chanco) are basal to the other wolf
'Wildlife Institute of India, RO. Box No. 18, clades and may have separated from them about
Chandrabani, Dehradun, Uttaranchal 248 001, India.
Email: [email protected] 800.000 years ago, while wolves westofKashmir
JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003 293
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ECOLOGY AND CONSERVATION OF THE INDIAN WOLF
belong to the widespread wolf- dog clade. Thus, Status, Distribution and Habitat Needs
the peninsular and Himalayan wolf lineages of
India are very ancient and unique to the Indian C.l. pallipes is considered endangered in
subcontinent (Sharma et al., in press). India and features on Schedule 1 of the Indian
Almost nothing is known about the ecology Wildlife (Protection) Act of 1972 and ofCITES
and status ofthe Himalayan wolflineage. In these (Shahi 1982). Killing ortrading ofC.l. pallipesis
regions, persecution of wolves is common by prohibited by law in India. Shahi (1982) in his
pastoralists (Fox and Chundawat 1992), and it is preliminary survey estimatedthe wolfpopulation
likely that the Himalayan lineage would probably in India to be about 800 individuals that were
be one of the most endangered canids in the patchily distributed over peninsular India (Fig. 1).
world. The account that follows is of the More intensive surveys in the state of Gujarat
comparatively well-studied peninsular lineage of and Rajasthan have shown that wolfdistribution
the Indian wolf(Plate 1, Fig. 1). is continuous in these states. The population of
Adultmale and female Indian wolves weigh wolves in these two states was estimated to be
between 19-25 kg and 17-22 kg respectively. between 450 and 620 (Jhala and Giles 1991).
Adult wolves measure between 103 and 145 cm Analysis of recent surveys, coupled with the
from nose tip to tail tip, and between 57 and dispersal capability ofwolves (Mech 1995) makes
72 cm at shoulder height. Coat colour is sandy it likely that the wolfpopulation within the states
brown with black hair tips. Some wolves have a of Gujarat, Rajasthan, Haryana, Uttar Pradesh,
more rufous tinge to their coats. Adult wolves Madhya Pradesh, Maharashtra, Karnataka and
can be distinguished from juveniles in the field Andhra Pradesh is continuous (Fig. 1).
by their size, behaviour, and white markings that The current estimated distribution ofwolves
develop above the eyes, on the chin and under covers amuch largerarea(Fig. than was reported
1)
the throat. Colour variation is rare among Indian earlier. This does not imply range extension by the
wolves, black wolves being reported only from wolf, but is a result ofmore intensive surveys and a
Ladakh. In much of the wolfs range in India, better understanding ofwolfdistribution. The wolf
summer temperatures up to 47 °C are not is still far from safe in most ofits range and occurs
uncommon, while winter temperatures at low densities (about 1 wolfper 100-200 sq. km,
occasionally approach 0 °C. In summer, most of JhalaandGiles 1991). Highdensitywolfpopulations
the fur is shed, only sparse long hair remains on (upto 5 wolvesper 100 sq. km) are foundto occur in
the wolfs body. This gives the wolf a scrawny, somehabitatpocketsandpreserves. Breedingpacks
long-legged appearance. ranging from 4 to 14 wolves have been reported
Though the wolfis believedto have evolved from the Bhal, Dwarka, Banni, and Abdasa area in
as a temperate species, the Indian wolf is aptly Gujarat and Kutch; Kumbhalgarh, Gudda-Bishnoi,
adapted for living in semi-arid and hot and Pali-Barmer and Jodhpur areas in Rajasthan;
environments. The small body size reduces food Nannaj, Rehukuri, Nasik, and Phaltan areas in
demands, permitting it to sustain its populations Maharashtra; Neoradehi in Madhya Pradesh;
on smaller ungulates, lagomorphs and rodents; Rollapadu in Andhra Pradesh; Melkote and
the shedding of under fur and behavioural Ranibennur in Karnataka; and Mahuadaur,
thermoregulation permits this canid to live in hot HazaribagandPalamauareas in Bihar(Fig. 1). Such
and aridregions. The Indian wolfstill needs ample high-density habitats are extremely important for
drinking water, and in that sense is not truly wolf conservation, since these pockets serve as
adapted to desert living like the chinkara Gazella successful breeding and recruitment areas from
(
bennettii). wherewolves disperseto occupy marginal habitats.
294 JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003
ECOLOGY AND CONSERVATION OF THE INDIAN WOLF
Fig. 1 : Distribution ofthe wolfCams lupuspallipes in India. Locations ofknown breeding populations
ofwolves, areas where wolfstudies have been conducted and areas where wolfattacks
on children have been reported are shown
In the light of current information and in The eastern population of C.l. pallipes
,
concurrencewith GinsburgandMacDonald (1990), found in Orissa, Bihar and parts of West Bengal,
apopulation estimate ofabout2,000-3,000 wolves is an exception and occurs in moister forested
for the Indian peninsula seems more realistic habitats (Shahi 982), but even here wolves are
1
(Jhala 2000) and may well be a conservative not reported where thick forests occur. Wolves
underestimate. occur on the periphery of protected forest areas
Since wolves are believed to have evolved as is seen in Kumbhalgarh and Kailadevi
in boreal forest systems as predators of large Sanctuaries in Rajasthan; Panna, Kanha, and
ungulates, it is rather surprising that the Indian Bandhavgarh National Parks in Madhya Pradesh;
wolfrarely lives in forests and prefers scrubland, and Girin Gujarat. These peripheral areas are under
grassland and semi-arid pastoral/agricultural heavy biotic pressure from surrounding human
landscapes. The Indian wolf probably evolved populations and are reduced to scrub forests. It
duringthe drier spells ofthe Pleistocene to exploit is these scrublands that wolves primarily use.
a relatively unoccupied niche as a top carnivore Wolves do not seem to be restricted in their
ofthe arid zones. movements at night and even approach very close
JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003 295
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ECOLOGY AND CONSERVATION OF THE INDIAN WOLF
to human dwellings and villages (Jhala 1991). In 1,000 sq. km (Jhalaand Sharma 1997). Food, water,
Maharashtra, radio-collared wolves were and availability of habitat for denning and
observed foraging in the Ojhar township ofNasik, rendezvous sites are factors determining territory
at night. However, during the day, wolves are size (Fuller 1989, Jhala 1991). Wolves subsisting
extremely selective in their habitat use, limiting on wild prey in areas ofhigh prey densities were
themselves to habitat pockets that offer visual observed to have small territories (100 sq. km),
cover and shade, away from human disturbance. while wolves subsisting primarily by scavenging
Such habitat pockets are crucial for denning and and by predation on domestic livestockhad larger
as rendezvous sites (areas where pups are kept homeranges (250-300 sq. km) coveringthegrazing
afterthey leave the natal den), and could be in the grounds of several villages. Territoriality acts as
form of broken terrain, rocky outcrops, rivulets, a spacing mechanism to limit the number ofpacks
and grass or scrub patches. In VelavadarNational in an area. Prey density and prey size regulate
Park and the rest ofthe Bhal area, wolves preferred pack size and territory size. These two factors act
to use moderately dense Prosopis chilensis in synchrony to socially regulate wolf density in
patches (Jhala 1991; Jethva and Jhala, in prep.), an area.
while in Nanaj area wolves used forestry
plantations (Kumarand Rahmani 1995); in Banner Social Organisation
and Pali districts of Rajasthan ‘Aorans’ (sacred
groves) were crucial wolfhabitats. In Ojhar-Nasik In canids, sociality increases with body size
and near Jodhpur, airforce and military bases (Moehlman 1990, 1992). Wolves beingthe largest
provide refugia for denning and rendezvous sites of the canids, have a highly developed social
amongst a semi-urban landscape. In the Bhal area, system. Wolf society is organised around the
where wolves were studied through telemetry for breeding or alpha pair, the alpha pair and its
over eight years (Jhala 2001), core areas ofwolf offspring living together as a pack. Thus, a pack
territories where dens and rendezvous sites were is normally a family unit that may have been
located were between 5-10 sq. km and were established by related or unrelated wolves. The
characterised by low human disturbance, good pack stakes out and defends a resource territory
cover and presence offresh drinkingwater(Jethva from otherwolfpacks. Territorial defence is done
2002; Jethva and Jhala, in prep.). by scent marking, howling and by actual strife
Wolves living in the western dry zone of between neighbouring packs (Mech 1970). As
India were considered to be nomadic, primarily juvenile wolves mature, they either tend to
following pastoralists during their annual disperse from their natal pack or stay back as
migrations. However, telemetry studies in the Bhal helpers to their parents. Indian wolf pups may
and Kutch area ofGujarat, and in Ojhar in Nasik disperse as early as 7-8 months of age (Fig. 2).
district of Maharashtra, have shown that Indian Dispersers wander in search ofmates and available
wolves are territorial like all the other subspecies habitatto establish their own territories and packs,
of wolves. It seems likely that under certain while helpers bide their time till they become
ecological conditions territorial boundaries are breeders themselves by replacing and/or
more fluid. Territories of Indian wolves can be displacing their parent (Packard and Mech 1980).
quite large, with profound seasonal use of only Telemetry data has shown packs to be quite
some areas within them, giving an appearance of tolerant to intruders. Single non-pack members
“nomadism” to a casual observer. The area were observed in close proximityto pack members
covered by a wolf that predated on children, in and even shared kills on some occasions. Two
eastern Uttar Pradesh in 1996, was over packs were observed to intermingle without any
296 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003
ECOLOGY AND CONSERVATION OF THE INDIAN WOLF
Jhala, Yadvendradev: Canis lupus pallipes Plate
1
Fig. 1: The Indian wolfrepresents an ancient lineage of Canis lupus that is unique to peninsular India and
parts of Pakistan. Considered to be endangered, its numbers are believed to be between 2,000 to 3,000.
JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003 297
)
ECOLOGY AND CONSERVATION OF THE INDIAN WOLF
—
12 _J 1 I 1-
Whelping Rendezvous sites
Dispersal
Join the Pack
Natal Den
Mating
N
GO
JC
o
CO
0
.
4 -
Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May. Jun. Jul.
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Fig. 2: Major annual events and the average monthly pack size (with standard deviation)
ofthree wolfpacks in the Bhal region ofGujarat
strife on two occasions and at one time shared limited to the reserves. Most of the studies on
the same core area for a rendezvous site. This food habits reported here (Table 1) have been
may be an adaptation to a different set of done in protected areas and, therefore,
ecological conditions, like living offclumped food overestimate the contribution ofwild prey to the
resources (MacDonald 1979, Pisapio and wolfs diet. It would be safe to conclude that the
Theberge 2000) in the form ofdomestic livestock majority ofwolves in India subsist on small size
carcasses, or having limited access to undisturbed livestock, primarily goats and sheep (Shahi 1982,
habitats in a sea ofhuman modified landscapes. Jhala and Giles 1991).
Reduction in body size reduces food
Food Habits, Predation and Consumption demands and permits Indian wolves to subsist
even on small prey like hare {Lepus nigricollis
Wolves are the top predators of and rodents. Besides these, wolves also eat
blackbuck {Antilope cervicapra) and chinkara locusts, other insects, reptiles, birds, and
(Gazella bennettii), both medium-sized to small vegetable matter, like the pods of Prosopis
ungulates, in much ofthe arid and semi-arid areas chilensis and fruits ofZizyphns spp. (Sharma 978;
1
ofIndia (Jhala 1991, 1993, Sharma 1978). Food Jhala 1 993; Jethva and Jhala, in press). In Kutch,
habits of wolves have been studied from seven wolves were also recorded to prey on donkeys
different regions in India (Table 1). The majority and camel calves. Wolves ofVelavadar National
of the wolf population in India occurs outside Park subsisted almost exclusively on wild prey,
wildlife reserves, while wild ungulates are mostly even though domestic livestock were present in
JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003 299
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300 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003
ECOLOGY AND CONSERVATION OF THE INDIAN WOLF
the area (Jhala 1993). Consumption by wolves unsuccessful. Long chases of over 1 km were
preying on blackbuck was estimated at 4.62 observed when wolves hunted blackbuck calves.
(SE 0.11) kg/wolf/kill. Wolves killed blackbuck at Large prey is usually eviscerated and dies due to
an average interval of3.5 (SE 0.5)days(Jhala 1993). shock and blood loss. Wolves sometimes chase
Daily consumption by wolves estimated by three prey towards other pack members waiting in
independent studies was 1.33 (Jhala 1993), 1.008 ambush. The technique ofhunting blackbuck and
(Kumar2000), and 1 .80 kg/wolf(Jethvaand Jhala, chinkara by a lone wolf is by stalking, followed
in press). Jethva (2002) continuously monitored by a quick rush. Ifthe prey is caught, it is normally
three radio-collared wolf packs in the Bhal for killed by a bite on the throat or nape. The prey
periods ranging from 148 to 342 hours (total 1994 usually dies due to asphyxia. The majority ofkills
hours) and quantified feeding and intake rates of were made at night, however, during winter and
these packs. Wolfpredation was biased towards monsoon, when the days were not very hot,
adultmale blackbuckwhich contributed about70% wolves were seen hunting during daylight hours.
to the total biomass consumption of these wolf Wolfpredation is targeted towards ungulates that
packs (Jethva and Jhala, in press). Predation on are old, very young or in poor condition.
domestic livestock (cattle calves) was limited to
8%, while scavenging off cattle carcasses Breeding
contributed 14%tothe biomass consumed (Jethva
and Jhala, in press). Wolfpredation was a major C.l. pallipes is the only subspecies of
limiting factor responsible for shaping the age wolfthat breeds in winter. Births throughout the
and sex structure ofthe blackbuck population in wolf’s range seem to be well synchronised. Births
the Bhal (Jhala et al. 2001). in Gujarat (n=22 litters), Uttar Pradesh (n=1), and
Largerpacks (6 to 12 wolves)were recorded Maharashtra (n=4) occurred between December
in areas where wild ungulate prey was abundant, 15 and January 15. Mating occured sometime
e.g. in Velavadar National Park, Gudda Bishnoi, during OctobertoNovember(Fig. 2), between the
Nannaj, Dwarka and Rollapadu. In areas where dominantmale and female ofthepack. The mating
domestic livestock formed the major component pair is locked in a copulatory tie that lasts for 20-
ofthe diet, packsize ranged from 1 to4 individuals 30 minutes (Sheldon 1992). The gestation lasts
(Jhala and Giles 1991). Large numbers are more for62-63 days (Mech 970). The breeding (alpha)
1
likely to be detected by vigilant pastoralists and female begins excavating dens about a month to
their dogs, and may prove to be a disadvantage 15 days prior to whelping. It takes 2-10 days to
while hunting domestic prey. Moreover, most dig a complete den, and one or more dens are
large domestic livestockkills are underutilised due simultaneously excavated. The alpha male
to human disturbance to feeding wolves and a sometimes assists in the excavation butthe female
major portion ofthe kill is lost to scavengers like does most ofthe digging. Dens are excavated in
dogs andvultures. Utilisation ofkills in wilderness dry river embankments. Sometimes, fox and
areas by wolves is high due to lack of human porcupine holes are enlarged. Hollow trunks of
disturbance. Salvadora oleoides stacks of harvested cotton
,
Wolves are capable of hunting prey much stems, and rock crevices are also used (Jhala 1991,
largerthan themselves. This is primarily achieved Kumar2000, Sharma 1978). There seemsto be site
by hunting in packs. Wolves when hunting in fidelity for denning. Four study packs in the Bhal
packs use different “strategies” like stalking and region and two packs in Kutch excavated dens in
rushing or chasing. Most chases do not last more the same area (1 km radius) for 4-7 consecutive
than 0.5 km and several of the chases are years, even after the alpha females ofthree packs
JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003 301
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ECOLOGY AND CONSERVATION OF THE INDIAN WOLF
were replaced. A wolfden in the Gudda Bishnoi summer (March to early June), the rendezvous
area near Jodhpur excavated in the hollow trunk sites in and around a stream bed were intensively
ofa Salvadora tree, showed evidence ofuse over used. The pups would spend the hot hours ofthe
several years. Wolf dens that we observed had day in cool depressions dug out in the stream
oblong entrances with an average diameter of bank. The depressions were 25-30 cm deep and
48 cm (n=8, SE 14.5). The tunnels were about3 m 25-50 cm wide. These dug out depressions were
long with a single chamber at the end measuring also used by adult wolves for lying up. The pups
m m
about 1 . 1 by 0.7 m, having an average height of were restricted to an area of250 radius at their
m m
0.25 andwasatadepth of0.6 fromthe surface. first rendezvous site. Wolftracks converged from
The mother confines herselfto the vicinity all directions alongwell used trails. The area smelt
ofthe den 5-7 days prior to birth. In wild wolves strongly ofwolf odour and urine. Pup and adult
studied in the Bhal, Kutch, andNasik, mean litter wolf scats accumulated in and around the area.
size was 4.8 (n=28 litters) and ranged between 4 Kill remains like bones and hides were often strewn
to 6 pups. Pups are bom blind, develop blurred around. The description of rendezvous sites of
vision by the age of 15 days and by 1 8-20 days, timberwolves inNorth America(Joslin 1967, Mech
occasionally emerge from the den to play and 1970) matches extremely well withthe rendezvous
nurse. At the age of about 30-40 days, the pups sites of Indian wolves. Wolf pups were taken to
are normally moved to another den. They are feed on blackbuck kills as far as 3 km from the
moved between 4-5 rendezvous sites between the rendezvous site, by the age of 2.5 months in
age of40 days to theirjoining the pack at the age VelavadarNational Park.
of5-6 months.
Canidmilkis quite dilute (Oftedal 1984). The Conflict with Humans
lactating female needs to drink a lot ofwater so as
to produce milk. Denning sites and rendezvous Since the majority ofthe wolfpopulations
sites are so selected that fresh water is always in India live outside wildlife reserves, in human
availablewithin a radius of2 km. Wateravailability, dominated landscapes, they subsist primarily on
remoteness from human disturbance, visual cover livestock. A major occupation of the people in
and shade, seem to be the critical parameters for much of the wolfs range consists of livestock
selecting rendezvous and denning sites. At these rearing. Large herds of cattle, sheep, and goats
rendezvous sites, pups may continue to use dens graze the semi-arid landscape. Most of these
that are excavated by adult wolves and by the livestock are malnourished and die ofdisease and
pups themselves up to the age of5-6 months (till starvation. Since humans rarely consume cattle
June/July). This behaviour has not been reported in India, much ofthese carcasses are available for
in otherwolfsubspecies and may be an adaptation scavenging by dogs, vultures, jackals, hyenas,
in the Indian wolfto escape high ambient summer and wolves. Besides scavenging, the wolf also
temperatures. Afterthe pups leftthe natal den, they predates on livestock like goats, sheep, and cattle
were rarely observed using a single den at calves. Wolf predation severely affects the
rendezvous sitesbutwere distributed intwo ormore economy ofthe pastoral communities that barely
dens. Thisbehaviourwasprominentin Kutch, where manage to eke out a living from the highly
persecution of pups by herdsmen was severe. overgrazed and degraded landscape ofsemi-arid
Rarely were all the pups killed when shepherds India. The pastoral community invests
smoked wolfdens other than the natal den. significantly in measures to protect their stock
Nine rendezvous sites were used by the from wolf predation. These measures include
Velavadar pack over2 breeding years. Duringthe night vigils, maintaining guard dogs, building
302 JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 100(2&3), AUG.-DEC. 2003
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