Problem-Elephant Translocation: Translocating the Problem and the Elephant?
Citation: Fernando P, Leimgruber P, Prasad T, Pastorini J (
Problem-Elephant Translocation: Translocating the Problem and the Elephant?
Prithiviraj Fernando 0 1
Peter Leimgruber 0 1
Tharaka Prasad 0 1
Jennifer Pastorini 0 1
Matt Hayward, Australian Wildlife Conservancy, Australia
0 Funding: This work was funded by the U.S. Fish and Wildlife Service, Asian Elephant Conservation Fund, Alexander Abraham Foundation, Sidney S. Byers Charitable Trust, Eco Health Alliance, Friends of the National Zoo (FONZ) , Circus Knie , Smithsonian Women's Committee, Vontobel Stiftung, and the PAM-WCP Project of the DWC. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript
1 1 Centre for Conservation and Research , Rajagiriya , Sri Lanka , 2 Smithsonian Conservation Biology Institute, Front Royal, Virginia, United States of America, 3 Department of Wildlife Conservation , Battaramulla , Sri Lanka , 4 Anthropologisches Institut, Universita t Zu rich , Zu rich , Switzerland
Human-elephant conflict (HEC) threatens the survival of endangered Asian elephants (Elephas maximus). Translocating ''problem-elephants'' is an important HEC mitigation and elephant conservation strategy across elephant range, with hundreds translocated annually. In the first comprehensive assessment of elephant translocation, we monitored 16 translocations in Sri Lanka with GPS collars. All translocated elephants were released into national parks. Two were killed within the parks where they were released, while all the others left those parks. Translocated elephants showed variable responses: ''homers'' returned to the capture site, ''wanderers'' ranged widely, and ''settlers'' established home ranges in new areas soon after release. Translocation caused wider propagation and intensification of HEC, and increased elephant mortality. We conclude that translocation defeats both HEC mitigation and elephant conservation goals.
Translocation is defined as the deliberate and mediated
movement of wild individuals or populations from one part of
their range to another . It is a commonly used tool in
conservation, for establishing, re-establishing and augmenting
populations of managed species . It is also employed in
managing problem-wildlife, although a number of studies have
questioned its use in this context . Due to ethical concerns
and mounting objections to lethal control [3,7], translocation is
increasingly viewed as a panacea for all wildlife problems . The
main objective of problem-animal translocation is eliminating
problems caused by wildlife  and secondly, saving the animals
responsible. In conservation use, translocated animals are usually
released in empty habitats . In problem-animal translocation
they are more likely released in areas fully occupied by conspecifics
. Translocated animals may be first acclimatized at the release
site (soft-release) or released immediately (hard-release), the latter
being more common in problem-animal translocation [2,3]. Many
thousands of problem-animals are translocated annually [3,4].
Mostly applied to nuisance or dangerous animals it is
taxonomically biased towards mammals. Species so translocated
include squirrels , raccoons , deer, bear, rodents , wolves
, foxes, wild cats , cougars , leopards , tigers ,
elephants [16,17], geese , eagles , Gila monsters ,
snakes  and crocodilians .
The Asian elephant (Elephas maximus) is an endangered species
on the IUCN Red List and is listed in CITES Schedule I [23,24].
The global population estimate for Asian elephants is 35,000
50,000 [24,25], one tenth that of African elephants (Loxodonta
africana and L. cyclotis). Asian elephants are now extinct in 78% of
their historic range . Currently they are limited to a number of
fragmented and isolated populations in 13 south and south-east
Asian states [24,25,27]. With only 16% of their remaining range
protected , most Asian elephants are compelled to share space
with humans, leading to frequent conflict. For example, over 70%
of about 6,000 elephants in Sri Lanka live outside protected areas,
where annually human-elephant conflict (HEC) claims the lives of
over 70 humans and 200 elephants . Today, HEC is a major
conservation, socio-economic and political issue across Asian
elephant range .
Elephant social organization is sexually dimorphic with
groupliving adult females and young, and mainly solitary adult males
. Males display a higher propensity for crop raiding,
accessing superior resources to gain in size hence reproductive
advantage, in a high-risk high-gain strategy . Some males
raid crops, break into houses for stored grain, and react
aggressively to confrontation, causing human injury and death.
Considered problem-elephants, such individuals are responsible
for the majority of HEC incidents .
While lethal control is preferred in some parts of Africa ,
translocation remains one of the main elephant management tools
and hundreds of elephants are translocated annually
[17,28,35,36]. Translocating problem-elephants aims to mitigate
HEC by removing them from human proximity. It also attempts
to further elephant conservation, assuming higher mortality if
problem-elephants remain in their original home ranges. The
modus operandi for translocating problem-elephants is capture by
drug immobilization, transport by truck and release in a protected
area. In Sri Lanka and India, elephants so translocated are
exclusively males, while in Malaysia, Indonesia and some African
countries it may involve both sexes [35,37].
Elephants have comparatively large home ranges and can cover
long distances quickly . Often they also inhabit poor
visibility habitat and actively avoid humans [39,41]. Consequently,
monitoring individual elephants without radio-telemetry is
ineffective and with VHF transmitters is at best difficult. Only a few
translocations have been previously monitored with
radio-telemetry, consisting of one elephant in India , 11 in Kenya 
and six in South Africa  that were tracked with VHF, two
tracked with satellite-PTT transmitters in Malaysia  and one
with GPS in Kenya . Anecdotal accounts  and the few
monitoring studies, suggest that some translocated elephants
return while others settle in release areas.
In this paper, we report on the first comprehensive assessment
of problem-elephant translocation. Using remote-download GPS
collars, we monitored 12 males translocated 16 times and 12 males
resident in their normal home ranges. Here we compare and
contrast the behavior and HEC involvement of translocated and
resident elephants, and discuss the relevance of findings for
All elephants in our study were adults and were classified as
mature-adults or young-adults, corresponding approximately to
above and below 30 years of age. Individuals displaying
a combination of the following characters were identified as
mature adults: shoulder height over 270 cm; well developed
secondary sexual characters such as wide trunk base, prominent
nasal protuberance, deep temporal depression and large penis/
penile bulge; characters indicating active musth such as temporal
gland discharge and urine dribbling; and age related characters
such as completely folded top edge of ear and heavy
All 12 translocated elephants were identified as
problemelephants by the Department of Wildlife Conservation Sri Lanka
(DWC) based on HEC incidents and information from villagers.
The resident males consisted of two (Kandula and Kavan) that did
not cause HEC and 10 problem-elephants. Reported incidents of
crop raiding, house breaking or human injury and death, and
entering areas of human habitation by monitored elephants were
taken to indicate causation of HEC.
Collars and Collaring
Translocated elephants were fitted with radio-collars at the time
of capture. The collars consisted of a GPS unit, VHF transmitter
beacon, satellite or GSM transmitter for data download (Table 1)
and batteries packaged into one integrated unit. Sky orientation of
the functional unit for satellite detection was achieved by
a counterweight. Collars that became non-functional were not
removed as it was determined that the risk to the elephant and
collaring team in tranquilization was not acceptable for the
purpose of collar removal. Collar belting degraded and broke off
within a period of 24 years (unpublished data).
All translocated elephants were captured outside protected areas
and released inside national parks (Fig. 1). All release locations
were within current elephant range and had ample water and
fodder. Two males (Ravana and Tzu Chi) were translocated twice
and one (Homey) was translocated three times. Translocated
elephants were hard-released and the time from capture to
release was 13 days.
Translocated elephants were tracked using the VHF beacon on
the collar and observed opportunistically.
The study was mandated by the DWC and conducted
collaboratively by the DWC and the Centre for Conservation and Research
(CCR). Under the Fauna and Flora Protection Ordinance of Sri
Lanka, the DWC is legislated as the government institution that is the
sole authority on wildlife management in Sri Lanka and there is no
requirement or procedure to obtain separate approval for activities
conducted by the DWC. Elephants were captured and translocated as
part of the routine activities of the DWC for mitigating HEC and
conserving elephants. Collaring of resident elephants was done as part
of another on-going study by the DWC and CCR to obtain baseline
information to better elephant conservation and HEC mitigation.
Tranquilizing elephants for collaring was done by a DWC team of 15
20 personnel led by two DWC veterinarians according to guidelines
set out by the DWC.
All efforts were made to prevent and minimize suffering of
animals concerned and to ensure the safety of animals and
personnel involved in research activities. Radio darts were used to
maximize the safety of darted animals by reducing search time and
minimizing possibilities of complications of tranquilization under
field conditions. Throughout the tranquilized period, a
veterinarian monitored the status of the elephant to prevent any
complications. Tranquilized elephants were given a health check
and were treated by wound cleaning and injection of antibiotics as
indicated (eg. gunshot wounds, abscesses).
Collars were programmed to collect GPS locations every 4 or 8
hours and transmit the data every 8, 24 or 48 hours (Table 1). In
Telonics and Vectronic collars data were also stored on-board and
were directly downloadable if the collar was recovered.
Data received from collars were processed with the
corresponding manufacturers software. GPS locations obtained were
tabulated in Excel, exported into ArcMap (EsriArcGIS) version
9.2 or Quantum GIS version 1.7 (QGIS) and plotted on satellite
imagery or 1:50,000 topographic sheets. Home ranges and use
areas were calculated as 100% Minimum Convex Polygons in
QGIS (single minimum convex hull function).
To simplify directional analysis we re-projected the movement
data after release so that all elephant release sites were at the
coordinate origin (0,0) and capture sites were oriented at 180u (to the
left) from the release location. To assess movement orientation after
release, we calculated the spatial mean of all GPS positions acquired
during the first 10 days of tracking and computed the movement angle
between the release site and this spatial mean. Angles ,90u and
.270u (in right hemisphere) were taken to represent movement
orienting away from the capture site, and all others (90u270u)
towards the capture site. To test whether elephants more often
oriented towards the capture site than expected by chance alone, we
used a binomial test and calculated confidence intervals. All data
manipulations and statistical tests for assessing movement direction
were performed using R statistical software (R Development Core
Team 2011, ,www.R-project.org.).
Translocated and resident individuals were tracked for periods
of 262.56279.4 (range 171,009) and 314.86298.6 (range 34
1,022) days respectively, giving total periods of 4,200 days of
Africa Wildlife Tracking
Africa Wildlife Tracking
Africa Wildlife Tracking
translocated and 3,777 days of resident elephant tracking (Tables 1
and 2). The mean translocation distance was 134.8672.7 (range
37.4289.1) km (Table 1). All translocated elephants were released
inside national parks. Two elephants were shot dead within the
parks where they were released (Tzu Chi and Ravana) and all the
others left those parks (time to exit: 33.3669.3, range 1263 days,
Over the first 10 days post-release, in 11 of 16 translocations,
elephants oriented towards the capture site (Fig. 2). No aggression
was observed between translocated elephants and resident park
elephants, and no injuries caused by other elephants were
observed on the five translocated males that died (Table 3). All
areas where translocated elephants settled had resident elephants.
Two elephants (Galli and Ekes) were observed to associate with
resident bulls post-release.
Individual Variation in Response
We classified the translocated elephants as homers, wanderers
and settlers based on response.
In five translocations homers Chandi, Homey and Kabaraya
returned to the capture site thrice and showed movements
consistent with successful homing twice (Fig. 3B). Chandi
translocated 93.4 km, returned in 29 days. Homey after his first
and second translocations over 48.2 and 46.2 km homed back in 5
and 41 days respectively. Homey on his third translocation of
161.7 km showed homing movement for 62.0 km in 4 days but
entered a town causing conflict. Chased back to the release
location, he settled at the perimeter of the park, raided
surrounding villages, was shot repeatedly and died 15 months
after from gunshot injuries. Kabaraya translocated 116.8 km, after
an initial period in the release area, showed homing movement.
However, the collar stopped functioning at 92 days, 81.4 km from
the capture point. Homey and Kabaraya showed well directed
homing movements while Chandi took a more circuitous route
back (Fig. 3B).
Wanderers Wasaba, Siyak, Brigadier, Ravana, and Barbar
showed misdirected long distance movements (Fig. 3A). Wasaba
and Siyak travelled 127.0 and 43.0 km respectively till obstructed
by the sea, returned and settled proximal to the release area.
Brigadier showed directional movement for 95.9 km. When
confronted by the sea he swam out, was providentially spotted
5 km offshore by the Sri Lanka Navy, noosed underwater by scuba
divers and brought back to shore. He then settled in a new area,
continued to cause conflict and died from falling into a well 6
months after. Ravana entered a major town, created conflict and
was shot in the leg. He then took refuge in a forest patch where he
remained for 3 months. He was recaptured and translocated to
another national park, raided cultivations within the park and was
found shot dead 8 months later. Babar traveled 95.9 km in 19 days
before exiting the park where he was released and the collar came
off 16 days later.
Settlers Galli, Ekes, Tzu Chi and Nalagiri settled proximal to
the park where they were released, without any directional long
distance movements away from the release site (Fig. 3C). Galli
shifted his new home range twice after 6 and then 3 months.
Gallis first home range was in the park (176 km2) and the others
(115 and 73 km2) outside. Between his first and second home
ranges, for 2 months Galli used only an 8 km2 area along
a perimeter electric fence of the park. Ekes new home range was
162 km2, largely adjacent to the park where he was released. He
ventured back into the park 16 times, spending 35 days within, in
the 1,009 days period tracked. He raided regularly, making
nocturnal forays into villages and taking cover in forested habitat
during day. Tzu Chi was translocated 37.4 km northeast from his
capture site. After 29 days he left the park and two weeks later
settled in an area 8.1 km south of the release point, where he
continued to cause conflict. He was re-captured eight months later
and translocated 289.1 km northwest from the original capture
site. Upon release he moved south and was found shot dead 55
days later, 18.3 km from the release point and 355 m from the
park boundary. Nalagiri established a new home range partly
outside the park where he was released, regularly raided nearby
villages and was found shot dead 5 months after release.
Relation to Conflict
On six instances translocated elephants confronted electric
fences on park boundaries. Another two elephants were released
within a holding ground with a high specification perimeter
electric fence plus elephant-trench (Table 3). None of them were
contained by such barriers, except Ravana who was killed within
the park. Translocated elephants had average use-areas of
1,09061,276 km2, (range 604,380 km2; Table 1). The 12
resident males had significantly smaller home ranges of
2826222 km2 (range: 63643 km2; Mann-Whitney U test,
P = 0.0488, Table 2). Homey had home ranges of 153 km2 and
132 km2 between his translocations and use-areas of 311, 570
and 435 km2 during them. Chandi had a use-area of 4,126 km2
during the translocation and a home range of 336 km2 after
Four of the 12 translocated elephants (Homey, Chandi,
Wasaba, Ravana) but none of the 12 resident males entered
major towns. The incursions created chaos with human injury and
death, damage to property including vehicles and killing of a
waterbuffalo. The 12 translocated elephants killed 5 people. No deaths
were caused by the 12 resident males, one of whom was shot dead
during the study period.
The majority of translocated elephants displayed post-release
movements oriented towards the capture site (Fig. 2). Homing
upon translocation has been observed in a range of species,
including bears [48,49], cougars , wolves , foxes , deer
, elephant seals , eagles , crocodiles , Gila
monsters , and newts . Home ranges and spatial
organization of individuals reflect resource use and strategies
adopted by individuals to maximize fitness [53,54]. Familiarity
with ones environment and neighbors is positively correlated with
individual fitness . Thus, the drive of translocated animals to
return, maybe due to the increased fitness accruing from
occupying a familiar home range. In long-lived and highly social
species such as elephants, selection on home range fidelity, hence
drive to home back is likely to be stronger.
Asian elephants have well defined home ranges with high
fidelity [39,56] and it is likely that translocated elephants left the
parks where they were released, in attempts to return home. All
six parks where elephants were released had abundant water, wild
fodder and female herds. Thus, it is unlikely that the decision to
leave was related to resource deficiency. Some translocated
elephants associated with resident park elephants and we saw no
evidence of agonistic encounters between translocated and
resident elephants. Therefore, consistent with non-territoriality of
elephants , translocated individuals are also unlikely to have
left the parks due to antagonism by resident elephants.
Individual elephants responded variably to translocation by
homing back, wandering or settling, the type of response being
unrelated to translocation distance. In an assessment of elephant
re-introductions in South Africa, no factors including distance
explained translocation failure . We found 3 of 5
matureadults and none of 7 young-adults displayed successful homing
movements, suggesting a tendency of successful homing by older
individuals (Table 3). Many species show individually variable
responses to translocation with some returning to the capture site
and others settling at the release location [10,11]. In some species
the probability of returning home is inversely related to distance
translocated (wolves , bears , foxes , Gila monsters
) and in some, those that return are more likely to be adults
(cougars , wolves ). Sex bias with males more likely to
leave has been observed in cougars  and black bears .
Individual response to translocation may also be related to
environmental factors such as relative resource availability of
capture and settling/release locations, physiographic and
anthropogenic barriers; behavioral factors such as social status, and
covert aggression of conspecifics; and innate factors such as
physiological and psychological states of individuals. However,
such aspects are difficult to test empirically.
Extent of Ranging
Use-areas of translocated elephants were significantly larger
than home ranges of resident elephants. On the three instances
translocated elephants returned home, their use-areas between
release and return were greater than their post-return home
Table 3. Data summary for translocated elephants.
*Elephant broke through electric fence on a park boundary.
#Holding ground, which is a specially fenced off portion (25.5 km2) of the park.
ranges. Wider ranging upon translocation has been documented in
many species  including cougars , black bears , snakes
 and crocodiles . In addition to attempted homing,
animals released in occupied habitats may show increased ranging
due to competition with residents and exploration. Given the
apparent resource abundance and the absence of overt aggression
from conspecifics, the increased post-translocation ranging
observed in our study maybe primarily explained by attempted
homing and secondarily by exploration.
Relation to HEC
Practically all translocated elephants were involved in HEC
post-release. They ranged widely with Homers and wanderers
venturing outside normal elephant range, some even entering
highly populated cities. Thus, problem-elephant translocation
resulted in wider propagation of HEC.
Translocated elephants roamed in environments alien to them,
in ignorance of the lay of the land. This increased the likelihood of
unanticipated encounters and conflict with humans. The 12
resident males did not cause any human deaths. This finding is
consistent with the annual elephant induced human mortality rate
in Sri Lanka (including that by about 14 elephants translocated
annually) of 0.04 humans/adult male or 0.01 humans/elephant
. In contrast, human mortality caused by the 12 translocated
elephants monitored was an order of magnitude higher at 0.42
humans/elephant (Fishers exact test, p,0.0001). Therefore
problem-elephant translocation intensified HEC.
Most translocated elephants resumed raiding after release
(Table 3). Elephants in shared landscapes are preferential, rather
than obligate raiders . Therefore, raiders are likely to be
compulsive and continue to raid irrespective of changed
circumstances. Post-release assessments of behaviors characterizing
problem-animals have been few, but most have found lack of
reform [3,5]. A study of house-denning raccoons found the
majority to persist with the behavior after removal . Of four
tigers translocated because of livestock predation, two immediately
moved to human dominated habitats . Three of four
translocated stock-raiding leopards resumed raiding . A survey
of leopard translocations found a positive correlation between
translocations and conflict . Translocation was found to be
largely unsuccessful at keeping problem wolves out of livestock
production areas . Our findings are consistent with these
observations and suggest that successful problem-animal
translocation most likely translocates not only the animal but also the
Galli and Wasaba did not raid post-release. Translocation is the
culmination of a train of events, usually instigated by a major
incident like human death or house breaking by elephants.
Capture occurs days to weeks after the incident. Elephants in Sri
Lanka have home ranges of 41643 km2 (Table 2) .
Consequently, by the time of capture the elephant responsible
may no longer be in the vicinity. Additionally, most HEC incidents
occur at night and even if witnessed, the perpetrator cannot be
identified with certainty. Thus, Galli and Wasaba may not have
been problem-elephants but victims of mistaken-identity.
Reviews of avian and mammal translocations have generally
found a greater number of successful translocations with
hardrelease [2,61]. While IUCN guidelines for African elephant
translocation recommend soft-release , some re-introduced
African elephants so translocated still left the release area .
Effect of release type has mostly been assessed in re-introductions,
where settling in the release area denotes success. All the elephants
in our study were hard-released and some settled near release
areas but reverted to raiding. It is unlikely that release type would
have much bearing on the outcome in problem-animal
translocation, where eliminating the problem is the primary objective
 and its translocation signifies failure.
Soft-release is also advocated in African elephant translocation
for educating elephants to respect electric fences during
acclimatization . All elephants who encountered electric
fences in our study broke through them. In Gallis case, breakout
occurred only after months of fence patrolling, suggesting
sustained effort to overcome fences rather than a lack of respect
for them. Therefore, the effect of release type on fence breaking is
debatable. However, the adequacy of the fences that translocated
elephants were confronted with could be a confounding factor.
Survival of Translocated Elephants
The 12 resident males tracked had a death rate of 0.10 per
tracked-elephant-year. This is consistent with the annual mortality
rate of adult male elephants in Sri Lanka of around 78% . All
12 translocated elephants survived to adulthood in their original
home ranges. However, five of them died within 8 months of
release (Table 3), amounting to 42% mortality or 0.44 deaths per
tracked-elephant-year. Additionally, translocation carries a
mortality rate of approximately 6% during capture and transport .
Therefore, although translocation aimed to safeguard
problemelephants, in reality it greatly reduced their survival.
Increased mortality of translocated individuals has been
observed in raccoons , cougars , wolves , elephants
 and snakes . Similar survivability to resident populations
has been reported in muskoxen . Some studies found increased
mortality in black bears  while others did not [49,57]. Higher
mortality of translocated animals may be related to their wider
ranging in unfamiliar environments. Additionally,
problemanimals are individuals with a greater predilection for conflict
with people and the probability of encounters hence conflict is
increased by translocation. Therefore, as seen in our study,
mortality is likely to be much higher in translocated
Translocation caused elephants to behave abnormally,
increased their mortality, and presumably subjected individuals to
extreme stress. Elephants are a highly social species with a network
of relationships even amongst males . Translocation disrupts
such relationships at both capture and release locations. Elephants
are also an intelligent and long-lived species. Consequently,
profound negative experiences may have extensive and long-term
psycho-physiological effects on their brains and behavior .
Therefore, from an elephant welfare point of view, translocation is
not an acceptable management tool.
We conclude that problem-elephant translocation causes
intensification and broader propagation of HEC and increased
elephant mortality, hence defeats both HEC mitigation and
elephant conservation goals. The driver of translocation is public
and political pressure. Capturing and translocating an elephant
from the vicinity of major HEC incidents may defuse tension
hence be of relevance in particular contexts. However we found
that even if the original problem is solved by translocation, the
same or more likely worse is created at another location.
Based on our results we advocate phasing out problem-elephant
translocation, for which public awareness is key. In the interim,
translocations should only be undertaken with monitoring through
GPS-telemetry, and contingency plans to address unintended
outcomes. Problem-elephant translocation without either,
amounts to reckless disregard for the safety and welfare of people
and elephants. In the long term, attention needs to be shifted
towards preventing the genesis of problem-elephants. Such
a strategy requires eliminating elephant management and crop
protection methods that promote elephant aggression and increase
HEC, and implementing land-use plans that minimize crop
We thank the Department of Wildlife Conservation (DWC) Sri Lanka for
collaboration and H.K. Janaka, S. Ekanayaka, H.G. Nishantha, M.
Gunawardena and DWC officers for helping with the fieldwork.
Conceived and designed the experiments: PF PL. Performed the
experiments: PF JP TP. Analyzed the data: JP PL. Wrote the paper: PF
JP PL. Led the collaring operation of elephants: TP.
1. IUCN ( 1998 ) Guidelines for Re-introductions . Prepared by the IUCN/SSC Reintroduction Specialist Group . Gland, Switzerland and Cambridge, UK: IUCN.
2. Wolf CM , Griffith B , Reed C , Temple SA ( 1996 ) Avian and Mammalian Translocations : Update and Reanalysis of 1987 Survey Data . Conserv Biol 10 : 1142 - 1154 .
3. Linnell JDC , Aanes R , Swenson JE , Odden J , Smith ME ( 1997 ) Translocation of carnivores as a method for managing problem animals: a review . Biodiv Conserv 6 : 1245 - 1257 .
4. Craven S , Barnes T , Kania G ( 1998 ) Toward a professional position on the translocation of problem wildlife . Wildl Soc Bull 26 : 171 - 177 .
5. Loveridge AJ , Wang SW , Frank LG , Seidensticker J ( 2010 ) People and wild felids: conservation of cats and management of conflicts . In: Macdonald DW, Loveridge AJ , editors. The Biology and Conservation of Wild Felids . New York : Oxford University Press . 161 - 195 .
6. Massei G , Quy RJ , Gurney J , Cowan DP ( 2010 ) Can translocations be used to mitigate human-wildlife conflicts? Wildl Res 37 : 428 - 439 .
7. Treves A , Karanth KU ( 2003 ) Human-carnivore conflict and perspectives on carnivore management worldwide . Conserv Biol 17 : 1491 - 1499 .
8. Loredo-Prendeville I , Vuren DV , Kuenzi AJ , Morrison ML ( 1994 ) California ground squirrels at Concord naval weapons station: alternatives for control and the ecological consequences . In: Halverson WS, Crabb AC, editors. Proceedings of the Sixteenth Vertebrate Pest Conference . Davis, CA: University of California.
9. O'Donnell MA , DeNicola AJ ( 2006 ) Den site selection of lactating female raccoons following removal and exclusion from suburban residences . Wildl Soc Bull 34 : 366 - 370 .
10. Rogers LL ( 1988 ) Homing tendencies of large mammals: a review . In: Nielsen L, Brown R , editors. Translocation of Wild Animals. Milwaukee, WI: Wisconsin Humane Society . 76 - 92 .
11. Bradley EH , Pletscher DH , Bangs EE , Kunkel KE , Smith DW , et al. ( 2005 ) Evaluating wolf translocation as a nonlethal method to reduce livestock conflicts in the northwestern United States . Conserv Biol 19 : 1498 - 1508 .
12. Lenain DM , Warrington S ( 2001 ) Is translocation an effective tool to remove predatory foxes from a desert protected area ? J Arid Environ 48 : 205 - 209 .
13. Ruth TK , Logan KA , Sweanor LL , Homocker MG , Temple LJ ( 1998 ) Evaluating cougar translocations in New Mexico . J Wildl Manag 62 : 1264 - 1275 .
14. Athreya V , Odden M , Linnell JDC , Karanth KU ( 2010 ) Translocation as a tool for mitigating conflict with leopards in human-dominated landscapes of India . Conserv Biol 25 : 133 - 141 .
15. Goodrich JM , Miquelle DG ( 2005 ) Translocation of problem Amur tigers Panthera tigris altaica to alleviate tiger-human conflicts . Oryx 39 : 1 - 4 .
16. Fernando P ( 1997 ) Keeping jumbo afloat . Sri Lanka Nature 1 : 4 - 12 .
17. Dublin HT , Niskanen LS (editors) ( 2003 ) IUCN/SSC AfESG Guidelines for the in situ Translocation of the African Elephant for Conservation Purposes . Gland, Switzerland: IUCN.
18. Holevinski RB , Malecki RA , Curtis PD ( 2006 ) Can hunting of translocated nuisance Canada geese reduce local conflicts ? Wildlife Soc Bull 34 : 845 - 849 .
19. Boshoff AF , Vernon CJ ( 1988 ) The translocation and homing ability of problem eagles . S Afr J Wildlife Res 18 : 38 - 40 .
20. Sullivan BK , Kwiatkowski MA , Schuett GW ( 2004 ) Translocation of urban Gila monsters: a problematic conservation tool . Biol Conserv 117 : 235 - 242 .
21. Butler H , Malone B , Clemann N ( 2005 ) The effects of translocation on the spatial ecology of tiger snakes (Notechis scutatus) in a suburban landscape . Wildlife Res 32 : 165 - 171 .
22. Read MA , Grigg GC , Irvin SR , Shanahan D , Franklin CE ( 2007 ) Satellite tracking reveals long distance coastal travel and homing by translocated estuarine crocodiles . Crocodylus porosus. PloS ONE 9 : 1 - 5 .
23. IUCN ( 2011 ) IUCN Red List of Threatened Species . Version 2011 .2. Available: http://www.iucnredlist.org. Accessed 2012 Nov 10 . (Current version: 2012 .2.).
24. Santiapillai C , Jackson P ( 1990 ) The Asian Elephant: An Action Plan for its Conservation . Gland, Switzerland: IUCN/SSC Asian Elephant Specialist Group.
25. Fernando P , Pastorini J ( 2011 ) Range-wide status of Asian elephants . Gajah 35 : 15 - 20 .
26. Fernando P , Leimgruber P ( 2011 ) Asian elephants and dry forests . In: McShea WJ , Davies SJ , Phumpakphan N , Pattanavibool A , editors. The Ecology and Conservation of Seasonally Dry Forests in Asia . Washington, DC: Smithsonian Institution Scholarly Press. 151 - 163 .
27. Leimgruber P , Gagnon JB , Wemmer C , Kelly DS , Songer MA , et al. ( 2003 ) Fragmentation of Asia's remaining wildlands: implications for Asian elephant conservation . Anim Conserv 6 : 347 - 359 .
28. Fernando P , Jayewardene J , Prasad T , Hendavitharana W , Pastorini J ( 2011 ) Current status of Asian elephants in Sri Lanka . Gajah 35 : 93 - 103 .
29. Moss CJ , Poole JH ( 1983 ) Relationships and social structure of African elephants . In: Hinde RA, editor. Primate Social Relations: An Integrated Approach . Oxford: Blackwell Scientific Publications. 315 - 325 .
30. Sukumar R ( 1989 ) The Asian Elephant: Ecology and Management . Cambridge : Cambridge University Press.
31. Fernando P , Lande R ( 2000 ) Molecular genetic and behavioral analyses of social organization in the Asian elephant . Behav Ecol Sociobiol 48 : 84 - 91 .
32. Sukumar R ( 1991 ) The management of large mammals in relation to male strategies and conflict with people . Biol Conserv 55 : 93 - 102 .
33. Fernando P ( 2011 ) Managing 'problem elephants' . Loris 25 ( 6 ): 32 - 36 .
34. Slotow R , Whyte I , Hofmeyr M , Kerley GHI , Conway T , et al. ( 2008 ) Lethal management of elephants . In: Scholes RJ, Mennell KG, editors. Elephant Management: A Scientific Assessment of South Africa . Johannesburg: Witwatersrand University Press.
35. Pinter-Wollman N ( 2009 ) Spatial behaviour in translocated African elephants (Loxodonta africana) in a novel environment: using behaviour to inform conservation . Behaviour 146 : 1171 - 1192 .
36. Saaban S , Othman NB , Yasak MNB , Burhanuddin MN , Zafir A , et al. ( 2011 ) Current status of Asian elephants in Peninsular Malaysia . Gajah 35 : 67 - 75 .
37. Stuewe M , Abdul JB , Nor BM , Wemmer CM ( 1998 ) Tracking the movements of translocated elephants in Malaysia using satellite telemetry . Oryx 32 : 68 - 74 .
38. Douglas-Hamilton I , Krink T , Vollrath F ( 2005 ) Movements and corridors of African elephants in relation to protected areas . Naturwissenschaften 92 : 158 - 163 .
39. Fernando P , Wickramanayake ED , Janaka HK , Jayasinghe LKA , Gunawardene M , et al. ( 2008 ) Ranging behavior of the Asian elephant in Sri Lanka . Mammal Biol 73 : 2 - 13 .
40. Roy M , Choudhury SP , Kamalakanth P , Dutta C , Kundu S , et al. ( 2010 ) Translocation of a wild elephant from southern West Bengal to northern West Bengal-An approach to reduce elephant-human conflict . Gajah 33 : 8 - 11 .
41. Lin L , Feng L , Pan W , Guo X , Zhao J , et al. ( 2008 ) Habitat selection and the change in distribution of Asian elephants in Mengyang Protected Area , Yunnan, China. Acta Theriol 53 : 365 - 374 .
42. Slotov R , Van Dyk G ( 2004 ) Ranging of older male elephants introduced to an existing small population without older males: Pilanesberg National Park . Koedoe 47 : 91 - 104 .
43. Banks J ( 1979 ) The translocation of the Deduru Oya herd . What was left of it . Loris 15 : 113 - 115 .
44. Lahiri-Choudhury DK ( 1993 ) Problems of wild elephant translocation . Oryx 27 : 53 - 55 .
45. Garai ME , Carr RD ( 2001 ) Unsuccessful introductions of adult elephant bulls to confined areas in South Africa . Pachyderm 31 : 52 - 57 .
46. Arivazhagan C , Sukumar R ( 2008 ) Constructing age structures of Asian elephant populations: A comparison of two field methods of age estimation . Gajah 29 : 11 - 16 .
47. Varma S , Baskaran N , Sukumar R ( 2012 ) Field Key for Elephant Population Estimation and Age and Sex Classification . Bangalore: Asian Nature Conservation Foundation , Innovation Centre , Indian Institute of Science.
48. Miller SD , Ballard WB ( 1982 ) Homing of transplanted Alaskan brown bears , J Wildl Manage 46 : 869 - 876 .
49. Rogers LL ( 1986 ) Effects of translocation distance on frequency of return by adult black bears . Wildl Soc Bull 14 : 76 - 80 .
50. Eberhardt LL , Pickens HC ( 1979 ) Homing tendencies in mule deer . Southwest Nat 24 : 705 - 706 .
51. Oliver GW , Morris PA , Thorson PH , Le Boeuf BJ ( 1998 ) Homing behavior of juvenile northern elephant seals . Marine Mammal Sci 14 : 245 - 256 .
52. Phillips JB , Adler K , Borland SC ( 1995 ) True navigation by an amphibian . Anim Behav 50 : 855 - 858 .
53. Burt WH ( 1943 ) Territoriality and home range concepts as applied to mammals . J Mammal 24 : 346 - 352 .
54. Sandell M ( 1989 ) The mating tactics and spacing patterns of solitary carnivores . In: Gittleman JL, editor. Carnivore Behaviour, Ecology and Evolution . Ithaca: Cornell University Press. 164 - 182 .
55. Shier DM , Swaisgood RR ( 2012 ) Fitness costs of neighborhood disruption in translocations of a solitary mammal . Conserv Biol 26 : 116 - 123 .
56. Baskaran N , Desai AA ( 1996 ) Ranging behavior of the Asian elephant (Elephas maximus) in the Nilgiri biosphere reserve , South India. Gajah 15 : 41 - 57 .
57. Benson JF , Chamberlain MJ ( 2007 ) Space use, survival, movements, and reproduction of reintroduced Louisiana black bears . J Wildl Manag 71 : 2393 - 2403 .
58. Ekanayaka SKK , Campos-Arceiz A , Rupasinghe M , Pastorini J , Fernando P ( 2011 ) Patterns of crop raiding by Asian elephants in a human-dominated landscape in Southeastern Sri Lanka . Gajah 34 : 20 - 25 .
59. Weilenmann M , Gusset M , Mills DR , Gabanapelo T , Schiess-Meier M ( 2010 ) Is translocation of stock-raiding leopards into a protected area with resident conspecifics an effective management tool? Wildl Res 37 : 702 - 707 .
60. Fritts SH , Paul WJ , Mech LD ( 1984 ) Movements of translocated wolves in Minnesota . J Wildl Manag 48 : 709 - 721 .
61. Short J ( 2009 ) Australian Animal Welfare Strategy-The Characteristics and Success of Vertebrate Translocations within Australia . Canberra, Australia: Australian Government Department of Agriculture, Fisheries and Forestry.
62. Allen TJ ( 1986 ) Evaluation of movements, harvest rate, vulnerability and survival of translocated raccoons in southern West Virginia . Trans Northeast Sect Wildlife Soc 43 : 64 .
63. Le Henaff D , Crete M ( 1989 ) Introduction of muskoxen in northern Quebec: the demographic explosion of a colonizing herbivore . Can J Zool 67 : 1102 - 1105 .
64. Chiyo PI , Archie EA , Hollister-Smith JA , Lee PC , Moss CJ , Alberts SC ( 2011 ) Association patterns of African elephants in all-male groups: the role of age and genetic relatedness . Anim Behav 81 : 1093 - 1099 .
65. Bradshaw GA , Schore AN , Brown JL , Poole JH , Moss CJ ( 2005 ) Elephant breakdown . Nature 433 : 807 .