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This article appeared in a journal published by Elsevier. The attachedcopy is furnished to the author for internal non-commercial researchand education use, including for instruction at the authors institution
and sharing with colleagues.
Other uses, including reproduction and distribution, or selling orlicensing copies, or posting to personal, institutional or third party
websites are prohibited.
In most cases authors are permitted to post their version of thearticle (e.g. in Word or Tex form) to their personal website orinstitutional repository. Authors requiring further information
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Conflicts between Amur (Siberian) tigers and humans in the Russian Far East
John M. Goodrich a,⇑, Ivan Seryodkin a,b, Dale G. Miquelle a, Sergei L. Bereznuk c
a Wildlife Conservation Society, 2300 Southern Boulevard, Bronx, NY 10460, USAb Institute of Geography, Far Eastern Branch of the Russian Academy of Sciences, 7 Radio Street, Vladivostok 69004, Russiac Phoenix Fund, 2 Petra Velikogo St., office 409, Vladivostok 690091, Russia
a r t i c l e i n f o
Article history:Received 16 May 2010Received in revised form 30 September2010Accepted 25 October 2010Available online 19 November 2010
Keywords:Human–tiger conflictAmur (Siberian) tigerPanthera tigris altaicaRussian Far East
a b s t r a c t
In 1999, the Russian Federation created a Tiger Response Team (TRT) to investigate and intervene inhuman–tiger conflicts. We examined data collected on human-Amur tiger (Panthera tigris altaica) con-flicts from January 2000 through February 2009 to: (1) summarize and characterize human–tiger con-flicts in the area, (2) examine causes of human–tiger conflicts, and (3) attempt to evaluate theeffectiveness of the TRT. The team investigated 202 conflicts. Both attacks on humans and depredationswere greatest in winter (X2 = 9, df = 3, P = 0.03 and X2 = 64, df = 3, P < 0.001, respectively). Tiger depreda-tion on domestic animals was the most common type of conflict reported (57%), followed by tigers nearhuman habitations (22%), miscellaneous conflicts (12%), and attacks on humans (9%). Dogs were killedmore commonly than other domestic animals (63% of 254 animals), likely because livestock were wellmanaged. Nineteen attacks on humans were recorded resulting in 11 injuries and 2 deaths. Nearly fourtigers per year (n = 32 tigers) died, were killed, or were removed from the wild, and all but 1 of 20 tigerskilled or removed from the wild by the TRT were considered unfit to survive in the wild. Attacks onhumans (n = 19) were most often (77%) by wounded tigers (80% of injuries were human-caused) andcommonly provoked (47% of attacks). The effectiveness of interventions focused on reducing depredationon domestic animals was unclear, but data suggested that removal of injured and other unhealthy tigersfrom the wild by the Tiger Response Team resulted in fewer human deaths. Our recommendationsinclude that the TRT continues to work to reduce conflict by rapidly removing debilitated tigers fromthe wild, explores different methods and technologies for reducing depredation on domestic animals,and increases their efforts to maintain tigers in the wild through telemetry monitoring, translocation,and rehabilitation of orphaned cubs. Further, standardized data should be collected to evaluate all inter-ventions, with information from evaluation guiding an adaptive management component of theirhuman–tiger conflict mitigation activities. This process should occur across tiger landscapes in Asia toallow rapid assessment of interventions.
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1. Introduction
Human-carnivore conflict is one of the leading causes of declinein large carnivore populations, particularly for large felids, andreducing conflict is critical to successful conservation efforts (In-skip and Zimmermann, 2009; Treves and Karanth, 2003; Woodr-offe et al., 2005a). Among the large cats, tigers (Panthera tigris)are perhaps most notorious for conflict with man because they his-torically killed large numbers of people (Boomgaard, 2001; McDo-ugal, 1987; Nyhus et al., 2010). Today conflict between humansand tigers occurs almost everywhere there are tigers (Barlowet al., 2010; Goodrich, 2010; Karanth and Gopal, 2005; McDougal,
1987; Nyhus and Tilson, 2010), and negative impacts of human–ti-ger conflict include loss of human life and livelihood (e.g. when ti-gers kill domestic animals), negative attitudes towards tigerconservation, retaliation killing of tigers, and increased poachingwhen poachers take advantage of tigers depredating domestic ani-mals near villages (Karanth and Gopal, 2005; Matyushkin, 1985;Miquelle et al., 2005; Nikolaev and Yudin, 1993; Nyhus and Tilson,2010). Thus, both directly and indirectly, human–tiger conflict re-sults in increased tiger mortality; yet, decreasing human-causedmortality of tigers is critical to population persistence (Chapronet al., 2008; Goodrich et al., 2008; Kenney et al., 1995).
Human–tiger conflicts have declined considerably over the pastcentury (Boomgaard, 2001; Nyhus et al., 2010) with dramatic de-clines in tiger numbers range-wide (Walston et al., 2010). How-ever, as tiger populations become smaller and more isolated(Dinerstein et al., 2007), the impact of conflict-related mortalityon population viability increases (Nyhus and Tilson, 2010). Further,current tiger conservation initiatives, including programs by inter-
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national NGOs (Non-Governmental Organizations) and national ti-ger recovery plans of some range states, aim to increase tiger pop-ulations by 50–100% (e.g. Department of Wildlife and NationalParks Peninsular Malaysia, 2008; Panthera, 2010; Walston et al.,2010). If these initiatives are successful, the potential for human–tiger conflicts will increase as populations in protected areas reachcarrying capacity, forcing young or infirm animals into human-dominated landscapes (Karanth and Gopal, 2005). Thus, efforts toreduce conflict are critical.
There are about 128,000 km2 of occupied Amur tiger (P. t. alta-ica) habitat in the Russian Far East, 93% of which is multiple-useforest land where tigers and humans coexist (Matyushkin et al.,1999; Miquelle et al., 1999; Sanderson et al., 2006). Human useof these lands includes logging, agriculture, livestock grazing,hunting, and collection of non-timber forest products. Unlike muchof South Asia, which can be characterized as islands of tiger habitatsurrounded by a sea of humanity, the situation in the Russian FarEast is the opposite; humans live in small settlements that are is-lands dispersed in a sea of tiger habitat (Miquelle et al., 2010a;Sanderson et al., 2006). Hence, while density of both humans(12/km2 in Primorski Krai) and tigers (<1/100 km2) are low in theregion, reports of human–tiger encounters became common inRussia after the tiger population rebounded from a low of about40 in the 1940s (Gorokhov, 1983; Khramtsov, 1995; Matyushkin,1973; Miquelle et al., 2010b; Nikolaev and Yudin, 1993; Shishkin,1967) because many settlements are completely surrounded by ti-ger habitat, and people extensively exploit tiger habitat for com-mercial and personal uses (Miquelle et al., 2005).
Attacks on humans occur annually in several areas of the tiger’srange, including Russia, and because they provoke a strong re-sponse from local people, they can negate conservation efforts(Barlow et al., 2010; Chowdhury and Sanyal, 1985; Gurung et al.,2009; Karanth and Gopal, 2005; Miquelle et al., 2005; Nyhus andTilson, 2004). These attacks are poorly understood and there is lit-tle published data on the nature and causes of such attacks (Gur-ung et al., 2009; McDougal, 1987; Quigley and Herrero, 2005).Yet an improved understanding should result in fewer attacks, im-proved local attitudes towards tigers, and fewer human and tigerdeaths. The same is true for tiger depredation on domestic animals.
Prior to the 1990s, human–tiger conflict was mitigated in Russiathrough a State program that compensated for damages caused bytigers and by occasionally killing ‘problem’ tigers (Nikolaev, 1985).However, government support was discontinued during the polit-ical and economic turmoil of Perestroika and human–tiger conflictincreased dramatically in the 1990s (Miquelle et al., 2005). In late1999, Inspection Tiger (Ministry for Natural Resources, RussianFederation) created a special ‘‘Tiger Response Team’’ (TRT) withthe goal of reducing threats and perceived threats caused by tigersto humans, and reducing tiger mortality associated with human–tiger conflict (Miquelle et al., 2005). To our knowledge, Russia isthe first country ever to designate a government team whose pri-mary responsibility was to respond to human–tiger conflicts. Weexamined data collected by the TRT from 2000 through February2009 to: (1) summarize and characterize human–tiger conflict inthe Russian Far East, (2) examine causes of human–tiger conflict,and (3) attempt to evaluate the effectiveness of the TRT’s interven-tions. We use the results to develop recommendations forimprovement of interventions into human–tiger conflict.
2. Methods
2.1. Study area
The TRT worked throughout the range of Amur tigers in Russia,which is primarily confined to the Russian Far East provinces
(Krais) of Primorye and Khabarovsk (Miquelle et al., 1999). In2005, there were estimated to be <400 adult and subadult tigersin Russia (Miquelle et al., 2007), with about 95% occurring in thefoothills and main ranges of the Sikhote-Alin Mountains, the crestof which runs north–south through most of the tigers’ currentrange in Russia. The area consists of two Tiger Conservation Land-scapes (Sanderson et al., 2006) representing a zone where EastAsian coniferous–deciduous forests and the boreal forests merge,resulting in a mosaic of forest types (Miquelle et al., 2010a).
In contrast to most tiger habitat in Asia, the climate of Amur ti-ger range in Russia is marked by cold, snowy winters that likelystress tigers and ultimately limits their geographic distribution(Miquelle et al., 2010a). The climate is monsoonal, with 75–85%precipitation (650–800 mm in central Sikhote-Alin) occurring asrain between April and November. Snow depth varies greatly by lo-cale and year, but averages 22.6 cm in mid-February (60-yearmean) in the inland central Sikhote-Alin and only 13.7 cm on thecoast (data from Primorye Weather Bureau). However, heavysnowfalls, most common in March, can have dramatic impacts onungulate and tiger populations. The January monthly mean tem-perature on Sikhote-Alin Biosphere Zapovednik (roughly the geo-graphic center of tiger range) is �22.6 �C on the inland side ofthe central Sikhote-Alin Mountains, but the Sea of Japan moderatestemperatures and snow depths on the coastal side of the range,when the January monthly mean is �12.4 �C. The July monthlymean temperature is 15 �C on the coast and 18.4 �C inland.
2.2. Characteristics of human–tiger conflicts
We used records of human–tiger conflict collected by the TRTand compiled and reported by Phoenix Fund, as well as supple-mental data collected by the Wildlife Conservation Society whileworking with the TRT from January 2000 through February 2009.‘‘Conflicts’’ were defined as any reports of contact between tigersand humans that were responded to and confirmed by the TRT.We categorized conflicts as: depredation on domestic animals(hereafter referred to as ‘‘depredation(s)’’, attack on humans, tigersnear isolated human habitations (e.g. hunting cabins), or miscella-neous, and recorded location as: in villages, near isolated humanhabitations, in the forest, or unknown. We used a chi-square testto assess seasonal differences in number of conflicts, with seasonsdefined as winter (December–February), spring (March–May),summer (June–August), and autumn (September–November). Wetallied the result of each conflict in terms of injuries and deathsof humans, domestic animals, and tigers.
2.3. Causes of human–tiger conflicts
We analyzed tigers involved in conflict to determine if health-issues were associated with conflicts. In most cases when tigerswere captured, we collected blood samples for disease analysis(Goodrich et al., 2005; Quigley et al., 2010) and estimated physicalcondition through gross physical examination (Goodrich et al.,2001). Dead tigers were necropsied by a variety of different organi-zations, including WCS, the Veterinary School at the PrimorskayaState Academy of Agriculture (Ussurisk, Primorski Krai, Russia),and Lazovsky State Zapovednik (Lazo, Primorski Krai, Russia).Health status of tigers was categorized non-mutually exclusivelyas apparently healthy, wounded by people, wound cause naturalor unknown, emaciated/diseased, and orphaned cubs. Where morethan one category was applicable, the category that was believedresponsible for an animal’s condition was chosen. For example,animals categorized as wounded were often emaciated as well,but we assumed that the wound resulted in emaciated condition.Cubs captured or found dead with no evidence of a mother wereassumed to be orphaned.
Where tigers attacked humans, we used data on behavior of thepeople and tiger involved to elucidate causal factors and catego-rized attacks as provoked (the tiger was intentionally approachedor shot by people), accidental meetings (a person and tiger cameinto close proximity accidentally, provoking an aggressive re-sponse from the tiger), predation attempt (a tiger approachedand attacked a person without provocation), and mothers defend-ing cubs.
2.4. Interventions and evaluation
Responses by the TRT were categorized as monitoring (TRT per-sonnel or local informant in place to report if tiger returned), scar-ing or hazing, successful capture, unsuccessful capture, and tigershot in self defense. Usually, tigers were captured and anesthetizedas described by Goodrich et al. (2001, 2010a,b), but some, espe-cially emaciated cubs, were captured by the TRT or local villagers.Captured tigers were either released on site, translocated and re-leased, removed to zoos, or euthanized; all released tigers wereequipped with radio-collars (Goodrich and Miquelle, 2005a,b). Col-lared tigers were monitored and hazing was attempted when a ti-ger approached human habitations (Goodrich and Miquelle,2005b).
We evaluated the TRT’s interventions in four ways. First, weused a chi-square test to compare the frequency of repeat offensesfor cases where hazing was used to those where no action beyondmonitoring was used. We predicted that if hazing was effective,there would be fewer repeat offenses following hazing. Second,we monitored six tigers using radio-telemetry and snow-tracking(Goodrich and Miquelle, 2005a,b; 2010) and attempted to hazethem using pyrotechnics when they approached settlements. Haz-ing was considered successful if the tiger ran away and left thearea. Third, we extrapolated our data from 2000 to February2009 through the end of 2009 assuming a constant attack ratefor the decade, and then used a chi-square test to compare thenumber of attacks on humans to the previous two decades usingdata reported in Miquelle et al. (2005). Fourth, we used simple lin-ear regression analysis to test for changes in number of depreda-tions from 2000 to 2009. The effectiveness of translocations wasassessed elsewhere (Goodrich and Miquelle, 2005a).
3. Results
3.1. Characteristics of human–tiger conflicts
From January 2000 through February 2009, 202 cases of hu-man–tiger conflict were recorded by Inspection Tiger (Table 1).The number of conflicts reported per year (�x = 22 ± 14, range 6–39) jumped an order of magnitude between 2000–2003(8.5 ± 6.2) and 2004–2008 (32.6 ± 13.9), so we tested post hoc forstatistical significance (t = �6.9, df = 7, P < 0.001). Conflict variedseasonally, with levels highest in winter and declining steadilythrough the rest of the year (Fig. 1). Both attacks on humans and
depredations were greater in winter (X2 = 9, df = 3, P = 0.03 andX2 = 64, df = 3, P < 0.001). Tiger depredation on domestic animalswas the most common type of conflict reported (57%), followedby tigers near human habitations (22%), miscellaneous conflicts(12%), and attacks on humans (9%).
Tigers killed 254 domestic animals and injured 23. Dogs (Canislupus familiaris) were killed most commonly (63% of 254 animals),followed by cows (Bos primigenius) (17%), horses (Equus ferus)(12%), sheep (Ovis aries) (6%), and ‘‘farmed’’ sika deer (Cervus Nip-pon) (4%). Most depredations occurred in villages (57%) or at iso-lated farms, cabins, and logging camps (30%).
Nineteen attacks on humans were recorded by Inspection Tiger,resulting in 11 injuries and 2 deaths. Most attacks occurred in re-mote forested areas (53%), while 26% occurred in villages, 16% nearisolated farms, cabins, or logging camps. In only 1 case was a per-son eaten by a tiger, but it was not known if this person was killedby the tiger.
Of 45 reports of tigers near villages or isolated human habita-tions, 35 were reports of tracks and 10 were visual sightings. In 4cases, depredations were later reported in the same town and in1 case, a tiger later attacked a person in the same town. Of 10 vi-sual sightings, 4 were of diseased or wounded tigers that were latercaptured; of these, 2 were diagnosed with canine distemper virus(Goodrich et al., 2005; Quigley et al., 2010).
Of 25 miscellaneous conflicts reported, 12 (48%) were orphanedcubs found dead or in poor condition and 8 (32%) were woundedtigers (in 1 case, a poacher reported wounding a tiger because hefeared the tiger might attack someone), 2 were animals injuredor killed in collisions with vehicles, 1 was a tiger killed by anothertiger, 1 was a tiger that stole a pumpkin from a shed, and 1 was atiger that destroyed a border-guard mannequin on the Russia-Chi-na border.
Of the 202 cases investigated by the TRT, 32 tigers, or nearly 4tigers per year, died or were removed from the wild (Table 1). Ofthese, 13 were cubs deemed unable to survive in the wild due toyoung age or poor physical condition, 8 were wounded, 6 wereemaciated/diseased, the condition of 4 was not reported, and 1was apparently healthy. Twelve of the 32 tigers that died were
Table 1Responses to human–tiger conflict by the Tiger Response Team in the Russian Far East, January 2000–February 2009.
Type of conflict No. of conflicts Response
Monitor Hazinga Capture attempt(successes)
Tiger released Tiger removed orkilled by TRT
Tiger diedotherwise
Attack on person 19 10 5 3 (2) 1 7 0Depredation on domestic animals 115 62 41 24 (12) 4 7 2Approach human habitations 45 17 10 6 (6) 1 1 5Other 25 0 0 6 (6) 1 5 5
a In the text, 54 cases of hazing were used in analysis while 56 are reported here because 2 cases of livestock depredation also involved attacks on humans.
0
20
40
60
80
100
120
winter spring summer fall
Num
ber o
f Con
flcits
.
total depredation people
Fig. 1. Seasonality of tiger–human conflict in the Russian Far East, January 2000–February 2009.
found dead by the TRT and 20 were removed from the wild, eutha-nized, or killed in self-defense by the TRT.
3.2. Causes of human–tiger conflicts
The health of tigers was assessed in only 15 of 115 (13%) depre-dation cases. Of those, 6 tigers were orphaned cubs, 5 were emaci-ated and/or diseased, 4 were wounded animals, and 4 wereapparently healthy animals (multiple categories in four instances).For tigers approaching human habitations, the health condition ofthe tiger was assessed in 10 cases and included 2 wounded ani-mals, 4 emaciated and/or diseased animals (2 diagnosed with ca-nine distemper), and 4 cases of orphaned cubs. For miscellaneousconflicts, the health condition of the tiger was assessed in 13 casesincluding 4 cases of emaciation/disease, 8 orphaned cubs, and 1healthy animal. The latter was a tiger rescued from a poacher’ssnare and released. The health of tigers that attacked people wasassessed in 13 of 19 cases (Fig. 2). Tigers had human-causedwounds in 62% of the cases and in half of these cases, the tiger at-tacked a person who wounded it in a poaching attempt.
We examined the role of human and tiger behavior in attacks onpeople. Attacks were directly provoked by humans in 9 cases (47%),accidentally provoked by humans in 7 cases (37%), predation at-tempts in 2 cases (11%), and defense of cubs in one case (5%)(Fig. 3). Attacks were provoked by humans attempting to poach ti-gers (n = 5) and by approaches by the TRT (n = 4) to wounded ti-gers. Accidental provocations were the result of 2 cases in whichtigers chased dogs that ran to their owners, 4 cases when peopleaccidentally approached too closely (two related to depredations),and 1 case of a tiger defending a kill (no human injuries). Therewere two cases of apparent predation attempts by tigers on peopleand 1 case (5% of attacks) of a tiger defending a cub that had beenkilled in a vehicle collision; no one was injured.
3.3. Interventions and evaluation
Of 202 confirmed reports, the TRT monitored 89 situations, at-tempted to scare the tiger using pyrotechnics (fireworks or signalrockets), gunshot, or by chasing with a vehicle in 56 cases, initiatedcapture activities in 39 cases, and shot the tiger in self defense in 6cases (four provoked attacks and two accidental meetings). Of 26tigers captured, 10 were removed from the wild because of physi-cal impairments such as wounds or emaciation, 5 died fromwounds or disease unrelated to capture, 4 were euthanized, 3 werereleased on site, and 4 were translocated (Goodrich and Miquelle,2005a). Of 7 tigers released, 3 were held in captivity for 7 days to1 year. The TRT killed 10 tigers; 6 were shot in self defense and 4were euthanized. Of the tigers euthanized, all but one adult hadphysical impairments that rendered them unfit to survive in thewild.
In 54 cases where hazing was used, the tiger continued to causeconflicts in 16 cases (30%), but tigers continued to cause conflict inonly 4 of 57 cases where hazing or other interventions beyondmonitoring were not used ({2 = 8.1, df = 1, P = 0.004). In 6 caseswhere tigers were radio-collared and released, tigers caused fur-ther conflict in 4 cases, but in only 1 case were we able to effec-tively monitor and haze the tiger.
The number of people killed by tigers in the 2000s decreasedsignificantly ({2 = 7.1, P = 0.008) from the previous decade, butthe number of people injured did not ({2 = 0.04, df = 1, P = 0.8;Fig. 4). The differences in mean number of conflicts per year re-ported in 2000–2003 and 2004–2008 were related to administra-tive changes in Inspection Tiger (see Section 4.1), so only datafrom 2004 to 2008 were used for regression analysis to test forchanges in depredations per year. The number of depredationsper year decreased steadily during this time (R2 = 0.86, P = 0.02).
4. Discussion
4.1. Characteristics of human–tiger conflicts
The difference in number of reports between 2000–2003 and2004–2008 was probably related to poor reporting by TRT to Phoe-nix Fund in early years; indeed, there were a number of conflictsthat WCS responded to with TRT from 2000 to 2003 that werenot in the Phoenix Fund database. Administrative changes in2003 resulted in improved reporting and all conflicts to whichWCS responded after 2003 were included in the Phoenix Funddatabase. However, we believe that there was no difference inreporting of attacks on humans between the two periods becauseWCS worked very closely with the TRT and responded with themto most cases where tigers attacked people and such attacks were
0
10
20
30
40
50
60
70
Wounded by person
wound cause unknown
emaciated/disease
Perc
ent o
f tig
ers
at
tack
ing
peop
le .
Fig. 2. Condition of 13 tigers that attacked people in the Russian Far East, January2000–February 2009.
05
101520253035404550
Provoked Accidental meeting
Predation Defending Cubs
Perc
ent o
f atta
cks
Reason for attack
Fig. 3. Reason for 19 tiger attacks on people in the Russian Far East, January 2000–February 2009.
0
5
10
15
20
25
30
1970s 1980s 1990s 2000s
Num
ber o
f inc
iden
ts
people killed people injured total attacks on people
Fig. 4. Number of tiger attacks on people per decade in the Russian Far East.
widely publicized (Miquelle et al., 2005) and media searches didnot reveal attacks on people that were not in our database.
Depredation was most common in winter despite the fact thatlivestock were mostly stall-fed in winter so were less available totigers, and most depredations occurred in villages. Other studieshave found depredation to be greatest in seasons when livestockwere free-ranging and not in villages (Johnson et al., 2006; Liet al., 2009; Sangay and Vernes, 2008). However, these studieswere conducted in areas where livestock were allowed to rangeunattended in tiger habitat, so tigers were likely taking advantageof seasonal increases in livestock availability. In contrast, livestockwere normally tended by herders and were grazed along theperiphery of tiger habitat in Russia. We believe that depredationsincreased in winter in Russia because Amur tigers were morestressed physically due to intense cold, deep snow, and in somecases, lower prey availability, particularly of small prey (e.g. bad-gers [Meles meles]) that are easily captured and hence importantto sick or wounded animals. This likely forces some tigers into vil-lages in search of domestic prey.
Attacks on humans were also more common in winter, possiblybecause hunting season occurs in late fall through winter, resultingin an increase in number of people in the forest and an increase intiger–human encounters (Miquelle et al., 2005). Also, tigers areeasier to pursue in winter when tracks can be followed in snow,congregations of avian scavengers make kills obvious, and lack ofdense herbaceous vegetation in the forest understory make tigerseasier to see (Yudakov and Nikolaev, 1987). This likely resultedin increased poaching in winter and consequently, increased con-flict associated with wounding of tigers. Attacks on humans havebeen positively correlated with seasonal use of tiger habitat by hu-mans in Bangladesh (Barlow, 2009), but not in some other areas(Sanyal, 1987; Gurung et al., 2009).
Both depredations and attacks on humans were rare relative tomany other countries (Barlow et al., 2010; Chowdhury and Sanyal,1985; Gurung et al., 2009; Karanth and Gopal, 2005; McDougal,1987; Nyhus and Tilson, 2004, 2010; Sangay and Vernes, 2008),probably due to the relatively low density of tigers, humans, andlivestock in Russia. Additionally, livestock were well cared for inRussia and humans were often armed and able to prevent attacks.In contrast to other areas (e.g. Madhusudan, 2003; Wang and Mac-donald, 2006), dogs were killed more frequently in Russia thanwere livestock, probably because dogs were often staked out nearthe edges of yards, making them vulnerable to tigers. However,depredation on dogs is not unique to Russia (Nugraha and Sugardj-ito, 2009). While dogs are often used to prevent human-carnivoreconflict elsewhere (e.g. Green et al., 1984; Khan, 2009), in Russiadogs were the focus of many conflicts, including attacks on peoplewhen dogs chased by tigers ran to their owners (this study; Niko-laev and Yudin, 1993). While the loss of a dog usually results inlower direct economic loss for local people than the loss of acow, the indirect economic loss can be high because many peopledepend on dogs for hunting and hunting for income and food(Miquelle et al., 2005). Hence, depredation on dogs elicits a strongresponse and has been a primary reason for retaliation killing(Gorokhov, 1983).
Most depredations (87%) occurred in villages or at isolated hab-itations, likely because domestic animals were generally not al-lowed to range freely in the forest. This is a stark contrast tomost parts of the tiger’s range, where improved livestock manage-ment is considered one of the most important aspects of prevent-ing conflict (Johnson et al., 2006; Karanth and Madhusudan, 2002;Li et al., 2009; Nyhus and Tilson, 2010; Treves and Karanth, 2003).Thus, while managing domestic animals in ways that reduce theiravailability to tigers is important to preventing depredation, the fo-cus in Russia should be on dogs, so long as local people continue tomanage their livestock well. Hunters should be discouraged from
using dogs as much as possible, but many hunters will continueto use dogs regardless of the risk. In Russia, dogs are commonly al-lowed to roam free or are kept as guard dogs chained to a doghouse near homes. Dogs in high risk areas, such as remote huntingcabins and farms, and those in villages surrounded by tiger habitat,should be kept indoors, or in covered pens at night, which wouldallow them to continue to function as guard dogs.
In some cases, reports of tigers near human habitations mayhave been precursors to depredation, as was found in Indonesia(Nugraha and Sugardjito, 2009), but whether or not the tiger waspresent in an area prior to depredation was generally not known.In two known cases, tigers that spent several days near loggingcamps eventually attacked people. In other cases, tigers approach-ing human habitations were diseased – 2 animals diagnosed withcanine distemper entered a village and a house (without aggres-sion), and in one case a tiger suspected of having rabies attackeda logger (Goodrich et al., 2005).
Miscellaneous incidences primarily included orphaned cubsand wounded animals. Orphaned cubs were found and capturedmore than once per year and in many cases, the mother was be-lieved poached. Poaching of mothers is the leading cause of cubmortality in Russia (Kerley et al., 2003; Goodrich et al., 2008; Niko-laev and Yudin, 1993). In 2 cases, orphaned cubs were translocatedand released back into the wild and at least 1 of these cubs sur-vived (Goodrich and Miquelle, 2005a). In the remaining cases,the cubs either died or were sent to zoos. High removal rates ofcubs can have serious impacts on the Amur tiger population (Cha-pron et al., 2008) and recovery from a major decline in Amur tigernumbers in the mid-20th Century was hindered by high removalrates of cubs (Abramov, 1958). An alternative to removal is tomaintain cubs in the wild, especially those that were orphanedafter 6 months of age. Goodrich and Miquelle (2005b) report 3cases (i.e., 3 litters totaling six cubs) when cubs orphaned atP6 months of age successfully survived (with artificial feeding in1 case).
In many cases, small amounts of blood in tiger tracks in snowassociated with superficial wounds on pads of paws (which arecommonly noted in normal healthy tigers) led to false reports ofwounded animals. However, in 3 cases of reports of wounded ti-gers, animals were wounded in poaching attempts, including acase when the poacher himself reported wounding a tiger and an-other case in which a tiger was found alive in a poacher’s snare.
4.2. Causes of human–tiger conflicts
The condition of most (87%) tigers that depredated on domesticanimals was unknown because the tiger was not captured, killed,or found dead. In many of these cases, the offending individualmay have survived and returned to the forest. However, in severalcases, it was rumored that the tiger was subsequently poached.Such retaliation killings have been reported in the past (Gorokhov,1983; Miquelle et al., 2005; Nikolaev and Yudin, 1993), but areprobably more common now that a strong black market has devel-oped for tiger parts in Russia with the opening of its borders fol-lowing Perestroika. Poaching has been linked to depredationelsewhere and may be an incentive to poorly manage livestock(Johnson et al., 2006; Karanth and Gopal, 2005; Kawanishi et al.,2010).
Of 15 depredating tigers for which information was available,only 20% were apparently healthy; the remaining animals werewounded, emaciated/diseased, or orphaned cubs. Further, thatmost conflicts occurred in winter when tigers were most stressed(Miquelle et al., 2010a) suggests that depredation was related toseasonal shortages of wild foods. Sick or wounded animals thatpresumably had greater difficulty obtaining food more commonlypreyed on domestic animals than healthy animals. In many cases,
samples were not analyzed for infectious disease and necropsieswere often incomplete, so we likely underestimated rates ofwounding and disease.
Most tigers that attacked people were wounded and 63% werewounded by people. None that were assessed were consideredhealthy, although data may be biased towards detecting unhealthyanimals because healthy tigers were more likely to escape and dis-appear into the forest. Nonetheless, the data clearly indicate thatwounding of tigers by people, usually in botched poaching at-tempts, often leads to attacks on people. The person attackedmay be the poacher who wounded the tiger, or an innocent personwho later encounters that tiger. Further, nearly half of all attackswere provoked by people intentionally pursuing the tiger eitherto poach it, or, in the case of attacks on the TRT, to investigate aconflicts caused by the wounded tiger. Similar trends have beenobserved in the past in Russia (Matyushkin, 1985; Nikolaev andYudin, 1993; Zhivotchenko, 1976). Gurung et al. (2009) found that>50% of tigers that attacked humans had physical impairments,and injuries have been tied to conflict in other large cat species(e.g. Hoogesteijn et al., 1993; Loveridge et al., 2010; Rabinowitz,1986).
Although man-eating tigers were reportedly common in the19th century in Russia, over the past century it has become extre-mely rare (Khramtsov, 1995; Miquelle et al., 2005; Shishkin, 1967)and that trend continues as in other parts off the tiger’s range(Boomgaard, 2010; Nyhus et al., 2010). Both cases of predationon humans over this study period were only presumed predation,i.e., evidence was not conclusive. The tigress that attacked a loggerhad symptoms consistent with rabies, but samples were not col-lected by TRT. The logger was treated for rabies as standard proce-dure for wild animal bites. Otherwise, attacks on humans weremostly provoked by people attempting to poach the tiger. Reduc-ing poaching of tigers may significantly reduce the number of at-tacks on people in Russia.
4.3. Interventions and evaluation
Our capacity to evaluate the effectiveness of the TRT at reducingdepredations on domestic animals was limited for a number of rea-sons. First, data on trends in conflict over time were dependent onconsistent reporting of conflict by local people and data collectionand reporting by the TRT. Reporting by TRT differed between2000–2003 and 2004–2008 (see above), and reporting of conflictby local people may have also varied, e.g. as the existence of theTRT became better known later in the study. Second, many caseswere not followed-up on or follow-up investigations were not re-ported. Third, because tigers were rarely seen, it was usually notpossible to determine the effectiveness of hazing techniques suchas pyrotechnics, because proximity to the tiger and the tiger’s re-sponse were rarely known. Last, the data were confounded byother factors, such as a decline in tiger abundance. Nonetheless,some conclusions can be made.
Incidents of repeated conflict were more common in caseswhen tigers were hazed than in cases when tigers were not – theopposite of what we would predict if hazing were effective. This re-sult is likely due to bias in that hazing was generally used in caseswhere there was already repeated conflict (usually depredations)or evidence that the tiger was still in the area; whereas hazingwas not used if it appeared that the tiger had left the area. None-theless, tigers continued to cause conflict 30% of the time followinghazing, suggesting that it may be of limited effectiveness. In thesingle case where a radio-collared tiger was hazed, it appeared thatsignal rockets were effective only when used in very close (10 m)proximity to the tiger. Using radio-telemetry to monitor tigersand deter them from causing further conflict failed in 3 of 4 casesbecause tigers moved over too large of an area (hundreds to thou-
sands of km2; Goodrich et al., 2010a,b) to monitor them effectively.However, the use of GPS collars with satellite downloads may in-crease effectiveness (Goodrich and Miquelle, 2005a, 2010). Furtherproblems included that the TRT often arrived at depredation scenesafter the tiger had left the area because the Team had to cover vastareas over difficult roads in winter conditions. They sometimes hadto leave conflict areas too soon because they had to respond to re-ports of conflict in other areas. Creating more teams would be anobvious solution, but may be unjustifiably expensive. However, lo-cal wildlife law enforcement officials could be given basic trainingand equipment to respond to many types of human–tiger conflicts,while the TRT focused on cases requiring intensive intervention.
Salkina (2000) used similar types of hazing on Amur tigers withpositive results and a variety of visual and acoustical repellents(e.g. fireworks, signal flares, cracker shells, lights, sirens) and pro-jectiles (e.g. rubber bullets) have been used on other carnivoreswith some success (review in Breitenmoser et al., 2005), suggestingthat further attempts at hazing are warranted, especially if equip-ment designed specifically for wildlife were to be used. The 43%decrease in mean number of depredations per year from 2004 to2008 may have been related to the TRT’s interventions. However,the decrease may have been associated with a decline in tigernumbers in Russia (Miquelle et al., 2009).
Data do suggest that the TRT was effective at reducing attackson humans and particularly human deaths. The reduction in num-ber of human deaths between 1990s and 2000s, was likely the re-sult of TRT killing or removing from the wild wounded andemaciated or diseased animals that were a danger to people. TheTRT was charged seven times and killed the offending tiger in selfdefense in 6 cases, suggesting that most of these tigers were a dan-ger to people. All seven of these tigers were considered unfit to sur-vive in the wild because of wounds or emaciated condition.Removal or lethal control of tigers that attack people has beencommonly practiced and sometimes recommended as a methodfor reducing attacks on humans (Barlow et al., 2010; Gurunget al., 2009; Karanth and Gopal, 2005; Kawanishi et al., 2010; Ny-hus and Tilson, 2010; Smith et al., 2010; Treves and Naughton-Tre-ves, 2005). However, some have advocated fitting healthy tigerswith telemetry devices and monitoring them to keep people awayfrom the tigers and haze the tigers when necessary (Barlow et al.,2010; Smith et al., 2010). There were no repeated cases of a tigerkilling people between 2000 and 2009, but there were 2 cases ofwounded tigers killing >1 person during the previous decade(Miquelle et al., 2005), suggesting that rapid removal of woundedanimals is important to reducing attacks on humans. Further,poaching rates were very high in the 1990s following the turmoilof Perestroika, resulting in high rates of wounding (Nikolaev andYudin, 1993) which may have resulted in greater attacks on hu-mans. Poaching, and hence wounding and related attacks on hu-mans, may have decreased during the following decade. Whilethere are no range-wide data available for comparison, poachingrates were lower during the first half of the 2000s on the Sikh-ote-Alin Zapovednik than in the 1990s (Goodrich et al., 2008,2010a,b). Last, the TRT conducted anti-poaching work while not in-volved in conflict work and likely contributed to a reduction in ille-gal killing of tigers.
In addition to reducing attacks on humans, the TRT served otherimportant functions. First, they removed from the wild several ti-gers that were not fit to survive. These included wounded and dis-eased animals that did not cause conflict, but that wandered intotowns, approached habitations, and even entered buildings. Suchsituations represent a potential danger to people, domestic ani-mals, and tigers and require intervention by trained personnel.Without the TRT, many of these animals would likely have beenpoached and some may have attacked people or domestic animals.Second, by responding to conflicts, the TRT was responding to the
needs and assuaging the fears of local people, and providing analternative to killing tigers. This may have resulted in increasedtolerance and decreased poaching of tigers, although data areneeded to test this. Hence, even if the TRT was not effective atreducing depredation, their response likely served an importantfunction. Last, because the TRT also had wildlife law enforcementauthority and often maintained a presence in areas with active hu-man–tiger conflict, they likely prevented conflict-related poachingof some tigers.
4.4. Recommendations
We believe that the TRT played an important role in reducinghuman–tiger conflicts by removing wounded, diseased, or other-wise unfit animals from the wild and that this, combined with theiranti-poaching activities and the likelihood that they improve localtolerance of tigers, justifies their existence. Their protocol of inves-tigate, monitor, haze, capture and evaluate, and then either releasethe tiger or remove it from the wild seems to be a logical and effec-tive approach. However, their effectiveness at reducing depreda-tion on domestic animals was dubious, and data are needed todetermine if they improved public acceptance of tigers or reducedpoaching. We recommend that they continue to quickly captureand assess tigers that attack people or appear to be wounded, dis-eased, or otherwise unfit for survival in the wild. For dealing withdepredations on domestic animals, we recommend that they ex-plore additional techniques and technologies (e.g. those reviewedin Breitenmoser et al., 2005) and collect data necessary for evalua-tion of effectiveness. Telemetry monitoring with GPS collars of ti-gers that repeatedly cause conflict may improve hazing efforts byhelping the TRT to get close to the tiger and will provide data onthe response of the tiger to hazing. The TRT is currently dependenton WCS for capture and telemetry expertise and we recommendcontinued training for TRT personnel so that they may operatemore independently.
To further reduce loss of tigers due to conflict, the TRT shouldcontinue to use and improve upon methods to maintain tigers inthe wild, including translocation and returning cubs to the wild,using artificial feeding or rehabilitation in captivity when neces-sary (Goodrich and Miquelle, 2005a,b). While translocations havemet with mixed success for tigers and other carnivores (Goodrichand Miquelle, 2005a; Linnell et al., 2005; Loveridge et al., 2010),it is becoming an important tool for re-establishing populationsor supplementing small populations (Hunter et al., 2007). We alsorecommend that the TRT continue its anti-poaching activities be-cause poachers take advantage of conflict situations to kill tigers(Kawanishi et al., 2010; Karanth and Gopal, 2005; Nugraha andSugardjito, 2009) and because it provides a function for the teamwhen there are no conflicts to manage.
Methodology for evaluating interventions, including standard-ized data sheets and a central data base, are needed (Nyhus andTilson, 2010; Woodroffe et al., 2005b). Measures of effectivenessshould reflect the TRT’s goal of reducing loss of human life and live-lihood and tiger mortality. In this respect, appropriate measuresshould include changes in number of human and domestic animalinjuries and deaths, and number of tiger mortalities (Barlow et al.,2010), with both measured relative to overall tiger numbers. Hu-man tolerance towards tigers should also be surveyed to determineif the TRT’s interventions should improve attitudes towards tigers.However, because there are many potential correlates to frequencyof human–tiger conflicts (e.g. decline in prey populations may re-sult in increased conflict), these measures must include appropri-ate correlates, and will likely be useful only for measuring long-term general trends.
Additionally, each type of intervention should be evaluated. Forexample, the TRT often attempted to frighten tigers with signal
rockets and fireworks, but collected few data to determine if thesetechniques frighten tigers or discourage future depredations. Inaddition to telemetry, camera traps could be used to determine iftigers exposed to hazing cause further conflict; and photographsof tigers involved in conflict could be used to further investigatethe link between conflict and poaching (animals photographed inconflicts later showing up as confiscated carcasses or skins) (Kar-anth and Nichols, 2002). There have been few attempts to evaluateinterventions to reduce human–tiger conflict anywhere within thetiger’s range (Nyhus and Tilson, 2010; Woodroffe et al., 2005b) andextrapolation to tigers from other species is difficult because of thetremendous black-market value of tiger parts. For example, insome areas, depredation on livestock may be seen as a financialopportunity to poach a tiger because the value of a tiger on theblack market far outweighs the value of livestock lost (Johnsonet al., 2006). Thus, while reduced depredation on livestock is incen-tive to improve livestock management in many areas where tigersdo not exist, greater incentives may be necessary in areas wherethere are tigers.
A coordinated response to human-carnivore interventionsshould include programs to prevent and mitigate conflict (e.g. im-proved livestock management and insurance, respectively), educa-tion and outreach programs, legal protection and local lawenforcement agents capable of dealing with most situations, anda well-trained, government-sponsored response team to addresssituations that require serious interventions (Barlow et al., 2010;Goodrich, 2010; Gurung et al., 2009; Karanth and Gopal, 2005;Smith et al., 2010; Tilson and Nyhus, 1998; Woodroffe et al.,2005b). Along with these programs, a clear, well defined protocolshould be developed to provide guidelines for appropriate inter-ventions. We suggest development of a decision-tree type processthat will allow quick reference by field teams (e.g. Goodrich, 2010;Nugraha and Sugardjito, 2009), supported by a more extensivewritten protocol that will educate and guide response teams, aswell as provide legal protection of such teams when serious prob-lems arise (Goodrich, 2010; Karanth and Gopal, 2005; Nugraha andSugardjito, 2009; Tilson and Nyhus, 1998).
In 1999, Russia was the first country to create a government-sponsored TRT. Today, response teams currently exist in severalcountries in some form, as do proposals to create response teamsin other countries (Barlow et al., 2010; Gurung et al., 2009; Nyhusand Tilson, 2010; Smith et al., 2010; Tilson and Nyhus, 1998). Ifproperly evaluated, the activities of Russia’s TRT may serve as amodel for developing teams in other areas. As these teams develop,the process of evaluation of interventions should occur across tigerlandscapes in Asia to allow rapid assessment of interventions.
As tiger populations shrink in size, mortalities associated withhuman–tiger conflicts could have important ramifications for per-sistence of these small, isolated populations (Chapron et al., 2008;Nyhus and Tilson, 2010). Further, at the Tiger Summit, schedule tobe held in Russia in November 2010, 13 tiger range states are ex-pected to commit to increasing their tiger populations (Walstonet al., 2010) and increases in tiger numbers will likely lead to in-creased conflict (Karanth and Gopal, 2005). Hence, effective inter-ventions will likely become a key component of managementwherever tigers persist. Because conflicts are relatively rare inany single location, coordinating efforts and standardizing datacollection across sites and countries to allow analyses of a meta-data set will be crucial to rapid identification and implementationof effective interventions.
Acknowledgements
Funding was provided by the Wildlife Conservation Society,21st Century Tiger, The Columbus Zoo and Aquarium ConservationFund, National Geographic Society, National Fish and Wildlife
Foundation’s Save the Tiger Fund, Exxon Corporation, the CharlesEngelhard Foundation, Disney Wildlife Fund, Turner Foundation,US Fish and Wildlife Service Rhino and Tiger Conservation Fund,Richard King Mellon, Avocet Charitable Lead Unitrust, RobertsonFoundation, Starr Foundation, Goldman Environmental Founda-tion, the Homeland Fund, Denver Zoo, and numerous private do-nors. A. Khobitnev, E. Tsarapkin, B. Schleyer, N. Rybin, A. Rybin,A. Kostirya, I. Seryodkin, V. Melnikov, A. Saphonov, V. Schukin,and E. Gishko assisted with data collection. A.A. Astafiev and M.Gromyko, Y. Potikha, and Y. Pimenova of Sikhote-Alin BiosphereZapovednik provided logistical and administrative support, andthe Russian Ministry of Natural Resources provided permits forcapture.
References
Abramov, K.G., 1958. To protect the tiger. Okhota i Oknotnicie Khozyaistvo 11, 23(in Russian).
Barlow, A.C.D., 2009. The Sundarbans Tiger: Adaptation, Population Status andConflict Management. PhD Thesis. University of Minnesota, Saint Paul, USA.
Barlow, A.C.D., Greenwood, C.J., Ahmad, I.U., Smith, J.L.D., 2010. Use of an action–selection framework for human–carnivore conflict in the BangladeshSundarbans. Conservation Biology 24, 1338–1347.
Boomgaard, P., 2001. Frontiers of Fear: Tigers and People in the Malay World, 1600–1950. Yale University Press, New Haven, Connecticut, USA.
Boomgaard, P., 2010. Tigers and people in the Malay world: four centuries ofconfrontation and co-existence in comparative Asian perspective. The Malayantiger. In: Tilson, R., Nyhus, P.J. (Eds.), Tigers of the World: The Science, Politics,and Conservation of Panthera tigris. Academic Press, Burlington, Massachusetts,USA, pp. 349–356.
Breitenmoser, U., Angst, C., Landry, J., Breitenmoser-Würsten, C., Linell, J.D.C.,Weber, J., 2005. Non-lethal techniques for reducing depredation. In: Woodruffe,R., Thirgood, S., Rabinowitz, A. (Eds.), People and Wildlife: Conflict or Co-existence? Cambridge University Press, Cambridge, United Kingdom, pp. 49–71.
Chapron, G., Miquelle, D., Lambert, A., Goodrich, J., Legendre, S., Clobert, J., 2008. Theimpact of poaching versus prey depletion on tigers and other large solitaryfelids. Journal of Applied Ecology 45, 1667–1674.
Chowdhury, M.K., Sanyal, P., 1985. Man-eating behaviour of the tigers of theSundarbans, West Bengal. Environment and Ecology 3, 243–248.
Department of Wildlife and National Parks Peninsular Malaysia, 2008. NationalTiger Action Plan for Malaysia, 2008–2020. Kuala Lumpur, Malaysia.
Dinerstein, E., Loucks, C., Wikramanayake, E., et al., 2007. The fate of wild tigers.Bioscience 57, 508–514.
Goodrich, J., 2010. Human–tiger conflict: a review and call for comprehensive plans.Integrative Zoology 5, 297–308.
Goodrich, J.M., Miquelle, D.G., 2005a. Translocation of problem Amur tigers toalleviate human–tiger conflicts. Oryx 39, 1–4.
Goodrich, J.M., Miquelle, D.G., 2005b. Giving tigers a second chance: rehabilitationand translocation of problem Amur tigers to alleviate human–tiger conflicts. In:Miquelle, D.G., Smirnov, E.N., Goodrich, J.M. (Eds.), Tigers of Sikhote-AlinZapovednik: Ecology and Conservation. PSP, Vladivostok, Russia, pp. 186–190(in Russian).
Goodrich, J.M., Miquelle, D.G., 2010. Tiger telemetry. In: Tilson, R., Nyhus, P. (Eds.),Tigers of the World: The Science, Politics, and Conservation of Panthera Tigris,second ed. Elsevier Limited, Oxford, United Kingdom, pp. 261–273.
Goodrich, J.M., Kerley, L.L., Schleyer, B.O., Miquelle, D.G., Quigley, K.S., Smirnov, E.N.,Nikolaev, I.G., Quigley, H.B., Hornocker, M.G., 2001. Capture and chemicalanesthesia of Amur tigers. Wildlife Society Bulletin 29, 533–542.
Goodrich, J.M., Kerley, L.L., Smirnov, E.N., Miquelle, D.G., McDonald, L., Quigley, H.B.,Hornocker, M.G., McDonald, T., 2008. Survival rates and causes of mortality ofAmur tigers on and near the Sikhote-Alin Biosphere Zapovednik. Journal ofZoology 276, 323–329.
Goodrich, J., Lewis, J., Quigley, K., Armstrong, D., 2010a. Tiger anesthesia. In: Tilson,R., Nyhus, P. (Eds.), Tigers of the World: The Science, Politics, and Conservationof Panthera tigris, second ed. Elsevier Limited, Oxford, United Kingdom, p. 262.
Gorokhov, G.G., 1983. The Causes of Illegal Shooting of the Amur Tiger. Redkie VidyMlekopitaiuschikh ikh Okhrana. Material 3 Vsesoiuznogo Soveschania.Moscow: IEMEZH RAN I VTO AN SSSR 3, pp. 88–89. (in Russian).
Green, I.S., Woodruff, R.A., Tueller, T.T., 1984. Livestock-guarding dogs for predatorcontrol: costs, benefits and practicality. Wildlife Society Bulletin 12, 44–50.
Gurung, B., Smith, J.L.D., McDougal, C., Karki, J.B., Barlow, A., 2009. Factorsassociated with human-killing tigers in Chitwan National Park, Nepal.Biological Conservation 141, 3069–3078.
Hoogesteijn, R., Hoogesteijn, A., Mondolfi, E., 1993. Jaguar predation andconservation: cattle mortality caused by felines on three ranches in theVenezuelan Llanos. In: Dunstone, N., Gorman, M.L. (Eds.), Mammals asPredators, Symposia of the Zoological Society of London, No. 65, London, UK,pp. 391–406.
Hunter, L., Skinner, J.D., Pretorius, K., Carlisle, L.C., Rickleton, M., Walker, C., Slotow,R., 2007. Reintroducing lions Panthera leo to northern Kwazulu-Natal, SouthAfrica: short-term biological and technical successes but equivocal long-termconservation. Oryx 41, 196–204.
Inskip, C., Zimmermann, A., 2009. Human-felid conflict: a review of patterns andpriorities worldwide. Oryx 43, 18–34.
Johnson, A., Vongkhamheng, C., Hedemark, M., Saithongdam, T., 2006. Effects ofhuman–carnivore conflict on tiger (Panthera tigris) and prey populations in LaoPDR. Animal Conservation 9, 421–430.
Karanth, K.U., Gopal, R., 2005. An ecology-based policy framework for human–tigercoexistence in India. In: Woodruffe, R., Thirgood, S., Rabinowitz, A. (Eds.), Peopleand Wildlife: Conflict or Co-existence? Cambridge University Press, Cambridge,United Kingdom, pp. 373–387.
Karanth, K.U., Madhusudan, M.D., 2002. Mitigating human–wildlife conflicts inSoutheast Asia. In: Terborgh, J. (Ed.), Making Parks Work: Strategies forPreserving Tropical Nature. Island Press, Washington, DC, pp. 250–263.
Karanth, K.U., Nichols, J.D., 2002. Monitoring Tigers and Their Prey: A Manual forResearchers, Managers and Conservationists in Tropical Asia. Center for WildlifeStudies, Bangalore, India.
Kawanishi, K., Gumal, M., Soosayraj, L.A., Goldthorpe, G., Shepherd, C.R.,Krishnasamy, K., Hashimm, A.K.A., 2010. The Malayan tiger. In: Tigers of theWorld: The Science, Politics, and Conservation of Panthera Tigris. AcademicPress, Burlington, Massachusetts, USA, pp. 367–376.
Kenney, J.S., Smith, J.L.D., Starfield, A.M., McDougal, C.W., 1995. The long-termeffects of tiger poaching on population viability. Conservation Biology 9, 1127–1133.
Khan, M.M., 2009. Can domestic dogs save humans from tigers Panthera tigris? Oryx43, 44–47.
Khramtsov, V.S., 1995. Tigers’ behavior in their encounters with man. Ecologia 3,252–254 (in Russian).
Li, B., Zhang, E., Liu, Z., 2009. Livestock depredation by Amur tigers in HunchunNature Reserve, Jilin, China. Acta Theriologica Sinica 29, 231–238.
Linnell, J.D., Nilsen, E.B., Lande, U.S., 2005. Zoning as a means of mitigating conflictswith large carnivores: principles and reality. In: Woodruffe, R., Thirgood, S.,Rabinowitz, A. (Eds.), People and Wildlife: Conflict or Co-Existence? CambridgeUniversity Press, Cambridge, United Kingdom, pp. 162–175.
Loveridge, A.J., Wang, S.W., Frank, L.G., Seidensticker, J., 2010. People and wildfelids: conservation of cats and management of conflict. In: MacDonald, D.W.,Loveridge, A.J. (Eds.), Biology and Conservation of Wild Felids. Oxford UniversityPress, Oxford, UK, pp. 161–195.
Madhusudan, M.D., 2003. Living amidst large wildlife: livestock and cropdepredation by large mammals in the interior villages of Bhadra tiger reserve,South India. Environmental Management 31, 466–475.
Matyushkin, E.N., 1973. Tiger and man – the problem of living together. Prirorda 12,82–88 (in Russian).
Matyushkin, E.N., 1985. Securing the future of the tiger. Okhota I OkhotnichieKhozyaistvo 9, 8–10 (in Russian).
Matyushkin, E.N., Pikunov, D.G., Dunishenko, Y.M., Miquelle, D.G., Nikolaev, I.G.,mirnov, E.N., Salkina, G.P., Abramov, V.K., Bazilnikov, B.I., Yudin, V.G., Korkiskho,V.G., 1999. Distribution and numbers of Amur tigers in the Russian Far East inthe mid-1990’s. In: Aristova, A.A. (Ed.), Rare Mammal Species of Russia andNeighboring Territories. Russian Academy of Sciences Theological Society,Moscow, pp. 242–271 (in Russian).
McDougal, C., 1987. The man-eating tiger in geographical and historical perspective.In: Tilson, R.L., Seal, U.S. (Eds.), Tigers of the World: the Biology, Biopolitics,Management, and Conservation of an Endangered Species. Noyes, Park Ridge,New Jersey, USA, pp. 435–438.
Miquelle, D.G., Merrill, W.T., Dunishenko, Y.M., Smirnov, E.N., Quigley, H.B., Pikunov,D.G., Hornocker, M.G., 1999. A habitat protection plan for the Amur tiger:developing political and ecological criteria for a viable land-use plan. In:Seidensticker, J., Christie, S., Jackson, P. (Eds.), Riding the Tiger: TigerConservation in Human-Dominated Landscapes. Cambridge University Press,Cambridge, United Kingdom, pp. 273–295.
Miquelle, D., Nikolaev, I., Goodrich, J., Litvinov, B., Smirnov, E., Suvorov, E., 2005.Searching for the coexistence recipe: a case study of conflicts between peopleand tigers in the Russian Far East. In: Woodroffe, R., Thirgood, S., Rabinowitz, A.(Eds.), People and Wildlife: Conflict or Coexistence? Cambridge UniversityPress, Cambridge, United Kingdom, pp. 305–322.
Miquelle, D.G., Goodrich, J.M., Smirnov, E.N., Stephens, P.A., Zaumyslova, O.Yu.,Chapron, G., Kerley, L., Murzin, A.A., Hornocker, M.G., Quigley, H.B., 2010a. TheAmur tiger: a case study of living on the edge. The Amur tiger: a case study ofliving on the edge. In: Loveridge, A.J., Macdonald, D.W. (Eds.), The Biology andConservation of Wild Felids. Oxford University Press, Oxford, United Kingdom,pp. 325–339.
Miquelle, D.G., Goodrich, J.M., Kerley, L.L., Pikunov, D.G., Dunishenko, Y.M.,Aramiliev, V.V., Nikolaev, I.G., Smirnov, E.N., Salkina, G.P., Endi, Z., Seryodkin,I.V., Carroll, C., Gapanov, V.V., Fomenko, P.V., Kostyria, A.V., Murzin, A.A.,Quigley, H., Hornocker, M.G., 2010b. Science-based conservation of Amur tigersin the Russian Far East and Northeast China. In: Tilson, R., Nyhus, P. (Eds.),Tigers of the World: The Science, Politics, and Conservation of Panthera Tigris,second ed. Elsevier Limited, Oxford, United Kingdom, pp. 399–419.
Nikolaev, I.G., 1985. Last winter’s loss of tigers. Okhota i Okhotnichie Khozyaistvo 9,18–19 (in Russian).
Nikolaev, I.G., Yudin, V.G., 1993. Tiger and man in conflict situations. BulletinMoskovsky Obschestva Ispytateley Priorody, Otdyel Biologica 98, 23–26 (inRussian).
Nugraha, R.T., Sugardjito, J., 2009. Assessment and management options of human–tiger conflicts in Kerinci Seblat National Park, Sumatra, Indonesia. MammalStudy 34, 141–154.
Nyhus, P.J., Tilson, R., 2004. Characterizing human–tiger conflict in Sumatra,Indonesia: implications for conservation. Oryx 38, 68–74.
Nyhus, P.J., Tilson, R., 2010. Panthera Tigris vs. Homo sapiens: conflict, coexistence,or extinction? In: Tilson, R., Nyhus, P.J. (Eds.), Tigers of the World: The Science,Politics, and Conservation of Panthera Tigris. Academic Press, Burlington,Massachusetts, USA, pp. 125–142.
Nyhus, P.J., Caitlin, E.D., Ambrogi, M.C., Hart, S.E., Caroll, C., Tilson, R., 2010. Human–tiger conflict over time. In: Tilson, R., Nyhus, P.J. (Eds.), Tigers of the World: TheScience, Politics, and Conservation of Panthera Tigris. Academic Press,Burlington, Massachusetts, USA, pp. 132–135.
Quigley, H.Q., Herrero, S., 2005. Characterization an prevention of attacks onhumans. In: Woodroffe, R., Thirgood, S., Rabinowitz, A. (Eds.), People andWildlife: Conflict or Coexistence? Cambridge University Press, Cambridge,United Kingdom, pp. 27–48.
Quigley, K.S., Evermann, J.F., Leathers, C.W., Armstrong, D.L., Goodrich, J., Duncan,N.M., Miquelle, D.G., 2010. Morbillivirus infection confirmed in a wild Siberiantiger in the Russian Far East. Journal of Wildlife Diseases 42, 1252–1256.
Rabinowitz, A.R., 1986. Jaguar predation on domestic livestock in Belize. WildlifeSociety Bulletin 14, 170–174.
Salkina, G., 2000. Experiments in scaring away tigers in Russia. Cat News 33, 6–8.Sanderson, E., Forrest, J., Loucks, C., Ginsberg, J., Dinerstein, E., Seidensticker, J.,
Leimgruber, P., Songer, M., Heydlauff, A., O’Brien, T., Bryja, G., Klenzendorf, S.,
Wikramanayake, E., 2006. Setting Priorities for the Conservation and Recoveryof Wild Tigers: 2005–2015. The Technical Assessment. WCS, WWF,Smithsonian, NFWF-STF, Washington (DC).
Sangay, T., Vernes, K., 2008. Human–wildlife conflict in the Kingdom of Bhutan:patterns of livestock depredation by large mammalian carnivores. BiologicalConservation, 1272–1282.
Sanyal, P., 1987. Managing the maneaters of the Sundarbans Tiger Reserve of India:a case study. In: Tilson, R.L., Seal, U.S. (Eds.), Tigers of the World: The Biology,Biopolitics, Management, and Conservation of an Endangered Species. Noyes,Park Ridge, New Jersey, USA, pp. 427–434.
Shishkin, I.B., 1967. Tiger and man. Priroda 5, 99–105 (in Russian).Smith, J.L.D., McDougal, C., Gurung, B., Shrestha, N., Shrestha, M., Allendorf, T., Joshi, A.,
Dhakal, N., 2010. Securing the future for Nepal’s tigers: lessons from the past andpresent. In: Tilson, R., Nyhus, P.J. (Eds.), Tigers of the World: The Science, Politics,and Conservation of Panthera Tigris, second ed., Elsevier Limited, Oxford, UnitedKingdom, pp. 331–344.
Tilson, R., Nyhus, P., 1998. Keeping problem tigers from becoming a problemspecies. Conservation Biology 12, 261–262.
Treves, A., Karanth, K.U., 2003. Human-carnivore conflict and perspectives oncarnivore management worldwide. Conservation Biology 17, 1491–1499.
Treves, A., Naughton-Treves, L., 2005. Evaluating lethal control in the managementof human–wildlife conflict. In: Woodroffe, R., Thirgood, S., Rabinowitz, A. (Eds.),People and Wildlife: Conflict or Coexistence? Cambridge University Press,Cambridge, United Kingdom, pp. 86–106.
Walston, J., Robinson, J.G., Bennett, E.L., et al., 2010. Bringing the tiger back from thebrink – the six percent solution. PLoS Biology 8, e1000485. doi:10.1371/journal.pbio.1000485.
Wang, S.W., Macdonald, D.W., 2006. Livestock predation by carnivores in JigmeSingye Wangchuck National Park, Bhutan. Biological Conservation 129, 558–565.
Woodroffe, R., Thirgood, S., Rabinowitz, A., 2005a. The impact of human–wildlifeconflict on natural systems. In: Woodroffe, R., Thirgood, S., Rabinowitz, A. (Eds.),People and Wildlife: Conflict or Coexistence? Cambridge University Press,Cambridge, United Kingdom, pp. 1–12.
Woodroffe, R., Thirgood, S., Rabinowitz, A., 2005b. The future of coexistence:resolving human–wildlife conflicts in a changing world. In: Woodroffe, R.,Thirgood, S., Rabinowitz, A. (Eds.), People and Wildlife: Conflict or Coexistence?Cambridge University Press, Cambridge, United Kingdom, pp. 389–405.
Yudakov, A.G., Nikolaev, I.G., 1987. Ecology of the Amur Tiger. Winter Long-termObservations in the Western Part of the Middle Sikhote-Alin, 1970–1973. NaukaPress, Moscow, Russia (in Russian).
Zhivotchenko, V.E., 1976. The Amur tiger. Okhota i Okhotnichie Khozyaistvo 7, 18–21 (in Russian).
