Forum Big Cats in Borderlands: Challenges and Implications for Transboundary Conservation of Asian Leopards

Home , Amur leopard, Indochinese leopard

Forum Big cats in borderlands: challenges and implications for transboundary conservation of Asian leopards

M OHAMMAD S. FARHADINIA,SUSANA R OSTRO-GARCÍA,LIMIN F ENG J AN F. KAMLER,ANDREW S PALTON,ELENA S HEVTSOVA,IGOR K HOROZYAN M OHAMMED A L -DUAIS,JIANPING G E and D AVID W. MACDONALD

Abstract Large carnivores have extensive spatial require- for the survival of these subspecies. Recent listing of the ments, with ranges that often span geopolitical borders. leopard in the Bonn Convention on the Conservation of Consequently, management of transboundary populations Migratory Species of Wild Animals is important, but more is subject to several political jurisdictions, often with hetero- international collaboration is needed to conserve these sub- geneity in conservation challenges. In continental Asia there species. We provide a spatial framework with which range are four threatened leopard subspecies with transboundary countries and international agencies could establish trans- populations spanning  countries: the Persian Panthera boundary cooperation for conserving threatened leopards pardus saxicolor, Indochinese P. pardus delacouri, Arabian in Asia. P. pardus nimr and Amur P. pardus orientalis leopards. We Keywords Asia, Bonn Convention on the Conservation of reviewed the status of these subspecies and examined the Migratory Species of Wild Animals, borderland, conserva- challenges to, and opportunities for, their conservation. tion geopolitics, leopard, Panthera pardus, security fence, The Amur and Indochinese leopards have the majority transboundary (–%) of their remaining range in borderlands, and the Persian and Arabian leopards have –% of their remaining ranges in borderlands. Overall, in  of  countries the majority of the remaining leopard range is in borderlands, and thus in most countries conservation of these Introduction subspecies is dependent on transboundary collaboration. arge carnivores have extensive spatial requirements that However, we found only two transboundary initiatives for may extend beyond geopolitical borders. Consequently, Asian leopards. Overall, we highlighted three key trans- L these wide-ranging animals can fall under several political boundary landscapes in regions that are of high importance jurisdictions, resulting in a diversity of conservation challenges and efforts (Bischof et al., ; Linnell et al., ).

MOHAMMAD S. FARHADINIA (Corresponding author, orcid.org/0000-0002- Neighbouring states may have different levels of techni- 5385-6254) Oxford Martin School and Department of Zoology, University of cal expertise, knowledge, capacity and financial resources Oxford, 34 Broad Street, Oxford OX1 3BD, UK (Karlstetter & Mallon, ). Importantly, persistence of E-mail [email protected] large carnivore populations in one country can depend, SUSANA ROSTRO-GARCÍA,JAN F. KAMLER* and DAVID W. MACDONALD Department – of Zoology, Wildlife Conservation Research Unit, University of Oxford, Oxford, because of source sink dynamics, on populations on the UK other side of an international border (Falcucci et al., ;   LIMIN FENG and JIANPING GE Northeast Tiger and Leopard Biodiversity National Farhadinia et al., ; Feng et al., ). Conservation of Observation and Research Station, Monitoring and Research Center for Amur large carnivores requires a holistic appreciation of the Tiger and Amur Leopard, National Forestry and Grassland Administration,  Ministry of Education Key Laboratory for Biodiversity Science and geopolitical context (Hodgetts et al., ) for maintaining Engineering & College of Life Sciences, Beijing Normal University, Beijing, source populations as a priority and, where possible, enhan- China cing links to ensure persistence of metapopulations. ANDREW SPALTON Office of the Minister, Diwan of Royal Court, Muscat, Oman Although geopolitical borderlands are typically rich in ELENA SHEVTSOVA Federal Government Budgetary Institution United biodiversity, protecting these landscapes is often challen- Administration of the State Nature Biosphere Reserve Kedrovaya Pad and ging. National conservation programmes usually stop at in- Land of the Leopard National Park, Ministry of Natural Resources and Environment of the Russian Federation, Vladivostok, Russia ternational borders, but the species they aim to conserve

IGOR KHOROZYAN Georg-August-Universität Göttingen, Göttingen, Germany and the threats they strive to halt often do not (Macdonald et al., ). Borderlands are characterized by dynamic MOHAMMED AL-DUAIS Foundation Endangered Wildlife & Department of Biology, Ibb University, Ibb, Republic of Yemen social, political, economic and sometimes even ecological *Also at: Panthera, New York, USA transitions that, at extremes, may involve armed conflict  Received  February . Revision requested  April . and political instability (McNeely, ). Man-made bar- Accepted  June . First published online  June . riers along geopolitical borders, intended to control

This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, Downloaded fromdistribution, https://www.cambridge.org/core and reproduction in any medium,. IP address: provided 170.106.202.8 the original, work on 27 is Sep properly 2021 cited. at 09:20:50, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/termsOryx, 2021, 55(3),. https://doi.org/10.1017/S0030605319000693 452–460 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000693 Big cats in borderlands 453

movement of people, can create an impediment for the extant) overlapping with borderlands, and length of border movement of large carnivores (Karlstetter & Mallon, ; lines, were calculated using QGIS .. (QGIS Development Linnell et al., ; Trouwborst et al., ). Expansion of Team, ). We assigned a range patch as transboundary if border security barriers is recognized as a threat to wildlife it overlapped with borderlands, which we defined as a buffer because they can cause mortality, obstruct access to season- zone of  km from the borderline. We chose this size be- ally important resources, and reduce effective population cause it is the maximum dispersal distance for leopards size and viability (Linnell et al., ). Range shifts across in Asia, recorded for the Persian subspecies using satellite political borders induced by global climate change are telemetry in north-east Iran (Farhadinia et al., ). We also potential challenges for many species (Hannah, ). acknowledge that this buffer is an approximation, but there These challenges can add to the already precarious circum- are no similar data for the other Asian subspecies. stances of many large carnivores, which often occur at low densities and are prone to demographic and environmental stochasticity. Transboundary ranges Although its broad geographical range and charismatic There are six leopard subspecies across continental Asia appeal make the leopard Panthera pardus one of the most (Jacobson et al., ; Stein et al., ). We focus on those potent mammalian ambassador species, it is experiencing subspecies that have transboundary populations and are a greater range loss than most other large terrestrial carni- categorized, or are being petitioned to be categorized, as En-  vores (Wolf & Ripple, ). In Asia, leopard subspecies cur- dangered or Critically Endangered on the IUCN Red List of ,  rently occur in % of their historical range, with the only Threatened Species: the Persian P. pardus saxicolor (Endan- – relatively large populations ( individuals) occur- gered), Indochinese P. pardus delacouri (Critically Endan-  ring in India, Sri Lanka and Iran (Jacobson et al., ; gered), Arabian P. pardus nimr (Critically Endangered), and  Stein et al., ). Persistence of many small populations Amur P. pardus orientalis (Critically Endangered) leopards – of leopards is dependent on source sink dynamics across (Table , Plate ). international borders (Khorozyan et al., ; Farhadinia et al., ; Rostro-García et al., ; Feng et al., ; Maharramova et al., ; Askerov et al., ). Persian leopard Various international legal instruments exist that have  a direct or indirect effect on the conservation of large The Persian leopard occurs across the rugged terrain of carnivores, including the  Bonn Convention on the countries in western Asia and the Caucasus, with a popula- –   .  Conservation of Migratory Species of Wild Animals tion of , individuals (Khorozyan, ) and % ’ (Karlstetter & Mallon, ; Trouwborst et al., ). The of the subspecies extant range within Iran (Jacobson et al.,   ’ Convention has evolved into an instrument focusing on ). In countries the Persian leopards range occurs   the conservation of migratory wildlife crossing international exclusively in borderlands (Fig. , Table ), with small pop- ,   borders (Trouwborst et al., ). Recent listing of the leop- ulations of generally individuals (Askerov et al., ;  ard in Appendix II of the Convention is a step towards ini- Avgan et al., ). These appear to be sink populations  tiating global or regional agreements for the conservation on the brink of extinction (Askerov et al., ; Avgan    and management of this species. However, it is not yet et al., ; Stein et al., ; Maharramova et al., ), clear what regional and global challenges are faced by leop- although under intensive conservation efforts they are able  ard conservation in Asia and what opportunities exist for to recolonize suitable habitats (Askerov et al., ). overcoming them. Here, we highlight the importance of implementing Indochinese leopard transnational strategies for the conservation of leopards that range across extensive areas in continental Asia. We The Indochinese leopard is probably extant in only –%of focus on the conservation status and challenges of trans- its historical distribution in South-east Asia, and the remain- boundary populations of threatened leopard subspecies, ing populations are small and isolated (Rostro-García et al., and identify initiatives with which conservation practi- ). This subspecies has been extirpated in Singapore, tioners can facilitate effective transboundary cooperation probably extirpated in Laos and Viet Nam, nearly extirpa- for the conservation of leopards, and perhaps also other ted in Cambodia and has a greatly reduced distribution in large mammals. China, Thailand, Myanmar and Peninsular Malaysia (Rostro- We reviewed the available literature to collate a dataset of García et al., ). Of the remaining extant and possibly the population size and distribution of each threatened sub- extant distribution, % lies within borderlands (Table ). species in Asia (Jacobson et al., ; Stein et al., ), and With the exception of Malaysia, most of the remaining the legal status of the leopard in each country. The spatial leopard range within each country is in borderlands metrics, specifically leopard range (extant and possibly (Fig. , Table ).

Oryx, 2021, 55(3), 452–460 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000693 Downloaded from https://www.cambridge.org/core. IP address: 170.106.202.8, on 27 Sep 2021 at 09:20:50, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319000693 454 M. S. Farhadinia et al.

TABLE 1 Details of an Endangered and three Critically Endangered leopard Panthera pardus subspecies with populations in borderlands of continental Asia.

No. of extant Borderline Total area of subspecies Borderland area, km2 (% of Subspecies range countries length (km) range (km2) subspecies overall range) Endangered Persian P. pardus saxicolor 13 3,415 933,597 247,035 (26) Critically Endangered Indochinese P. pardus delacouri 5 1,740 159,000 92,220 (58) Arabian P. pardus nimr 4 47 22,720 5,332 (23) Amur leopard P. pardus orientalis 3 248 9,270 9,270 (100) Total 231 5,450 1,124,587 353,857 (31)

 There are two leopard subspecies in both China and Russia, and therefore the total number of countries is  rather than .

Arabian leopard is increasing in China, and some reproduction has been documented  km from the Russian border (Wang et al., There are small populations of the Arabian leopard in ). Oman, Saudi Arabia, Israel, and Yemen (Fig. , Table ). Until relatively recently, this subspecies also occurred in Discussion Jordan and the United Arab Emirates, in both of which the populations would have been partly transbound- The importance of borderlands for leopard conservation ary with Israel and Oman, respectively (Jacobson et al.,  ). The main stronghold for the subspecies is the A relatively high proportion (–%) of the ranges of each Dhofar Mountains of Oman, which comprise three con- leopard subspecies occurs in borderlands. The Amur and tiguous mountain blocks: Jabal Samhan in the east, Jabal Indochinese leopards are most dependent on borderlands Al Qara in the centre and Jabal Al Qamar in the west, the as the majority of their remaining distribution occurs within latter contiguous with the mountains of Hawf in south- these areas. The Persian and Arabian leopards have the ma- east Yemen. Occasional records of leopard mortality jority of their remaining distribution beyond borderlands as indicate some small populations across the inner parts most of their populations are within Iran and Afghanistan  of Yemen. The largest population nucleus of up to for the Persian leopard, and Oman and possibly Yemen Arabian leopards is believed to inhabit Jabal Samhan, for the Arabian leopard.  – which is c. km from the Oman Yemen border. A fur- Transboundary conservation initiatives have been en- – ther leopards occur across Jabal Al Qara and Jabal dorsed for many of Asian leopard subspecies (Knight  Al Qamar, with individuals recorded up to km from et al., ; Askerov et al., ; Farhadinia et al., ; –  the Oman Yemen border (Spalton & Al Hikmani, ). Feng et al., ). However, only two transboundary initia- – Across the Oman Yemen border in Hawf, leopards were tives are actively working to facilitate leopard conservation   recorded in low numbers in (Khorozyan et al., ). across borders, in the Caucasus (Askerov et al., ) and in Elsewhere, there are thought to be a few scattered areas the Russian Far East and north-east China (Feng et al., ; of extant range for the Arabian leopard along the border Vitkalova et al., ). Transboundary conservation was between Yemen and Saudi Arabia (Zafar-ul Islam et al., not considered in the latest IUCN assessment of leopards  ). (Stein et al., ). Our review highlights the importance of encouraging conservation agencies to work across international borders in Asia. Amur leopard  There is only one population of the Amur leopard (Fig. ) Conservation challenges in borderlands shared between Russia’s Primorskii Krai and China’s Jilin Province (Jacobson et al., ; Feng et al., ). There Previous studies have highlighted poaching of leopards and are also occasional reports, based on snow tracking, from their prey, and habitat loss, as the main reasons for the decline North Korea (Jacobson et al., ). With no evidence of of leopards across most of their range in Asia (Farhadinia Amur leopards occurring elsewhere, this single transbound- et al., ; Jacobson et al., ; Rostro-García et al., ; ary population of – individuals represents the global Wang et al., ; Zafar-ul Islam et al., ). We identify population of this subspecies (Vitkalova et al., ). After four main challenges for the conservation of transboundary concerted conservation effort, the Amur leopard population populations of Asian leopards: () different levels of legal

PLATE 1 Photographic evidence of Asian leopards in borderlands: (a) Persian leopard P. pardus saxicolor with amputated leg along the Armenia–Azerbaijan–Iran border in the Caucasus (Photo: WWF), (b) an Arabian leopard P. pardus nimr in the Hawf, on the Yemen–Oman border (Photo: Foundation for Endangered Wildlife Yemen), (c) a Persian leopard in north-east Iran, with Turkmenistan’s mountains in the background (Photo: WildCRU/ FutureLeopards Foundation), (d) a Persian leopard after killing two domestic goats along the Iran–Iraq border (Photo: Iran Department of Environment/R. Khoshfarman), (e, f) Indochinese leopards P. pardus delacouri near the Cambodia– Viet Nam border (Photo: Panthera/ WildCRU/WWF Cambodia/Ministry of Environment), and Amur leopards P. pardus orientalis along the Russia–China border in (g) China (Photo: Beijing Normal University) and (h) Russia (Photo: FGBU/ Land of the Leopard).

protection and management across national jurisdictions, Kazakhstan and Afghanistan (Persian leopard) and Cambo- () military activities and armed conflict, ()poachingfor dia, China, Myanmar and Laos (Indochinese leopard) have illegal international wildlife trade and () infrastructure pro- not prioritized or supported conservation action plans for jects, such as road development. the leopard, which hampers conservation efforts made by There are varied levels of legal protection and management neighbouring states. In Oman, conservation of the Arabian for leopards across national jurisdictions and protection by law leopard benefits from strong government agencies, law en- across most range countries, with substantial monetary fines forcement and compensation programmes in cases of live- and/or imprisonment for illegal killing (Table ). However, stock depredation. In adjacent Yemen, however, where there there were time lags of several decades for legal protection in is an ongoing armed conflict, there is little or no active conser- the adjacent countries in several large borderland populations, vation or protection of leopards. such as Thailand–Myanmar and Iran–Iraq–Turkey (Table ). Military activities and armed conflicts occur within the Neighbouring countries may have different agendas, technical borderland ranges of all leopard subspecies except for the capacities, and resources available for leopard conservation, Amur leopard. Political unrest compromises law enforce- potentially hindering the recovery of transboundary pop- ment and effective conservation. Potential effects of mili- ulations. For example, the governments of Turkmenistan, tary activities and armed conflicts on leopards and the

FIG. 1 The current range of the Persian Panthera pardus saxicolor, Arabian P. pardus nimr, Indochinese P. pardus delacouri and Amur P. pardus orientalis leopard subspecies in Asia, and the locations of borderlands.

persistence of their prey require further research. The prin- leopard, are active in Harad and Al Khoba in Yemen and cipal conflicts currently affecting leopards are those between Saudi Arabia, respectively. Yemen and Saudi Arabia, the Nagorno-Karabakh conflict Existing and proposed infrastructure projects can have between Armenia and Azerbaijan, and political instabilities detrimental effects on leopards and other wildlife. For along the borders between northern Iraq, western Iran example, road development, which has increased in Asia, and Turkey, and between Afghanistan and Pakistan. In has direct negative impacts on wildlife, particularly large- Myanmar, the long-running Karen insurgency near the bodied mammals such as leopards (Clements et al., ). border with Thailand could be negatively affecting a local Amongst other impacts, roads can promote illegal wildlife leopard population (Oswell, ), although recent surveys trade, impede animal movement, and be precursors of conducted in Karen State found a high diversity of spe- overhunting and habitat destruction. In Myanmar, a major cies, including leopards (Moo et al., ). Other non- source of illegal wildlife, road networks have facilitated il- government controlled areas in northern Myanmar, along legal trade of mammals to border markets (Oswell, ). the borders with Thailand, Laos, China and India, play a Proposed major infrastructure projects are expected to major role in illicit activities, including poaching and facil- cause fragmentation of key landscapes, such as the Dawei itating the illegal trade of leopards and other wild felids Special Economic Zone road in the Northern Tenassarim (Oswell, ). Furthermore, mine-strewn borderlands oc- Forest Complex (Helsingen et al., ), one of the last strong- casionally kill leopards and other wildlife, including areas holds for the Indochinese leopard (Rostro-García et al., ) along the Thailand–Myanmar, Iran–Iraq and Armenia– as well as Kopet Dag ecoregion along the Iran–Turkmenistan Azerbaijan borders (Oswell, ; Avgan et al., ;IUCN border for the Persian leopard (Farhadinia et al., ). SSC Cat Specialist Group, ). Because development projects are increasing throughout Illegal wildlife trade, particularly intensive and indis- many areas of Asia, it is imperative that measures are taken criminate snaring over large areas, is the main cause of to minimize the impact that such projects have on leopards the dramatic decline of the Indochinese leopard (Rostro- and other wildlife. García et al., , ). This is particularly problem- Border security fencing and associated roads are concerns atic in borderlands because poachers may originate from for transboundary movement of wildlife in many parts of a neighbouring country (O’Kelly et al., ) and return Eurasia (Linnell et al., ). Although barbed wire is com- across the border before being apprehended. In general, mon in some range country borderlands, such as parts of weak enforcement of wildlife laws and lax border security the Caucasus region, it is unlikely to act as a barrier for have facilitated the illegal trade of wildlife products in leopard movement, but more impassable border fences South-east Asia (Oswell, ). Two border markets for may impede movements of leopards and their prey along the trafficking of exotic animals, including the Arabian the Iran–Turkmenistan and Yemen–Oman borders (Fig. ).

TABLE 2 Populations and legal status of the four threatened leopard subspecies across continental Asia.

Extant Year national pro- Population % country range Reference for popula- countries tection granted Fine for illegal killing (USD) size in borderlands tion size Persian leopard Afghanistan 2008 None 200–300 18 Khorozyan (2008) Armenia 1972 6,100 (outside protected area), ,10 100 Askerov et al. (2015) 30,500 (inside protected area)2 Azerbaijan 1976 1,950 (outside protected area), ,10 100 Askerov et al. (2015) 5,820 (inside protected area) Georgia 1982 19,000 ,3 100 Askerov et al. (2015) Iran 1965 9,200 550–850 28 Kiabi et al. (2002) Iraq 2010 8,350 ,10 100 Avgan et al. (2016) Kazakhstan 2004 9,220 Not known 100 Pakistan Not known Not known Not known 75 Russia 1956 2–9 years in prison plus a fine ,10 100 Khorozyan (2008) up to 45,700 Tajikistan 2008 424–25,000 Not known 100 Turkey 2003 13,600 ,10 100 Avgan et al. (2016) Turkmenistan 1970s 600 (outside protected area), ,60 91 O. Pereladova, pers. 1,700 (inside protected area) comm. (2020) Uzbekistan 1983 7,300 (for Uzbek citizens), Not known 100 Not known 40,000 (for foreign citizens) Indochinese leopard Cambodia 1994 Up to 24,500 18–178 100 Modified from Rostro-García et al. (2019) Malaysia 1972 24,000–120,000, depending 106–1,059 26 Modified from on age/sex of leopard Rostro-García et al. (2019) Myanmar 1994 Up to 7,680 169–1,693 59 Modified from Rostro-García et al. (2019) SE China 1988 .10 years in prison plus a Not known 100 fine (1,600) or confiscation of property Thailand 1960 Up to 1,200 59–591 98 Modified from Rostro-García et al. (2019) Arabian leopard Israel Not known Not known 8 100 Perez et al. (2006) Oman 1976 Up to 12,500 44–58 25 Spalton & Al Hikmani (2014) Saudi Arabia 1999 5,000 (outside protected area), ,50 6 Zafar-ul Islam et al. 13,000 (inside protected area) (2018) Yemen 2009 Not known Not known 37 Amur leopard1 NE China 1988 .10 years in prison plus a 70–108 100 Vitkalova et al. (2018) fine (1,600) or confiscation of property North Korea Not known Not known Not known 100 Russia 1956 2–9 years in prison plus a fine 70–108 100 Vitkalova et al. (2018) up to 45,700

 Population size is the total number across Russia and China.  An increase up to USD , is currently under consideration.

FIG. 2 Locations (from GPS fixes) of a collared Persian leopard that dispersed from Tandoureh National Park in north-east Iran to Turkmenistan (Farhadinia et al., ). The locations show that although the leopard moved freely across the international border, the security fence, which lies further north within Turkmenistan, was a barrier to the leopard’s movements.

Most borderland leopard populations in Asia occur along leopard. Despite the civil strife in Yemen, the border gover- unfenced borders, and thus transboundary movements are norate of Al Mahrah in Yemen adjoining Oman’s Dhofar still feasible. Mountains remainsless affected. For leopards that move freely across international Conservation paradigms need to be realigned with po- borders in Asia, the same individuals may be counted litical reality when examining geopolitical situations for in more than one country, inflating abundance estimates conservation (Linnell et al., ). Some Asian countries (Bischof et al., ; Vitkalova et al., ). This may be a are facing security challenges from neighbouring coun- problem in the Caucasus and along the Russia–China bor- tries, which could reduce opportunities for transboundary der (Vitkalova et al., ; Askerov et al., ). This empha- cooperation. Given that the majority of borderland leopard sizes the need for the establishment of internationally man- populations occur along unfenced international borders, aged monitoring programmes and for sharing monitoring conflicting countries can unilaterally enforce the conser- information. Transboundary information exchange can vation of their transboundary populations and shift their improve the accuracy and precision of population estimates, conservation investments towards the borderland. An ex- which can lead to a better understanding of the status of ample is leopard conservation in Armenia and Azerbaijan’s leopard populations. Nakhichevan Republic which, despite a political dispute, has succeeded in maintaining protected areas for leopards and Implications for policy supporting population recovery on both sides of the shared border (Askerov et al., ). Transboundary conservation areas are widely recommended In addition, effects of border fences on leopard move- as a means to encourage multilateral intergovernmental part- ments and demography need to be better understood. If nerships. The concept of international Peace Parks is being planned without compromising national security, joint promoted as a way of linking biodiversity conservation with population monitoring (Feng et al., ; Vitkalova et al., national security (McNeely, ), and such a park has been ) and satellite telemetry (Farhadinia et al., ) can established between Arevik National Park in Armenia and help elucidate the locations of corridors and source–sink Dizmar Protected Area in Iran, where leopards occur. In ad- dynamics across international borders. Citizen science moni- dition to the two existing transboundary leopard initiatives toring schemes supported by border security personnel in the Caucasus and Russia–China borderlands (Vitkalova could provide coarse-scale data to identify key corridors. et al., ; Askerov et al., ), we identified three key trans- There are two intergovernmental organizations, the boundary landscapes: () Kopet Dag ecoregion along the Association of Southeast Asian Nations (ASEAN) and Iran–Turkmenistan border for the Persian leopard, although Economic Cooperation Organization (ECO), of which some parts contain impassable fences (Fig. ), ()Northern most of the regional countries within the ranges of the Tenasserim Forest Complex along the Thailand–Myanmar Indochinese and Persian leopard, respectively, are mem- border for the Indochinese leopard, one of WWF’sfiveprior- bers. The ASEAN Wildlife Enforcement Network and ECO’s ity global landscapes (WWF, ), and ()Hawf–Dhofar Division on Social Welfare and Environment could pro- Mountains along the Yemen–Oman border for the Arabian vide a framework for the establishment of transboundary

cooperation for leopard conservation and the suppression of Initiative may create new supplies for illegal wildlife trade in illegal border trade. Given that the majority of the remaining large carnivores. Nature Ecology & Evolution, , –. leopard range in most countries is in borderlands, trans- FENG, L., SHEVTSOVA, E., VITKALOVA, A., MATYUKHINA, D., MIQUELLE, D., ARAMILEV, V. et al. () Collaboration brings boundary collaborations would foster the persistence of hope for the last Amur leopards. Cat News, , . small but important populations of Asian leopards. HANNAH,L.() A global conservation system for climate-change adaptation. Conservation Biology, , –. Acknowledgements We thank S. Karryeva, E. Askerov, H. Raza, HELSINGEN, H., MYINT, S.N.W., BHAGABATI, N., DIXON, A., O. Pereladova, B. Lortkipanidze, Y. Ilemin, T. Rosen, M. Kabir, OLWERO, N., KELLY, A.S. & TANG,D.() A better road to Dawei: N. Jahed, M. Gritsina and A. Gasparyan for contributing data on the protecting wildlife, sustaining nature, benefitting people. World legal status of the Persian leopard, Z. Islam for data on the Arabian Wildlife Fund, Yangon, Myanmar. leopard, and G.R. Clements, A.J. Lynam, and H. Naing for help with HODGETTS, T., BURNHAM, D., DICKMAN, A., MACDONALD, E.A. & collecting data on the Indochinese leopard. MSF was supported by a MACDONALD, D.W. () Conservation geopolitics. Conservation research fellowship from the Oxford Martin School at the University Biology, , –. of Oxford. IUCN SSC CAT SPECIALIST GROUP () Foot-amputated leopards in the Caucasus. Facebook,  May . facebook.com/ Author contributions Study design: MSF, DWM, data collection /photos/a./ and analysis: MSF, SR-G, LF, JFK, AS, ES, MA-D; writing: all authors. [accessed May ]. JACOBSON, A.P., GERNGROSS, P., LEMERIS, J.R., SCHOONOVER, R.F., Conflicts of interest None. ANCO, C., BREITENMOSER-WÜRSTEN, C. et al. () Leopard (Panthera pardus) status, distribution, and the research efforts Ethical standards This research abided by the Oryx guidelines on across its range. PeerJ, ,e. ethical standards. KARLSTETTER,M.&MALLON,D.() Assessment of gaps and needs in migratory mammal conservation in Central Asia. In A Report for the Convention on the Conservation of Migratory Species of Wild References Animals (CMS) and the Deutsche Gesellschaft für Internationale Zusammenarbeit (GIZ). United Nations Environment Programme/ ASKEROV, E., TALIBOV, T., MANVELYAN, K., ZAZANASHVILI, N., Convention on Migratory Species Secretariat, Bonn, Germany. MALKHASYAN, A., FATULLAYEV,P.&HEIDELBERG,A.() KHOROZYAN,I.() Panthera pardus ssp. saxicolor.InThe IUCN South-Eastern Lesser Caucasus: the most important landscape Red List of Threatened Species. Version .: e.TA for conserving the Leopard (Panthera pardus) in the Caucasus iucnredlist.org/species// [accessed  March ]. region (Mammalia: Felidae). Zoology in the Middle East, KHOROZYAN, I., STANTON, D., MOHAMMED, M., AL-RA’IL,W.& , –. PITTET,M.() Patterns of co-existence between humans and ASKEROV, E., TALIBOV, T., MANVELYAN, K., ZAZANASHVILI, N., mammals in Yemen: some species thrive while others are nearly FATULLAYEV,P.&MALKHASYAN,A.() Leopard (Panthera extinct. Biodiversity and Conservation, , –. pardus) reoccupying its historic range in the South Caucasus: a first KIABI, B., DARESHOURI, B., GHAEMI,R.&JAHANSHAHI,M.() evidence (Mammalia: Felidae). Zoology in the Middle East, Population status of the Persian Leopard (Panthera pardus saxicolor , –. Podock, ) in Iran. Zoology in the Middle East, , –. AVGAN, B., RAZA, H., BARZANI,M.&BREITENMOSER,U.() KNIGHT, M.H., SEDDON, P.J. & AL MIDFA,A.() Transboundary Do recent leopard Panthera pardus records from northern Iraq conservation initiatives and opportunities in the Arabian Peninsula. and south-eastern Turkey reveal an unknown population nucleus Zoology in the Middle East, , –. in the region? Zoology in the Middle East, , –. LINNELL, J.D.C., TROUWBORST, A., BOITANI, L., KACZENSKY,P., BISCHOF, R., BRØSETH,H.&GIMENEZ,O.() Wildlife in a HUBER, D., RELJIC, S. et al. () Border security fencing and politically divided world: insularism inflates estimates of brown wildlife: the end of the transboundary paradigm in Eurasia? bear abundance. Conservation Letters, , –. PLOS Biology, ,e. CLEMENTS, G.R., LYNAM, A.J., GAVEAU, D., YAP, W.L., LHOTA, S., MACDONALD, D.W., JOHNSON, D.D.P. & WHITEHOUSE,H.() GOOSEM, M. et al. () Where and how are roads endangering Towards a more natural governance of Earth’s biodiversity and mammals in Southeast Asia’s forests? PLOS ONE, ,e. resources. Conservation & Society, , –. FALCUCCI, A., MAIORANO, L., TEMPIO, G., BOITANI,L.&CIUCCI,P. MAHARRAMOVA, E., MOQANAKI, E.M., ASKEROV, E., FAEZI, S., () Modeling the potential distribution for a range-expanding ALINEZHAD, H., MOUSAVI, M. et al. () Transboundary leopard species: wolf recolonization of the Alpine range. Biological movement between Azerbaijan and Iran in the Southern Caucasus. Conservation, , –. Cat News, , –. FARHADINIA, M.S., AHMADI, M., SHARBAFI, E., KHOSRAVI, S., MCNEELY, J.A. () Conserving forest biodiversity in times of ALINEZHAD,H.&MACDONALD, D.W. () Leveraging violent conflict. Oryx, , –. trans-boundary conservation partnerships: persistence of Persian MOO, S.S.B., FROESE, G.Z.L. & GRAY, T.N.E. () First structured leopard (Panthera pardus saxicolor) in the Iranian Caucasus. camera-trap surveys in Karen State, Myanmar, reveal high diversity Biological Conservation, , –. of globally threatened mammals. Oryx, , –. FARHADINIA, M.S., JOHNSON, P.J., MACDONALD, D.W. & HUNTER, O’KELLY, H.J., ROWCLIFFE, J.M., DURANT, S.M. & MILNER- L.T.B. () Anchoring and adjusting amidst humans: ranging GULLAND, E.J. () Robust estimation of snare prevalence behavior of Persian leopards along the Iran–Turkmenistan within a tropical forest context using N-mixture models. Biological borderland. PLOS ONE, ,e. Conservation, , –. FARHADINIA, M.S., MAHESHWARI, A., NAWAZ, M.A., AMBARLI, H., OSWELL, A.H. () The Big Cat Trade in Myanmar and Thailand. GRITSINA, M.A., KOSHKIN, M.A. et al. () Belt and Road TRAFFIC Southeast Asia, Malaysia.

PEREZ, I., GEFFEN,E.&MOKADY,O.() Critically Endangered TROUWBORST, A., BLACKMORE, A., BOITANI, L., BOWMAN, M., Arabian leopards Panthera pardus nimr in Israel: estimating CADDELL, R., CHAPRON, G. et al. () International wildlife law: population parameters using molecular scatology. Oryx, , –. understanding and enhancing its role in conservation. BioScience, QGIS DEVELOPMENT TEAM () qgis.org [accessed  June ]. , –. ROSTRO-GARCÍA, S., KAMLER, J.F., ASH, E., CLEMENTS, G.R., VITKALOVA, A.V., FENG, L., RYBIN, A.N., GERBER, B.D., MIQUELLE, GIBSON, L., LYNAM, A.J. et al. () Endangered leopards: range D.G., WANG, T. et al. () Transboundary cooperation improves collapse of the Indochinese leopard (Panthera pardus delacouri) endangered species monitoring and conservation actions: a case in Southeast Asia. Biological Conservation, , –. study of the global population of Amur leopards. Conservation ROSTRO-GARCÍA, S., KAMLER, J.F., CLEMENTS, G.R., LYNAM, A.J. & Letters, ,e. NAING,H.() Panthera pardus ssp. delacouri.InThe IUCN Red WANG, T., FENG, L., MOU, P., WU, J., SMITH, J.L.D., XIAO, W. et al. List of Threatened Species : e.TA. dx.doi.org/ () Amur tigers and leopards returning to China: direct evidence ./IUCN.UK.-.RLTS.TA.en [accessed and a landscape conservation plan. Landscape Ecology, , –.  December ]. WOLF,C.&RIPPLE, W.J. () Range contractions of the world’s SPALTON, J.A. & AL HIKMANI, H.M. () Arabian Leopards of large carnivores. Royal Society Open Science, , . Oman. Stacey International, London, UK. WWF () The Dawna-Tenasserim Landscape. World Wildlife STEIN, A.B., ATHREYA, V., GERNGROSS, P., BALME, G., HENSCHEL,P., Fund. greatermekong.panda.org/discovering_the_greater_mekong/ KARANTH, U. et al. () Panthera pardus (errata version landscapes/dawna_tenasserim_landscape [accessed  May ]. published in ). In The IUCN Red List of Threatened Species : ZAFAR-UL ISLAM, M., BOUG, A., JUDAS,J.&AS-SHEHRI,A.() e.TA. dx.doi.org/./IUCN.UK.-.RLTS. Conservation challenges for the Arabian leopard (Panthera pardus TA.en [accessed  July ]. nimr) in Western Highlands of Arabia. Biodiversity, , –.