The Javan Leopard Panthera pardus melas (Cuvier, 1809) (Mammalia: Carnivora: Felidae) in West Java, Indonesia: estimating population density and occupancy

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Anton Ario
Senjaya Mercusiana
Ayi Rustiadi
Robi Gumilang
I Gede Gelgel Darma Putra Wirawan
Toni Ahmad Slamet


The Javan Leopard is endemic to the Indonesian island of Java and has been classified as Endangered. Reliable information about its population status, distribution, and density is lacking but are essential to guide conservation efforts and provide a benchmark for management decisions. Our study represents the first empirical density and occupancy estimates for the Leopard in West Java and provides baseline data for this region. We used camera trap data collected from February 2009 to October 2018 in six study areas comprising a sampling effort of 10,955 camera trap days in a total area of 793.5 km2. We identified 55 individual Leopards in these areas and estimated Leopard density using spatially explicit capture-recapture. Population density estimates range from 4.9 individuals/100 km2 in Gunung Guntur-Papandayan Nature Reserve to 16.04 individuals/100 km2 in Gunung Gede Pangrango National Park. Latter is among the globally highest Leopard densities. Based on detection data, we modelled single-season Leopard occupancy using three sampling covariates and eight site covariates. Modelling revealed that the two covariates forest cover and presence of Wild Boar are the strongest predictors for Leopard occupancy in our study areas. We recommend assessing and monitoring Leopard distribution, density and occupancy in other areas of Java and emphasize that a landscape approach for conservation of the Javan Leopard is imperative.

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Adhiasto, D.N., E. Wilianto & H.T. Wibisono (2020). Uncover the unrevealed data: the magnitude of Javan Leopard removal from the wild. Cat News 71: 5–6.

Ario, A., A.P. Kartono, L.B. Prasetyo & J. Supriatna (2018a). Habitat suitability of release site for Javan Gibbon (Hylobates moloch) in Mount Malabar Protected Forests, West Java. Jurnal Manajemen Hutan Tropika 24(2): 92–101. DOI:

Ario, A., Supian, E. Hidayat, R. Hidayatullah, A. Rustiadi, A. Gunawan, T. Triprajawan, I. Sopian, R. Zatnika, D. Yusup, W. Hindrayani, T. Pramesti, A. Mulyanto & D. Iskandar (2018b). Population dynamics and ecology of Javan Leopard, Panthera pardus melas, in Gunung Gede Pangrango National Park, West Java. Journal of Indonesian Natural History 6(1): 6–13.

Athreya, V., A. Srivathsa, M. Puri, K.K. Karanth, N.S. Kumar, K.U. Karanth (2015). Spotted in the news: using media reports to examine Leopard distribution, depredation, and management practices outside protected areas in southern India. PLoS One 10(11): e0142647. DOI:

Badan Informasi Geospasial (2013). Ina-Geoportal. Pusat Pengelolaan dan Penyebarluasan Informasi Geospasial [Center for Geospatial Information Management and Dissemination], Bogor, Indonesia. Electronic version at accessed on 10 December 2020.

Badan Pusat Statistik Provinsi Jawa Barat (2021). Provinsi Jawa Barat dalam angka. Badan Pusat Statistik Provinsi Jawa Barat. Bandung, 703 pp.

Badan Pusat Statistik (2020). Statistik Indonesia 2020. Badan Pusat Statistik Indonesia, Jakarta, 748 pp.

Balme, G.A., L. Hunter & A.R. Braczkowski (2012). Applicability of age-based hunting regulations for African Leopards. PloS One 7(4): e35209. DOI:

Bogie, I., Y.I. Kusumah & M.C. Wisnandary (2008). Overview of the Wayang Windu geothermal field, West Java, Indonesia. Geothermics 37(3): 347–365. DOI:

Borah, J., T. Sharma, D. Das, N. Rabha, N.K.A. Basumatary, M.F. Ahmed & J. Vattakaven (2014). Abundance and density estimates for Common Leopard Panthera pardus and Clouded Leopard Neofelis nebulosa in Manas National Park, Assam, India. Oryx 48(1): 149–155. DOI:

Boron, V., J. Tzanopoulos, J. Gallo, J. Barragan, L. Jaimes-Rodriguez, G. Schaller & E. Payán (2016). Jaguar densities across human-dominated landscapes in Colombia: the contribution of unprotected areas to long term conservation. PloS One 11(5): e0153973. DOI:

Burnham, K.P & D.R. Anderson (2002). Model Selection and Multimodel Inference: A Practical Information-theoretic Approach. Springer, New York, 488 pp.

Carranza, E.J.M., H. Wibowo, S.D. Barritt & P. Sumintadireja (2008). Spatial data analysis and integration for regional-scale geothermal potential mapping, West Java, Indonesia. Geothermics 37(3): 267–299. DOI:

Dormann, C.F., J. Elith., S. Bacher., C. Buchmann., G. Carl., G. Carre., J.R.G. Marquez., B. Gruber., B. Lafourcade., P.J. Leitao., T. Munkemuller., C. McClean., P.E. Osborne., B. Reineking., B. Schroder., A.K. Skidmore., D. Zurell & S. Lautenbach. (2013). Collinearity: a review of methods to deal with it and a simulation study evaluating their performance. Ecography 36: 27–46. DOI:

Dsikowitzky, L., A. Damar, S.C.A. Ferse, H.E. Irianto, T.C. Jennerjahn, M.C. Lukas, I. Nordhaus, T. Pohlmann, J. Schwarzbauer, K. Sugama & B. Sumiono (2019). Chapter 21 - Java Island, Indonesia, pp. 459–490. In: Sheppard, C. (ed). World Seas: An Environmental Evaluation (Second Edition), Volume II: the Indian Ocean to the Pacific. United Kingdom, 912 pp. DOI:

Efford, M.G. (2015). Spatially explicit capture-recapture models. R package version 2.9.5.

Efford, M.G. (2018). A tutorial on fitting spatially explicit capture–recapture models in secr. R package version 3.1.5. Downloaded at

Efford, M.G., D.K. Dawson & D.L. Borchers (2009). Population density estimated from locations of individuals on a passive detector array. Ecology 90(10): 2676–2682. DOI:

Erfanian, B., S.H. Mirkarimi, A.S. Mahini & H.R. Rezaei (2013). A presence-only habitat suitability model for Persian Leopard Panthera pardus saxicolor in Golestan National Park, Iran. Wildlife Biology 19(2): 170–178. DOI:

Farhadinia, M.S., B.T. McClintock, P.J. Johnson, P. Behnoud, K. Hobeali, P. Moghadas, L.T.B. Hunter & D.W. Macdonald (2019). A paradox of local abundance amidst regional rarity: the value of montane refugia for Persian Leopard conservation. Science Report 9: 14622. DOI:

Foster, R.J. & B.J. Harmsen (2012). A critique of density estimation from camera-trap data. The Journal of Wildlife Management 76(2): 224–236. DOI:

Giri, M.S. & A. Munawir (2021). Rumah kembali: kajian kesesuaian habitat dan penentuan lokasi pelepasliaran lima jenis satwa di kawasan Taman Nasional Gunung Halimun Salak Periode Tahun 2021–2025. Direktorat Jenderal Konservasi Sumber Daya Alam dan Ekosistem Kementerian Lingkungan Hidup dan Kehutanan, Jakarta, 66 pp.

Ghoddousi, A., A.K. Hamidi, T. Ghadirian, D. Ashayeri & I. Khorozyan (2010). The status of the endangered Persian Leopard Panthera pardus saxicolor in Bamu National Park, Iran. Oryx 44(4): 551–557. DOI:

Ghoddousi, A., B. Bleyhl, C. Sichau, D. Ashayeri, P. Moghadas, P. Sepahvand, A.K. Hamidi, M. Soofi & T. Kuemmerle (2020). Mapping connectivity and conflict risk to identify safe corridors for the Persian Leopard. Landscape Ecology 35: 1809–1825. DOI:

Ghoddousi, A., M. Soofi, A.K. Hamidi, T. Lumetsberger, L. Egli, S. Ashayeri & M. Waltert (2017). When pork is not on the menu: assessing trophic competition between large carnivores and poachers. Biological Conservation 209: 223–229. DOI:

Goldberg, J.F., T. Tempa, N. Norbu, M. Hebblewhite, L.S. Mills, T.R. Wangchuk & P. Lukacs (2015). Examining temporal sample scale and model choice with spatial capture-recapture models in the common Leopard Panthera pardus. PloS One 10(11): e0140757. DOI:

Gomez, L. & C.R. Shepherd (2021). The illegal exploitation of the Javan Leopard (Panthera pardus melas) and Sunda Clouded Leopard (Neofelis diardi) in Indonesia. Nature Conservation 43: 25–39. DOI:

Gray, T.N.E. & S. Prum (2012). Leopard density in post-conflict landscape, Cambodia: evidence from spatially explicit capture-recapture. Journal of Wildlife Management 76(1): 163–169. DOI:

Grassman, Jr. L.I. (1999). Ecology and behavior of the Indochinese Leopard in Kaeng Krachan National Park, Thailand. Natural History Bulletin of the Siam Society 47: 77–93.

Gunawan, H., S. Iskandar, V.S. Sihombing & R. Wienanto (2017). Conflict between humans and Leopards (Panthera pardus melas Cuvier, 1809) in western Java, Indonesia. Biodiversitas 18(2): 652–658. DOI:

Gunawan, H. & V.S. Sihombing (2017). Preferensi habitat Macan Tutul Jawa (Panthera pardus melas Cuvier 1809) di Jawa bagian barat. Jurnal Penelitian Hutan dan Konservasi Alam 1: 35–43. DOI:

Gunawan, H., L.B. Prasetyo, A. Mardiastuti & A.P. Kartono (2012). Habitat Macan Tutul Jawa (Panthera pardus melas Cuvier 1809) di lansekap hutan tanaman pinus. Jurnal Penelitian Hutan dan Konservasi Alam 9(1): 49–67. DOI:

Gunawan, H., L.B. Prasetyo, A. Mardiastuti & A.P. Kartono (2009). Studi karakteristik habitat dan daerah penyebaran Macan Tutul Jawa (Panthera pardus melas, Cuvier 1809) di Jawa Tengah dan Yogyakarta. Jurnal Penelitian Hutan dan Konservasi Alam 21(2): 95–110. DOI:

Harihar, A., B. Pandav & S.P. Goyal (2009). Density of Leopards (Panthera pardus) in the Chilla Range of Rajaji National Park, Uttarakhand, India. Mammalia 73: 68–71. DOI:

Harris, K.M. (1996). Mt. Gede Pangrango National Park Information book series. Volume 2. Gede Pangrango National Park. Cianjur, 106 pp.

Hayward, M.W., L. Boitani, N.D. Burrows, P.J. Funston, K.U. Karanth, D.I. MacKenzie, K.H. Pollock & R.W. Yarnell (2015). Ecologists need to use robust survey design, sampling and analysis methods. Journal of Applied Ecology 52(2): 286–290. DOI:

Hayward, M.W., J. O’Brien & G.I.H. Kerley (2007). Carrying capacity of large African predators: Predictions and tests. Biological Conservation 139(1–2): 219–229. DOI:

Havmøller, R.W., S. Tenan, N. Scharff & F. Rovero (2019). Reserve size and anthropogenic disturbance affect the density of an African Leopard (Panthera pardus) meta-population. PLoS One 14(6): e0209541. DOI:

Hedges, L., W.Y. Lam, A. Campos-Arceiz, D.M. Rayan, W.F. Laurance, C.J. Latham & G.R. Clements (2015). Melanistic Leopards reveal their spots: Infrared camera traps provide a population density estimate of Leopards in Malaysia. The Journal of Wildlife Management 79(5): 846–853. DOI:

Henschel, P. & J. Ray (2003). Leopards in African Rainforests: Survey and Monitoring Techniques. Wildlife Conservation Society, New York, 50pp.

Hines, J.E. (2006). PRESENCE. Software to estimate path occupancy and related parameters. USGS-PWRC, http://www.mbr-pwrc.usgs. gov/software/presence.html.

Islam, M.Z., A. Gavashelishvili, L. Kokiashvili, A. al Boug & A.A. Shehri (2021). Modelling the distribution and movement intensity of the Arabian Leopard Panthera pardus nimr (Mammalia: Felidae). Zoology in the Middle East 67(2): 106–118. DOI:

Kalle, R., T. Ramesh, Q. Qureshi & K. Sankar (2011). Density of Tiger and Leopard in a tropical deciduous forest of Mudumalai Tiger Reserve, southern India, as estimated using photographic capture-recapture sampling. Acta Theriologica 56: 335–342. DOI:

Kandel, S.R., B.R. Lamichhane & N. Subedi (2020). Leopard (Panthera pardus) density and diet in a forest corridor of Terai: implications for conservation and conflict management. Wildlife Research 47(6): 460–467. DOI:

Karanth, K.U. (1995) Estimating Tiger Panthera tigris populations from camera-trap data using capture–recapture models. Biological Conservation 71(3): 333–338. DOI:

Karanth, K.U. & M.E. Sunquist (2000). Behavioural correlates of predation by Tiger (Panthera tigris), Leopard (Panthera pardus) and Dhole (Cuon alpinus) in Nagarahole, India. Journal of Zoology 250(2): 255–265. DOI:

Kawanishi, K., A.M. Sahak & M. Sunquist (1999). Preliminary analysis on abundance of large mammals at Sungai Relau,Taman Negara. Journal of Wildlife and National Parks 17: 62–82.

Kebun Raya Bogor (2001). Laporan Eksplorasi Kawasan Hutan Gunung Ciremai. Kebun Raya Bogor, LIPI, Bogor, 17 pp.

Kittle, A.M. & A.C. Watson (2017). Density of Leopards (Panthera pardus kotiya) in Horton Plains National Park in the Central Highlands of Sri Lanka. Mammalia 82(2): 183–187. DOI:

Kittle, A.M., A.C. Watson & P.K.L. Samaranayake (2021). Edge effects and distribution of prey forage resources influence how an apex predator utilizes Sri Lanka’s largest protected area. Journal of Zoology 314(1): 31–42. DOI:

Kittle, A.M., A.C. Watson & T.S.P. Fernando (2017). The ecology and behaviour of a protected area Sri Lankan Leopard (Panthera pardus kotiya) population. Tropical Ecology 58(1): 71–86.

Kittle, A.M., A.C.Watson, S.A. Cushman & D.W. Macdonald (2018). Forest cover and level of protection influence the island-wide distribution of an apex carnivore and umbrella species, the Sri Lankan Leopard (Panthera pardus kotiya). Biodiversity Conservation 27: 235–263. DOI:

Khorozyan, I.G., A.G. Malkhasyan & A.V. Abramov (2008). Presence–absence surveys of prey and their use in predicting Leopard (Panthera pardus) densities: a case study from Armenia. Integrative Zoology 3(4): 322–332. DOI:

Kshettry, A., S. Vaidyanathan & V. Athreya (2017). Leopard in a tea-cup: a study of Leopard habitat-use and human-Leopard interactions in north-eastern India. PLoS One 12(5): e0177013. DOI:

Lamichhane, B.R., S. Lamichhane, R. Regmi, M. Dhungana, S.K. Thapa, A. Prasai, A. Gurung, S. Bhattarai, R.P. Paudel & N. Subedi (2021). Leopard (Panthera pardus) occupancy in the Chure range of Nepal. Ecology and Evolution 11(20): 13641–13660. DOI:

Linkie, M. (2008). Single-Species, Single-Season Occupancy Tutorials. DICE, University of Kent, 22pp.

MacKenzie, D.I., J.D. Nichols, J.A. Royle, K.H. Pollock, L.L. Bailey & J.E. Hines (2006). Occupancy estimation and modelling: Inferring Patterns and Dynamics of Species Occurrence. Academic Press, London, 324 pp.

MacKenzie, D.I. & L.L. Bailey (2004). Assessing the fit of site-occupancy models. Journal of Agricultural, Biological, and Environmental Statistics 9(3): 300–318. DOI:

Malau, P.W. (2013). Kesesuaian habitat Macan Tutul Jawa (Panthera pardus melas G. Cuvier) di Resort Gunung Botol, Taman Nasional Gunung Halimun Salak, Provinsi Jawa Barat [skripsi]. Institut Pertanian Bogor. Bogor, i-vii+42 pp.

Ministry of Environment and Forestry (2015). Pedoman peningkatan populasi 25 satwa terancam punah. Direktorat Jenderal Konservasi Sumberdaya Alam dan Ekosistem, Kementerian Lingkungan Hidup dan Kehutanan Republik Indonesia, Jakarta, Indonesia, 436 pp.

Ministry of Environment and Forestry (2016). Strategi dan rencana aksi konservasi macan tutul jawa (Panthera pardus melas): 2016–2026. Direktorat Jenderal Konservasi Sumberdaya Alam dan Ekosistem, Kementerian Lingkungan Hidup dan Kehutanan Republik Indonesia, Jakarta, Indonesia, 39 pp.

Ministry of Environment and Forestry (2018). Laporan Kinerja 2018. Direktorat Konservasi Keanekaragaman Hayati. Jakarta, Indonesia, i-ix+94 pp.

Mondal, K., K. Sankar & Q. Qureshi (2013). Factors influencing the distribution of Leopard in a semiarid landscape of western India. Acta Theriologica 58(2): 179–187. DOI:

Ngoprasert, D., A.J. Lynam & G.A. Gale (2007). Human disturbance affects habitat use and behaviour of Asiatic Leopard Panthera pardus in Kaeng Krachan National Park, Thailand. Oryx 41(3): 343–351. DOI:

Ngoprasert, D., A.J. Lynam & G.A. Gale (2017). Effects of temporary closure of a national park on Leopard movement and behaviour in tropical Asia. Mammalian Biology 82: 65–73. DOI:

Noor, A., Z.R. Mir, G.G. Veeraswami & B. Habib (2020). Density of Leopard in a moist-temperate forest of western Himalaya, India. Tropical Ecology 61: 301–310. DOI:

Nowell, K. & P. Jackson (1996). Leopard Panthera pardus (Linnaeus, 1758), pp. 24–30, 44–47, 78–82. In: Wild Cats: Status Survey and Conservation Action Plan. IUCN/SSC Cat Specialist Group, Gland, 382 pp.

O’Brien, T.G., M.F. Kinnaird & H.T. Wibisono (2003). Crouching Tigers, hidden prey: Sumatran Tiger and prey populations in a tropical forest landscape. Animal Conservation 6(2): 131–139. DOI:

Odden, M. & P. Wegge (2005). Spacing and activity patterns of Leopards Panthera pardus in the Royal Bardia National Park, Nepal. Wildlife Biology 11(2): 145–152.[145:SAAPOL]2.0.CO;2 DOI:[145:SAAPOL]2.0.CO;2

Otis, D.L., K.P. Burham, G.C. White & D.R. Anderson (1978). Statistical inference from capture data on closed animal populations. Wildlife Monographs 62: 1–135.

Pawar, D., H.P. Nelson, D.R.L. Pawar & S. Khanwilkar (2019). Estimating Leopard Panthera pardus fusca (Mammalia: Carnivora: Felidae) abundance in Kuno Wildlife Sanctuary, Madhya Pradesh, India. Journal of Threatened Taxa 11(5): 13531–13544. DOI:

Rahayu, N.D., B. Sasmito & N. Bashit (2018). Analisis pengaruh fenomena Indian Ocean Dipole (OID) terhadap curah hujan di Pulau Jawa. Jurnal Geodesi Undip 7(1): 57–67.

Rahman, D.A., P. Rianti, M. Muhiban, A. Muhtarom, U.M. Rahmat, Y. Santosa & S. Aulagnier (2018). Density and spatial partitioning of endangered sympatric Javan Leopard (Felidae) and Dholes (Canidae) in a tropical forest landscape. Folia Zoologica 67(3–4): 207–219. DOI:

Ramesh, T., R. Kalle, H. Rosenlund & C.T. Downs (2017). Low Leopard populations in protected areas of Maputaland: a consequence of poaching, habitat condition, abundance of prey, and a top predator. Ecology and Evolution 7: 1964–1973. DOI:

Ramesh, T. & C.T. Downs (2013). Impact of farmland use on population density and activity patterns of Serval in South Africa. Journal of Mammalogy 94: 1460–1470. DOI:

R Core Team (2018). R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.

Rosenblatt, E., S. Creel, M.S. Becker, J. Merkle, H. Mwape, P. Schuette & T. Simpamba (2016). Effects of a protection gradient on carnivore density and survival: An example with Leopards in the Luangwa valley, Zambia. Ecology and Evolution 6(11): 3772–3785. DOI:

Rostro-García, S., J.F. Kamler, R. Crouthers, K. Sopheak, S. Prum, V. In, C. Pin, A. Caragiulo & D.W. Macdonald (2018). An adaptable but threatened big cat: density, diet and prey selection of the Indochinese Leopard (Panthera pardus delacouri) in eastern Cambodia. Royal Society Open Science 5(2): 171–187. DOI:

Royle, J.A., J.D. Nichols, K.U. Karanth & A.M. Gopalaswamy (2009). A hierarchical model for estimating density in camera-trap studies. Journal of Applied Ecology 46: 118–127. DOI:

Santiapillai, C. & W.S. Ramono (1992). Status of the Leopard (Panthera pardus) in Java, Indonesia. Tiger Paper 29(2): 1–5.

Sehgal, J.J., D. Kumar, R.S. Kalsi, M.L. Allen & R. Singh (2022). Spatio-temporal overlap of Leopard and prey species in the foothills of Shiwalik, Himalaya. European Journal of Wildlife Research 68(2): 1–9. DOI:

Simbolon, H., M. Yoneda & J. Sugardjito (1998). Biodiversity and its conservation in Gunung Halimun National Park. Pp. 46–57 in: Simbolon, H., M. Yoneda & J. Sugardjito (eds.). Research and Conservation of Biodiversity in Indonesia. Vol. IV: Gunung Halimun, the last submontane tropical forest in West Java. JICA – LIPI – PHPA, Bogor, 153 pp.

Sharbafi, E., M.S. Farhadinia, H.R. Rezaie & A.R. Braczkowski (2016). Prey of the Persian Leopard (Panthera pardus saxicolor) in a mixed forest-steppe landscape in northeastern Iran (Mammalia: Felidae). Zoology in the Middle East 62(1): 1–8. DOI:

Sollmann, R., B. Gardner & J.L. Belant (2012). How Does Spatial Study Design Influence Density Estimates from Spatial Capture-Recapture Models? PLoS One 7(4): e34575. DOI:

Stein, A.B. & V. Hayssen (2013). Panthera pardus (Carnivora: Felidae). Mammalian Species 45(900): 30–48. DOI:

Stein, A.B., V. Athreya, P. Gerngross, G. Balme, P. Henschel, U. Karanth, D. Miquelle, S. Rostro-Garcia, J.F. Kamler, A. Laguardia, I. Khorozyan & A. Ghoddousi (2020). Panthera pardus. The IUCN Red List of Threatened Species 2020: e.T15954A163991139. Downloaded on 15 April 2020. DOI:

Steinmetz, R., W. Chutipong, N. Seuaturien & B. Poonnil (2009). Ecology and conservation of Tigers and their prey in Kuiburi National Park, Thailand. WWF Thailand, and Department of National Parks, Wildlife, and Plant Conservation, Bangkok, 58 pp.

Strampelli, P., L. Andresen, K.T. Everatt, M.J. Somers & J.M. Rowcliffe (2018). Habitat use responses of the African Leopard in a human-disturbed region of rural Mozambique. Mammalian Biology 89: 14–20. DOI:

Strampelli, P., L. Andresen, K.T. Everatt, M.J. Somers & J.M. Rowcliffe (2020). Leopard Panthera pardus density in southern Mozambique: evidence from spatially explicit capture–recapture in Xonghile Game Reserve. Oryx 54(3): 405–411. DOI:

Tan, C.K.W., D.G. Rocha, G.R. Clements, E. Brenes-Mora, L. Hedges, K. Kawanishi, S.W. Mohamad, D.M. Rayan, G. Bolongon, J. Moore, J. Wadey, A. Campos-Arceiz & D.W. Macdonald (2017). Habitat use and predicted range for the mainland Clouded Leopard Neofelis nebulosa in Peninsular Malaysia. Biological Conservation 206: 65–74. DOI:

Tobler, M.W., S.E. Carrillo-Percastegui, A.Z. Hartley & G.V.N. Powell (2013). High Jaguar densities and large population sizes in the core habitat of the southwestern Amazon. Biological Conservation 159: 375–381. DOI:

Traylor-Holzer, K., B. Holst, K. Leus & K. Ferraz (2020). Conservation Planning Workshops for the Javan Leopard (Panthera pardus melas). IUCN SSC Conservation Planning Specialist Group, Apple Valley, 61 pp.

Vinks, M.A., S. Creel, E. Rosenblatt, M.S. Becker, P. Schuette, B. Goodheart, C. Sanguinetti, K. Banda, C. Chifunte & C. Simukonda (2021). Leopard Panthera pardus density and survival in an ecosystem with depressed abundance of prey and dominant competitors. Oryx 56(4): 518–527. DOI:

Wang, S.W. & D.W. Macdonald (2009). The use of camera traps for estimating Tiger and Leopard populations in the high-altitude mountains of Bhutan. Biological Conservation 142(3): 606–613. DOI:

Wibisono, H.T., E. Wilianto, I.M.R. Pinondang, D.A. Rahman & D. Chandradewi (2021). Panthera pardus ssp. melas. The IUCN Red List of Threatened Species 2021: e.T15962A50660931. Downloaded on 11 September 2021. DOI:

Wibisono, H.T., H.A. Wahyudi, E. Wilianto, I.M.R. Pinondang, M. Primajati, D. Liswanto & M. Linkie (2018). Identifying priority conservation landscapes and actions for the Critically Endangered Javan Leopard in Indonesia: Conserving the last large carnivore in Java Island. PLoS One 13(6): e0198369. DOI:

Yang, H., B. Xie, G. Zhao, Y. Gong, P. Mou, J. Ge & L. Feng (2021). Elusive cats in our backyards: persistence of the North Chinese Leopard (Panthera pardus japonensis) in a human-dominated landscape in central China. Integrative Zoology 16(1): 67–83. DOI: