Activity patterns of the small and medium felid (Mammalia: Carnivora: Felidae) guild in northeastern India

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Shomita Mukherjee
Priya Singh
André Pinto Silva
Chandan Ri
Kashmira Kakati
Binod Borah
Tana Tapi
Sandesh Kadur
Prafull Choudhary
Shikha Srikant
Surabhi Nadig
R. Navya
Mats Björklund
Uma Ramakrishnan


Fifteen extant species of cats inhabit India, and the northeastern region of the country is among the richest with nine species.  Among these are the “standard fourâ€, an assemblage of Clouded Leopard Neofelis nebulosa, Asiatic Golden Cat Catopuma temminckii, Marbled Cat Pardofelis marmorata, and Leopard Cat Prionailurus bengalensis, which also occur across southeastern Asia.  Within India, despite several surveys in this region, very little information exists on the ecology of this assemblage to explain their co-occurrence.  In this paper, we put together data from several independent camera trapping studies over 10 sites across northeastern India to examine and interpret diel activity patterns of this group.  While we present results for all the four species, we focus on two species, the Marbled Cat and Leopard Cat, which are of very similar body size and are potential competitors.  We used kernel density estimates to measure diel activity patterns of all four species and overlap in activity between Marbled Cat and Leopard Cat at the regional scale as well as the point scale.  We obtained 783 captures of the standard four from >27,500 trap nights.  The Asiatic Golden Cat and Marbled Cat were strongly diurnal, Clouded Leopard largely crepuscular and nocturnal, and Leopard Cat largely nocturnal.  The degree of overlap between Marbled Cat and Leopard Cat activity was low and in consensus with other studies across southeastern Asia.  We interpret this as the differing niche spaces of the two cats due to their specific pre-existing adaptations, not restricted to the effects of competition.  The point scale analysis when both cats are captured at the same location and separately show no shift in activity pattern, supporting our hypothesis of pre-existing differences in resources, such as food, playing a major role in facilitating co-existence.  Our study, however, is preliminary and additional information with robust analysis is required to test this finding.

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Andheria, A.P., K.U. Karanth & N.S. Kumar (2007). Diet and prey profiles of three sympatric large carnivores in Bandipur Tiger Reserve, India. Journal of Zoology 273(2): 169–175;

Athreya, V.R. & A.V. Belsare (2006). Report providing the Maharashtra Forest Department Technical and Veterinary Support to better deal with Wild Animals that require Human Intervention. Wildlife Trust of India, New Delhi & Office of the Chief Wildlife Warden, Maharashtra, 29pp.

Azlan, J.M. & D.S. Sharma (2006). The diversity and activity patterns of wild felids in a secondary forest in peninsular Malaysia. Oryx 40(1): 36–41;

Borries, C., Z.M. Primeau, K. Ossi-Lupo, S. Dtubpraserit & A. Koenig (2014). Possible predation attempt by a marbled cat on a juvenile Phayre’s Leaf Monkey. Raffles Bulletin of Zoology 62: 561–565.

Brown, J. & E.O. Wilson (1956). Character displacement. Systematic Zoology 5: 49–64.

Caro, T.M. & C.J. Stoner (2003). The potential for interspecific competition among African carnivores. Biological Conservation 110(1): 67–75;

Champion, H.G. & S.K. Seth (1968). A Revised Survey of the Forest Types of India. Natraj Publishers, Dehradun, 404pp.

Cruz, P., M.E. Lezzi, C.D. Angelo, D. Varela, M.S.D. Bitetti & A. Paviolo (2018). Effects of human impacts on habitat use, activity patterns and ecological relationships among medium and small felids of the Atlantic forest. PloS ONE 13(8): e0200806;

Dayan, T., D. Simberloff, E. Tchernov & Y. Yom-Tov (1990). Feline canines: community-wide character displacement in the small cats of Israel. American Naturalist 136(1): 39–60;

Duckworth, J.W., A.J. Lynam & C. Breitenmoser-Würsten (2014). Non-Panthera cats in southeast Asia: present knowledge and recommendation. Cat News Special Issue 8: 62–67.

Grewal, B., R. Sreenivasan & B. Haralu (2012). Report on the Nagaland Wildlife & Biodiversity Conservation Project: An Action Document. Nagaland Wildlife Conservation Project, Dimapur, 156pp.

Grassman, L., A. Lynam, S. Mohamad, J.W. Duckworth, J. Borah, D. Wilcox, Y. Ghimirey, A. Reza & H. Rahman (2016). Neofelis nebulosa. In: The IUCN Red List of Threatened Species: e.T14519A97215090. Downloaded on 28 December 2018;

Gumal, M., A.B.B.M. Salleh, M.N. Yasak, L.S. Horng, B.P.Y.H. Lee, L.C. Pheng, H. Hamzah, D. Kong, D. Magintan, D.T.C. Yung, A.Z.B. Zalaluddin, A.B. Azmi, N.B. Khalid, T.P. Yen, V. Mufeng, F.C.F. Meng & S. Ng (2014). Small-medium wild cats of Endau Rompin Landscape in Johor Peninsular Malaysia. Cat News Special Issue 8: 10–18.

Hearn, A.J., S.A. Cushman, J. Ross, B. Goossens, L.T.B. Hunter & D.W. Macdonald (2018). Spatio-temporal ecology of sympatric felids on Borneo. Evidence for resource partitioning? PLoS ONE 13(7): e0200828;

Hunter, L. (2015). Wild Cats of the World, 1st Ed. Bloomsbury, London, 240pp.

Johnson, W.E., E. Eizirik, J. Pecon-Slattery, W.J. Murphy, A. Antunes, E. Teeling & S.J. O’Brien (2006). The late Miocene radiation of modern Felidae: a genetic assessment. Science 311(5757): 73–77;

Kaul, R. & K. Haridasan (1987). Forest types of Arunachal Pradesh—a preliminary study. Journal of Economic and Taxonomic Botany 9(2): 379–389.

Kitchener, A.C., C. Breitenmoser-Würsten, E. Eizirik, A. Gentry, L. Werdelin, A. Wilting, N. Yamaguchi, A.V. Abramov, P. Christiansen, C. Driscoll, J.W. Duckworth, W. Johnson, S.-J. Luo, E. Meijaard, P. O’Donoghue, J. Sanderson, K. Seymour, M. Bruford, C. Groves, M. Hoffmann, K. Nowell, Z. Timmons & S. Tobe (2017). A revised taxonomy of the Felidae. The final report of the Cat Classification Task Force of the IUCN/ SSC Cat Specialist Group. Cat News Special Issue 11: 1–80.

Leibold, M.A., M. Holyoak, N. Mouquet, P. Amarasekare, J.M. Chase & M.F. Hoopes (2004). The metacommunity concept: a framework for multi-scale community ecology. Ecology Letters 7(7): 601–613;

Longchar, S. (2013). Study on the Trends of Bushmeat Consumption and Traditional Hunting on Wild Fauna by Indigenous Community living near Protected Area in Nagaland. Master’s Dissertation. Saurashtra University, Rajkot.

Lorica, M.R.P & L.R. Heaney (2013). Survival of a native mammalian carnivore, the Leopard Cat Prionailurus bengalensis Kerr, 1792 (Carnivora: Felidae), in an agricultural landscape on an oceanic Philippine island. Journal of Threatened Taxa 5(10): 4451–4460;

Losos, J.B. (2000). Ecological character displacement and the study of adaptation. Proceedings of the National Academy of Sciences USA 97(11): 5693–5695;

Lynam, A.J., K.E. Jenks, N. Tantipisanuh, W. Chutipong, D. Ngoprasert, G.A. Gale, R. Steinmetz, R. Sukmasuang, N. Bhumpakphan, L.I. Grassman Jr., P. Cutter, S. Kitamura, D.H. Reed, M.C. Baker, W. McShea, N. Songsasen & P. Leimgruber (2013). Terrestrial activity patterns of wild cats from camera-trapping. The Raffles Bulletin of Zoology 61(1): 407–415.

Lyngdoh, S., S. Shrotriya, S.P. Goyal, H. Clements, M.W. Hayward & B. Habib (2014). Prey preferences of the snow leopard (Panthera uncia): regional diet specificity holds global significance for conservation. PloS ONE 9(2): e88349;

Mani, M.S. (ed.) (1974). Ecology and Biogeography in India. Dr. W. Junk B.V. Publishers, The Hague, 772pp.

McCarthy, J.L., H.T. Wibisono, K.P. McCarthy, T.K. Fuller & N. Andayani (2015). Assessing the distribution and habitat use of four felid species in Bukit Barisan Selatan National Park, Sumatra, Indonesia. Global Ecology and Conservation 3: 210–221;

McCarthy, J., S. Dahal, T. Dhendup, T.N.E. Gray, S. Mukherjee, H. Rahman, P. Riordan, N. Boontua & D. Willcox (2015). Catopuma temminckii. In: The IUCN Red List of Threatened Species: e.T4038A97165437. Downloaded on 28 December 2018;

Meredith, M. & M. Ridout (2018). Package ‘overlap’: estimates of coefficient of overlapping for animal activity patterns. R Package Version 0.3.2. Available online at

Mittermeier, R.A., R. Robles-Gil, M. Hoffman, J. Pilgrim, T. Brooks, C.G. Mittermeier, J. Lamoreux & G.A.B. Da Fonseca (2004). Hotspots Revisited: Earth’s Biologically Richest and Most Endangered Terrestrial Ecoregions. CEMEX, Mexico City, 392pp.

Morales, M.M. & N.P. Giannini (2010). Morphofunctional patterns in Neotropical felids: species co-existence and historical assembly. Biological Journal of the Linnean Society 100(3): 711–724;

Mukherjee, S., S.P. Goyal, A.J.T. Johnsingh & M.L. Pitman (2004). The importance of rodents in the diet of Jungle Cat (Felis chaus), Caracal (Caracal caracal) and Golden Jackal (Canis aureus) in Sariska Tiger Reserve, Rajasthan, India. Journal of Zoology 262(4): 405–411;

Mukherjee, S., R. Athreya, P.V. Karunakaran & P. Choudhary (2016). Report on Ecological Species Sorting in relation to Habitat Structure in the Small Cat Guild of Eaglenest Wildlife Sanctuary, Arunachal Pradesh. Salim Ali Centre for Ornithology and Natural History, Coimbatore, 52pp.

Nowell, K. & P. Jackson (eds.) (1996). Wild Cats: Status Survey and Conservation Action Plan, Vol. 382. IUCN SSC Cat specialist Group, Gland, 383pp.

Palomares, F., N. Fernández, S. Roques, C. Chávez, L. Silveira, C. Keller & B. Adrados (2016). Fine-scale habitat segregation between two ecologically similar top predators. PLoS ONE 11(5): e0155626;

Pocock, R.I. (1939). Family Felidae, pp191–330. In: The Fauna of British India, including Ceylon and Burma: Mammalia, Vol. 1, Primates and Carnivora. Taylor & Francis, Ltd., London, 572pp.

Pusparini, W., H.T. Wibisono, G.V. Reddy, T. Tarmizi & P. Bharata (2014). Small and medium-sized cats in Gunung Leuser National Park, Sumatra, Indonesia. Cat News Special Issue 8: 4–9.

R Core Team (2014). R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available online at

Ridout, M.S. & M. Linkie (2009). Estimating overlap of daily activity patterns from camera-trap data. Journal of Agricultural, Biological, and Environmental Statistics 14(3): 322–337;

Ross, J., A.J. Hearn, P.J. Johnson & D.W. Macdonald (2013). Activity patterns and temporal avoidance by prey in response to Sunda Clouded Leopard predation risk. Journal of Zoology 290(2): 96–106;

Ross, J., J. Brodie, S. Cheyne, A. Datta, A. Hearn, B. Loken, A. Lynam, J. McCarthy, C. Phan, A. Rasphone, P. Singh & A. Wilting (2016). Pardofelis marmorata. In: The IUCN Red List of Threatened Species: e.T16218A97164299. Downloaded on 28 December 2018;

Schoener, T.W. (1974). Resource partitioning in ecological communities. Science 185(4145): 27–39;

Sekhsaria, P. (2012). Report on the State of Wildlife in northeast India 1996–2011: A Compilation of News from the Protected Area Update. Foundation for Ecological Security, 293pp.

Shehzad, W., T. Riaz, M.A. Nawaz, C. Miquel & C. Poillot (2012). Carnivore diet analysis based next generation sequencing: application to the Leopard Cat (Prionailurus bengalensis) in Pakistan. Molecular Ecology 21(8): 1951–65;

Singh, P. & D.W. Macdonald (2017). Population and activity patterns of Clouded Leopards and Marbled Cats in Dampa Tiger Reserve, India. Journal of Mammalogy 98(5): 1453–1462;

Smith, J.L.D., M.E. Sunquist, K.M. Tamang & P.B. Rai (1983). A technique for capturing and immobilizing tigers. The Journal of Wildlife Management 47(1): 255–259.

Sunarto, S., M.J. Kelly, K. Parakkasi & M.B. Hutajulu (2015). Cat coexistence in central Sumatra: ecological characteristics, spatial and temporal overlap, and implications for management. Journal of Zoology 296(2): 104–115;

Sunquist, M. & F. Sunquist (2002). Wild Cats of the World. University of Chicago Press, Chicago, Illinois, 462pp.

Velho, N., U. Srinivasan, P. Singh & W.F. Laurance (2015). Large mammal use of protected and community-managed lands in a biodiversity hotspot. Animal Conservation 19(2): 199–208;

Xiong, M., D. Wang, H. Bu, X. Shao, D. Zhang, S. Li, R. Wang & M. Yao (2017). Molecular dietary analysis of two sympatric felids in the Mountains of southwest China biodiversity hotspot and conservation implications. Scientific Reports 7: 41909;

Zaw, T., T. Myint, S. Htun, S.H.T. Po, K.T. Latt, M. Maung & A.J. Lynam (2014). Status and distribution of smaller cats in Myanmar. Cat News Special Issue 8: 24–30.