Leopards at large: population density, habitat utilisation and spatio-temporal overlap of leopards (Panthera pardus) with competitors and prey in the Rajaji Tiger Reserve, northern India
Shashank Yadav
A , Manu Mohan A , Aditi Sharma B , Sanatan Sonker B and Ramesh Krishnamurthy A C *
A
B
C
Abstract
In the vicinity of the western Rajaji Tiger Reserve, a significant rise in conflicts between humans and leopards has occurred. These conflicts have resulted in a substantial number of human deaths, with 60% attacks resulting in deaths over a 4-year period.
This study aims to examine any potential connection among the non-breeding tigers, leopards, prey, humans and the increased occurrence of conflicts.
To address this objective, we conducted an analysis to evaluate the density of leopards, as well as their spatial and temporal behaviour with respect to tigers, prey and humans through on-site habitat use, circular statistics, diel-overlap as well spatial niche breadth and spatial-niche overlap.
Our findings showed one of the highest leopard-density estimates (excluding cubs) (mean ± s.e.) of 23.7 ± 4.8 per 100 km2. When examining on-site habitat use by leopards, we found three variables that had a negative influence, namely, human presence, slope, and the presence of tigers. The presence of a large prey (sambar) had a positive influence. Leopards, exhibiting the highest spatial-niche breadth, demonstrated a small overlap with both tigers and humans, while displaying a significant diel overlap with sambar. The niche overlap was notably high with sambar, but very low with humans and tigers.
Although tigers locally created a zone of spatial and temporal avoidance, at the population level the leopards experienced a surge in numbers in response to the reduced tiger numbers, indicated by their high density. This could be a cause of high leopard movement into nearby human settlements. This unique scenario could have contributed to conflict.
Understanding the coexistence among different species and the impact of transient animals is crucial to develop effective management strategies to mitigate conflict. This approach would facilitate and ensure the long-term survival of diverse large carnivores in the tropical forests of southern Asia through sustained support for conservation from local communities.
Keywords: activity pattern, habitat use, human–field conflict, leopard, Panthera pardus, population density, prey, southern Asia.
References
Allen ML, Wang S, Olson LO, Li Q, Krofel M (2020) Counting cats for conservation: seasonal estimates of leopard density and drivers of distribution in the Serengeti. Biodiversity and Conservation 29, 3591-3608.
| Crossref | Google Scholar |
Athreya VR, Thakur SS, Chaudhuri S, Belsare AV (2004) A study of the man-leopard conflict in the Junnar Forest Division, Pune District, Maharashtra. Wildlife Protection Society of India, New Delhi, India. pp. 89. Available at https://citeseerx.ist.psu.edu/document?repid=rep1&type=pdf&doi=37a9d778161b5f1b824b8e80fc5fb8c4154e4cd7
Balme GA, Slotow R, Hunter LTB (2010) Edge effects and the impact of non-protected areas in carnivore conservation: leopards in the Phinda–Mkhuze Complex, South Africa. Animal Conservation 13, 315-323.
| Crossref | Google Scholar |
Balme GA, Hunter L, Braczkowski AR (2012) Applicability of age-based hunting regulations for African leopards. PLoS ONE 7, e35209.
| Crossref | Google Scholar | PubMed |
Balme GA, Pitman RT, Robinson HS, Miller JRB, Funston PJ, Hunter LTB (2017) Leopard distribution and abundance is unaffected by interference competition with lions. Behavioral Ecology 28(5), 1348-1358.
| Crossref | Google Scholar |
Beier P (1991) Cougar attacks on humans in the United States and Canada. Wildlife Society Bulletin (1973-2006) 19, 403-412 Available at https://www.jstor.org/stable/3782149.
| Google Scholar |
Brown WL, Wilson EO, Jr (1956) Character displacement. Systematic Zoology 5, 49-64.
| Crossref | Google Scholar |
Burnham KP, Anderson DR (2004) Multimodel inference: understanding AIC and BIC in model selection. Sociological Methods & Research 33, 261-304.
| Crossref | Google Scholar |
Carter N, Jasny M, Gurung B, Liu J (2015) Impacts of people and tigers on leopard spatiotemporal activity patterns in a global biodiversity hotspot. Global Ecology and Conservation 3, 149-162.
| Crossref | Google Scholar |
Chase Grey JN, Kent VT, Hill RA (2013) Evidence of a high density population of harvested leopards in a montane environment. PLoS ONE 8, e82832.
| Crossref | Google Scholar |
Crall JP, Stewart CV, Berger-Wolf TY, Rubenstein DI, Sundaresan SR (2013) HotSpotter – patterned species instance recognition. In ‘2013 IEEE Workshop on Applications of Computer Vision (WACV)’, pp. 230–237. (IEEE) 10.1109/wacv.2013.6475023
Creel S (2001) Four factors modifying the effect of competition on carnivore population dynamics as illustrated by African wild dogs. Conservation Biology 15, 271-274.
| Crossref | Google Scholar |
Creel S, Christianson D (2008) Relationships between direct predation and risk effects. Trends in Ecology & Evolution 23, 194-201.
| Crossref | Google Scholar | PubMed |
Creel S, Creel NM (1998) Six ecological factors that may limit African wild dogs, Lycaon pictus. Animal Conservation Forum 1, 1-9.
| Crossref | Google Scholar |
Dalerum F, Cameron EZ, Kunkel K, Somers MJ (2009) Diversity and depletions in continental carnivore guilds: implications for prioritizing global carnivore conservation. Biology Letters 5, 35-38.
| Crossref | Google Scholar | PubMed |
Dennell RW, Coard R, Turner A (2008) Predators and scavengers in early pleistocene southern Asia. Quaternary International 192, 78-88.
| Crossref | Google Scholar |
Dröge E, Creel S, Becker MS, M’soka J (2017) Spatial and temporal avoidance of risk within a large carnivore guild. Ecology and Evolution 7, 189-199.
| Crossref | Google Scholar | PubMed |
Durant SM (1998) Competition refuges and coexistence: an example from Serengeti carnivores. Journal of Animal Ecology 67, 370-386.
| Crossref | Google Scholar |
Durant SM (2000) Living with the enemy: avoidance of hyenas and lions by cheetahs in the Serengeti. Behavioral Ecology 11(6), 624-632.
| Crossref | Google Scholar |
Efford M (2019) Habitat masks in the package Secr. p. 18. Available at https://www.otago.ac.nz/density/pdfs/secr-habitatmasks.pdf
Farhadinia MS, McClintock BT, Johnson PJ, Behnoud P, Hobeali K, Moghadas P, Hunter LTB, Macdonald DW (2019) A paradox of local abundance amidst regional rarity: the value of montane refugia for Persian leopard conservation. Scientific Reports 9, 14622.
| Crossref | Google Scholar | PubMed |
Fattebert J, Balme G, Dickerson T, Slotow R, Hunter L (2015) Density-dependent natal dispersal patterns in a leopard population recovering from over-harvest. PLoS ONE 10, e0122355.
| Crossref | Google Scholar | PubMed |
Goldberg JF, Tempa T, Norbu N, Hebblewhite M, Mills LS, Wangchuk TR, Lukacs P (2015) Examining temporal sample scale and model choice with spatial capture–recapture models in the common leopard Panthera Pardus. PLoS ONE 10, e0140757.
| Crossref | Google Scholar | PubMed |
Greenwood PJ (1980) Mating systems, philopatry and dispersal in birds and mammals. Animal Behaviour 28, 1140-1162.
| Crossref | Google Scholar |
Guisan A, Edwards TC, Jr, Hastie T (2002) Generalized linear and generalized additive models in studies of species distributions: setting the scene. Ecological Modelling 157, 89-100.
| Crossref | Google Scholar |
Harihar A, Pandav B, Goyal SP (2009) Density of leopards (Panthera pardus) in the Chilla Range of Rajaji National Park, Uttarakhand, India. Mammalia 73, 68-71.
| Crossref | Google Scholar |
Harihar A, Pandav B, Goyal SP (2011) Responses of leopard Panthera pardus to the recovery of a tiger Panthera tigris population. Journal of Applied Ecology 48, 806-814.
| Crossref | Google Scholar |
Hayward MW, Henschel P, O’Brien J, Hofmeyr M, Balme G, Kerley GIH (2006) Prey preferences of the leopard (Panthera pardus). Journal of Zoology 270, 298-313.
| Crossref | Google Scholar |
Jacobson AP, Gerngross P, Lemeris JR, Jr, et al. (2016) Leopard (Panthera pardus) status, distribution, and the research efforts across its range. PeerJ 4, e1974.
| Crossref | Google Scholar | PubMed |
Johnsingh AJT, Negi AS (2003) Status of tiger and leopard in Rajaji–Corbett Conservation Unit, northern India. Biological Conservation 111, 385-393.
| Crossref | Google Scholar |
Kalle R, Ramesh T, Qureshi Q, Sankar K (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.
| Crossref | Google Scholar |
Karanth KU, Sunquist ME (2000) Behavioural correlates of predation by tiger (Panthera Tigris), leopard (Panthera Pardus) and dhole (Cuon Alpinus) in Nagarahole, India. Journal of Zoology 250, 255-265.
| Crossref | Google Scholar |
Karanth KU, Srivathsa A, Vasudev D, Puri M, Parameshwaran R, Kumar NS (2017) Spatio-temporal interactions facilitate large carnivore sympatry across a resource gradient. Proceedings of the Royal Society B: Biological Sciences 284, 20161860.
| Crossref | Google Scholar |
Karra K, Kontgis C, Statman-Weil Z, Mazzariello JC, Mathis M, Brumby SP (2021) Global land use/land cover with Sentinel 2 and deep learning. In ‘2021 IEEE international geoscience and remote sensing symposium IGARSS’. pp. 4704–4707. (IEEE) doi:10.1109/igarss47720.2021.9553499
Kittle AM, Watson AC (2018) Density of Leopards (Panthera pardus kotiya) in Horton Plains National Park in the Central Highlands of Sri Lanka. Mammalia 82, 183-187.
| Crossref | Google Scholar |
Lamichhane BR, Persoon GA, Leirs H, et al. (2017) Are conflict-causing tigers different? Another perspective for understanding human-tiger conflict in Chitwan National Park, Nepal. Global Ecology and Conservation 11, 177-187.
| Crossref | Google Scholar |
Li Z, Wang T, Smith JLD, Feng R, Feng L, Mou P, Ge J (2019) Coexistence of two sympatric flagship carnivores in the human-dominated forest landscapes of Northeast Asia. Landscape Ecology 34, 291-305.
| Crossref | Google Scholar |
Linnell JDC, Strand O (2000) Interference interactions, co-existence and conservation of mammalian carnivores. Diversity and Distributions 6(4), 169-176.
| Crossref | Google Scholar |
Mandal D, Basak K, Mishra RP, Kaul R, Mondal K (2017) Status of leopard Panthera pardus and striped hyena Hyaena hyaena and their prey in Achanakmar Tiger Reserve, Central India. The Journal of Zoology Studies 4, 34-41.
| Google Scholar |
Maputla NW, Chimimba CT, Ferreira SM (2013) Calibrating a camera trap–based biased mark–recapture sampling design to survey the leopard population in the N’wanetsi concession, Kruger National Park, South Africa. African Journal of Ecology 51, 422-430.
| Crossref | Google Scholar |
Mattson D, Logan K, Sweanor L (2011) Factors governing risk of cougar attacks on humans. Human–Wildlife Interactions 5, 15.
| Crossref | Google Scholar |
Miller JRB, Pitman RT, Mann GKH, Fuller AK, Balme GA (2018) Lions and leopards coexist without spatial, temporal or demographic effects of interspecific competition. Journal of Animal Ecology 87, 1709-1726.
| Crossref | Google Scholar | PubMed |
Mondal K, Gupta S, Bhattacharjee S, Qureshi Q, Sankar K (2012) Response of leopards to re-introduced tigers in Sariska Tiger Reserve, western India. International Journal of Biodiversity and Conservation 4, 228-236.
| Crossref | Google Scholar |
Newsome TM, Greenville AC, Ćirović D, et al. (2017) Top predators constrain mesopredator distributions. Nature Communications 8, 15469.
| Crossref | Google Scholar | PubMed |
Odden M, Wegge P, Fredriksen T (2010) Do tigers displace leopards? If so, why? Ecological Research 25, 875-881.
| Crossref | Google Scholar |
Olson DM, Dinerstein E, Wikramanayake ED, et al. (2001) Terrestrial ecoregions of the world: a new map of life on earth: a new global map of terrestrial ecoregions provides an innovative tool for conserving biodiversity. BioScience 51, 933-938.
| Crossref | Google Scholar |
O’Brien TG, Kinnaird MF, Wibisono HT (2003) Crouching tigers, hidden prey: sumatran tiger and prey populations in a tropical forest landscape. Animal Conservation Forum 6, 131-139.
| Crossref | Google Scholar |
Palomares F, Caro TM (1999) Interspecific killing among mammalian carnivores. The American Naturalist 153, 492-508.
| Crossref | Google Scholar | PubMed |
Pianka ER (1974) Niche overlap and diffuse competition. Proceedings of the National Academy of Sciences 71(5), 2141-2145.
| Google Scholar |
Pokheral CP, Wegge P (2019) Coexisting large carnivores: spatial relationships of tigers and leopards and their prey in a prey-rich area in lowland Nepal. Écoscience 26, 1-9.
| Crossref | Google Scholar |
Prugh LR, Sivy KJ (2020) Enemies with benefits: integrating positive and negative interactions among terrestrial carnivores. Ecology Letters 23, 902-918.
| Crossref | Google Scholar | PubMed |
Prugh LR, Stoner CJ, Epps CW, Bean WT, Ripple WJ, Laliberte AS, Brashares JS (2009) The rise of the mesopredator. BioScience 59, 779-791.
| Crossref | Google Scholar |
R Core Team (2023) ‘R: a language and environment for statistical computing. Version 4.2.3.’ (R Foundation for Statistical Computing: Vienna, Austria) Available at https://www.R-project.org/
Ramesh T, Kalle R, Sankar K, Qureshi Q (2012) Spatio-temporal partitioning among large carnivores in relation to major prey species in western Ghats. Journal of Zoology 287, 269-275.
| Crossref | Google Scholar |
Ramesh T, Kalle R, Rosenlund H, Downs CT (2016) Native habitat and protected area size matters: preserving mammalian assemblages in the maputaland conservation unit of South Africa. Forest Ecology and Management 360, 20-29.
| Crossref | Google Scholar |
Rayan DM, Linkie M (2016) Managing conservation flagship species in competition: tiger, leopard and dhole in Malaysia. Biological Conservation 204, 360-366.
| Crossref | Google Scholar |
Ridout MS, Linkie M (2009) Estimating overlap of daily activity patterns from camera trap data. Journal of Agricultural, Biological, and Environmental Statistics 14, 322-337.
| Crossref | Google Scholar |
Ritchie EG, Johnson CN (2009) Predator interactions, mesopredator release and biodiversity conservation. Ecology Letters 12, 982-998.
| Crossref | Google Scholar | PubMed |
Sehgal JJ, Panda D, Kumar D, Kalsi R, Allen ML, Singh R (2023) Does the size of a protected area matter? An assessment of leopard population and habitat usage in a protected area of Shiwalik foothills, Himalaya. Mammal Research 68, 459-469.
| Crossref | Google Scholar |
Seidensticker J (1976) On the ecological separation between tigers and leopards. Biotropica 8, 225-234.
| Crossref | Google Scholar |
Simcharoen A, Simcharoen S, Duangchantrasiri S, Bump J, Smith JLD (2018) Tiger and leopard diets in western Thailand: evidence for overlap and potential consequences. Food Webs 15, e00085.
| Crossref | Google Scholar |
Sollmann R, Mohamed A, Samejima H, Wilting A (2013) Risky business or simple solution – relative abundance indices from camera-trapping. Biological Conservation 159, 405-412.
| Crossref | Google Scholar |
Surve NS, Sathyakumar S, Sankar K, Jathanna D, Gupta V, Athreya V (2022) Leopards in the city: the Tale of Sanjay Gandhi National Park and Tungareshwar Wildlife Sanctuary, Two Protected Areas in and Adjacent to Mumbai, India. Frontiers in Conservation Science 3, 787031.
| Crossref | Google Scholar |
Symonds MRE, Moussalli A (2011) A brief guide to model selection, multimodel inference and model averaging in behavioural ecology using Akaike’s information criterion. Behavioral Ecology and Sociobiology 65, 13-21.
| Crossref | Google Scholar |
Taylor RA, Ryan SJ, Brashares JS, Johnson LR (2016) Hunting, food subsidies, and mesopredator release: the dynamics of crop-raiding baboons in a managed landscape. Ecology 97, 951-960.
| Crossref | Google Scholar | PubMed |
Teichman KJ, Cristescu B, Nielsen SE (2013) Does sex matter? temporal and spatial patterns of cougar–human conflict in British Columbia. PLoS ONE 8, e74663.
| Crossref | Google Scholar | PubMed |
Turner A (1990) The evolution of the guild of larger terrestrial carnivores during the Plio-Pleistocene in Africa. Geobios 23, 349-368.
| Crossref | Google Scholar |
Van Valkenburgh B, Hayward MW, Ripple WJ, Meloro C, Roth VL (2016) The impact of large terrestrial carnivores on Pleistocene ecosystems. Proceedings of the National Academy of Sciences 113, 862-867.
| Crossref | Google Scholar |
Vijayan S, Pati BP (2002) Impact of changing cropping patterns on man-animal conflicts around Gir Protected Area with specific reference to Talala Sub-District, Gujarat, India. Population and Environment 23, 541-559.
| Crossref | Google Scholar |
Wegge P, Yadav SK, Lamichhane BR (2018) Are corridors good for tigers Panthera tigris but bad for people? An assessment of the Khata Corridor in lowland Nepal. Oryx 52, 35-45.
| Crossref | Google Scholar |
Williams ST, Williams KS, Lewis BP, Hill RA (2017) Population dynamics and threats to an apex predator outside protected areas: implications for carnivore management. Royal Society Open Science 4, 161090.
| Crossref | Google Scholar | PubMed |
Zuur AF, Ieno EN, Elphick CS (2010) A protocol for data exploration to avoid common statistical problems. Methods in Ecology and Evolution 1, 3-14.
| Crossref | Google Scholar |
