Tiger habitat use dynamics in Bardia-Banke Complex, Nepal using camera trap-based multi-season occupancy framework
Abstract
Understanding fine-scale habitat use dynamics within species home range is crucial for long term conservation planning. We used multi-season occupancy models incorporating field and landscape level covariates to tiger camera trap data collected during the systematic periodic survey from 2013 to 2022. Habitat use probabilities varied across primary sessions, peaking at 0.56 in 2013 and dropping to 0.49 in 2022, indicating negative and positive rate of change in tiger habitat use probabilities (average lambda λ2013-2018 = 0.60, - 40% decline; average lambda λ2018-2022 = 2.28, +128% increase) between survey period 2013–2018 and 2218–2022 respectively. Local colonization probabilities fluctuated between 0.39 (SE = 0.09) during 2013–2018 and 0.48 (SE = 0.04) during 2018–2022, while local extinction remained stable at 0.50 (SE = 0.005), suggesting high site turnover. The effects of covariates on tiger habitat use dynamics varied over time. Tiger habitat use probabilities increased with proximity to waterholes and distance from settlements and decreased with elevation. However, the influence of habitat productivity and prey availability on tiger habitat use was contrary to our expectations. Colonization probability increased with prey availability and proximity to waterholes. Habitat productivity positively influenced local extinction, contrary to a priori expectations. Our findings highlight the critical role of waterhole distribution in shaping tiger habitat use in water-limited landscape. Enhancing water and prey availability can support colonization, promoting long-term persistence of tigers. Tiger habitat use dynamics highlight the importance of targeted, site-specific conservation strategies aimed at enhancing habitat suitability and promoting landscape connectivity. We emphasize the importance of long-term, multi-year camera trap monitoring to track persistence of tigers across the Complex. The study’s combination of estimated tiger habitat use and grid-based approaches offers a valuable framework for implementing targeted conservation interventions.
References
Abade L., Cusack J., Moll R.J., Strampelli P., Dickman A.J., Macdonald D.W., et al. 2020. The relative effects of prey availability, anthropogenic pressure and environmental variables on lion (Panthera leo) site use in Tanzania's Ruaha landscape during the dry season. Journal of Zoology, 310(2): 135‒144. https://doi.org/10.1111/jzo.12731
Barber‐Meyer S.M., Jnawali S.R., Karki J.B., Khanal P., Lohani S., Long B., et al. 2013. Influence of prey depletion and human disturbance on tiger occupancy in Nepal. Journal of Zoology, 289(1): 10‒18. https://doi.org/10.1111/j.1469-7998.2012.00956.x
Burnham K.P. and Anderson D.R. 2002. Model Selection and Multimodel Inference: A practical Information - Theoretic Approach. Springer, New York.
Carter N.H., Shrestha B.K., Karki J.B., Pradhan N.M.B. and Liu J. 2012. Coexistence between wildlife and humans at fine spatial scales. Proceedings of the National Academy of Sciences, 109(38): 15360‒15365. https://doi.org/10.1073/pnas.1210490109
Carter N., Killion A., Easter T., Brandt J. and Ford A. 2020. Road development in Asia: Assessing the range-wide risks to tigers. Science Advances, 6(18), eaaz9619. https://doi.10.1126/sciadv.aaz9619
CBS Nepal. 2022. Nepal Census 2021. Central Bureau of Statistics. https:// censusnepal.cbs.gov. np/ Home/ Detai ls? tpid= 5& dcid= 3479c092-7749-4ba6-9369-45486cd67f 30& tfsid=17.
Cimatti M., Ranc N., Benítez‐López A., Maiorano L., Boitani L., Cagnacci F., et al. 2021. Large carnivore expansion in Europe is associated with human population density and land cover changes. Diversity and Distributions, 27(4): 602‒617. https://doi.org/10.1111/ddi.13219
Cristescu B., Domokos C., Teichman K.J. and Nielsen S.E. 2019. Large carnivore habitat suitability modelling for Romania and associated predictions for protected areas. PeerJ, 7, e6549. https://doi.org/10.7717/peerj.6549
Devarajan K., Morelli T.L. and Tenan S. 2020. Multi‐species occupancy models: Review, roadmap, and recommendations. Ecography, 43(11): 1612‒1624. https://doi.org/10.1111/ecog.04957
Di Bitetti, M.S. Paviolo A. and De Angelo C. 2014. Camera trap photographic rates on roads vs. off roads: location does matter. Mastozoología neotropical, 21(1): 37‒46, Mendoza. http://www.sarem.org.ar
DNPWC 2018. Annual report. Department of National Parks and Wildlife Conservation, Kathmandu, Nepal.
DNPWC 2019. Annual report. Department of National Parks and Wildlife Conservation, Kathmandu, Nepal.
DNPWC and DFSC. 2022. Status of Tigers and Prey in Nepal. Department of National Parks and Wildlife Conservation and Department of Forests and Soil Conservation. Ministry of Forests and Environment, Kathmandu, Nepal.
Dupont, W. D. (2002). Statistical modeling for biomedical researchers. Cambridge University Press, Cambridge
Essl F., Dullinger S., Rabitsch W., Hulme P.E., Pyšek P., Wilson J.R., et al. 2015. Historical legacies accumulate to shape future biodiversity in an era of rapid global change. Diversity and Distributions, 21(5): 534‒547. https://doi.org/10.1111/ddi.12312
Everatt K.T., Moore J.F. and Kerley G.I. 2019. Africa's apex predator, the lion, is limited by interference and exploitative competition with humans. Global Ecology and Conservation, 20, e00758. https://doi.org/10.1016/j.gecco.2019.e00758
Farris Z.J., Kelly M.J., Karpanty S. and Ratelolahy F. 2016. Patterns of spatial co‐occurrence among native and exotic carnivores in north‐eastern Madagascar. Animal Conservation, 19(2): 189‒198. https://doi.org/10.1111/acv.12233
Fasona M.I., Gregory A.J., Okimiji P.O. Oni and Egonmwan R.I. 2024. Unravelling the dynamics: Large mammal occupancy modelling in a West African tropical forest. African Journal of Ecology, 62(2), e13269. https://doi.org/10.1111/aje.13269
Feng J., Sun Y., Li H., Xiao Y., Zhang D., Smith J.L.D., et al. 2021. Assessing mammal species richness and occupancy in a Northeast Asian temperate forest shared by cattle. Diversity and Distributions, 27(5): 857‒872. https://doi.org/10.1111/ddi.13237
Garcia R.A., Cabeza M., Rahbek C. and Araújo M.B. 2014. Multiple dimensions of climate change and their implications for biodiversity. Science, 344(6183): 1247579. https://doi.org/10.1126/science.1247579
Goodrich J., Sanderson A.J., Chapman E., Gray S., Chanchani T.N.E., Harihar P., et al. 2022. Panthera tigris. The IUCN Red List of Threatened Species. https://doi.org/10.2305/IUCN.UK. 2022.
Gray T.N., Rosenbaum R., Jiang G., Izquierdo P., Yongchao J.I.N., Kesaro, L., et al. 2023. Restoring Asia’s roar: Opportunities for tiger recovery across the historic range. Frontiers in Conservation Science, 4: 1124340. https://doi.org/10.3389/fcosc.2023.1124340
Harihar A. and Pandav B. 2012. Influence of connectivity, wild prey and disturbance on occupancy of tigers in the human-dominated western Terai Arc Landscape. PloS one, 7(7), e40105. https://doi.org/10.1371/journal.pone.0040105
Hines J.E. 2006. PRESENCE3 - Software to estimate patch occupancy and related parameters. USGS - PWRC. http://www.mbr-pwrc.usgs.gov/software/presence.html
Hines J.E., Nichols J.D. and Collazo J.A. 2014. Multiseason occupancy models for correlated replicate surveys. Methods in Ecology and Evolution, 5(6): 583‒591. https://doi.org/10.1111/2041-210X.12186
Jhala Y.V., Mungi N.A., Gopal R. and Qureshi Q. 2025. Tiger recovery amid people and poverty. Science, 387(6733): 505‒515. https://doi.org/10.1126/science.adk4827
Kafley H., Gompper M.E., Sharma M., Lamichane B.R. and Maharjan R. 2016. Tigers (Panthera tigris) respond to fine spatial-scale habitat factors: occupancy-based habitat association of tigers in Chitwan National Park, Nepal. Wildlife Research, 43(5): 398‒410. https://doi.org/10.1071/WR16012
Karanth K.K., Nichols J.D., Hines J.E., Karanth K.U. and Christensen N.L. 2009. Patterns and determinants of mammal species occurrence in India. Journal of Applied Ecology, 46(6): 1189‒1200. https://doi.org/10.1111/j.1365-2664.2009.01710.x
Karanth K.U. and Nichols J.D. 2011. Estimating tiger abundance from camera trap data: field surveys and analytical issues. Camera traps in animal ecology: methods and analyses, 97‒117. https://doi.org/10.1111/cobi.13517
Karanth K.U., Nichols J.D., Kumar N.S., Link W.A. and Hines J.E. 2004. Tigers and their prey: predicting carnivore densities from prey abundance. Proceeding. Natl. Acad. Sci. USA 101, 4854–4858. https://doi.org/10.1073/pnas.0306210101.
Kshettry A., Vaidyanathan S. and Athreya V. 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. https://doi.org/10.1371/journal.pone.0177013
Kuussaari M., Bommarco R., Heikkinen R.K., Helm A., Krauss J., Lindborg R., Öckinger E, et al. 2009. Extinction debt: a challenge for biodiversity conservation. Trends in ecology & evolution, 24(10): 564‒571. https://doi.org/10.1016/j.tree.2009.04.011
Lamichhane B.R., Lamichhane S., Regmi R., Dhungana M., Thapa S.K., Prasai A., et al. 2021. Leopard (Panthera pardus) occupancy in the Chure range of Nepal. Ecology and Evolution, 11(20): 13641‒13660. https://doi.org/10.1002/ece3.8105
MacKenzie D.I. 2011. Monitoring carnivore populations at the landscape scale: occupancy modelling of tigers from sign surveys. Journal of Applied Ecology, 48(4): 1048‒1056.
https://doi.org/10.1111/j.1365-2664.2011.02002.x
MacKenzie D.I., Nichols J.D., Hines J.E., Knutson M.G. and Franklin A.B. 2003. Estimating site occupancy, colonization, and local extinction when a species is detected imperfectly. Ecology, 84(8): 2200‒2207. https://doi.org/10.1890/02-3090
MacKenzie D.I., Nichols J.D., Royle J.A., Pollock K.H., Bailey L. and Hines J.E. 2006. Occupancy estimation and modeling: inferring patterns and dynamics of species occurrence. Elsevier, San Diego. https://doi.org/10.1016/C2012-0-01164-7
MacKenzie D.I., Nichols J.D., Royle J.A., Pollock K.H., Bailey L.L. and Hines J.E. 2018. Occupancy estimation and modeling: Inferring patterns and dynamics of species occurrence (2nd ed.). London, UK: Academic Press. https://doi.org/10.1016/C2012-0-01164-7
Magurran A.E. and Henderson P.A. 2010. Temporal turnover and the maintenance of diversity in ecological assemblages. Philosophical Transactions of the Royal Society B: Biological Sciences, 365(1558): 3611‒3620. https://doi.org/10.1098/rstb.2010.0285
Mondal K., Sankar K. and Qureshi Q. 2013. Factors influencing the distribution of leopard in a semiarid landscape of Western India. Acta Theriologica, 58: 179‒187. https://doi.org/10.1007/s13364-012-0109-6
Oriol‐Cotterill A., Valeix M., Frank L.G., Riginos C. and Macdonald D.W. 2015. Landscapes of coexistence for terrestrial carnivores: the ecological consequences of being downgraded from ultimate to penultimate predator by humans. Oikos, 124(10): 1263‒1273. https://doi.org/10.1111/oik.02224
Oswell A. 2010. The big cat trade in Myanmar and Thailand. https://www.traffic.org/site/assets/files/2735/big_cat_trade_in_myanmar_report.pdf
Pettorelli N., Ryan S., Mueller T., Bunnefeld N., Jędrzejewska B., Lima M., et al. 2011. The Normalized Difference Vegetation Index (NDVI): unforeseen successes in animal ecology. Climate research, 46(1): 15‒27. https://doi.org/10.3354/cr00936
Qi J., Gu J., Ning Y., Miquelle D.G., Holyoak M., Wen D., et al. 2021. Integrated assessments call for establishing a sustainable meta-population of Amur tigers in northeast Asia. Biological Conservation, 261: 109‒250. https://doi.org/10.1016/j.biocon.2021.109250
Ram A.K., Lamichhane B.R. Subedi, N. Yadav, N.K. Karki A., Pandav B., et al. 2024. Dynamic occupancy modelling of Asian elephants (Elephas maximus) reveals increasing landscape use in Nepal. Scientific Reports, 14(1): 20023. https://doi.org/10.1038/s41598-024-70092-4
Riley S.J., DeGloria S.D. and Elliot R. 1999. Index that quantifies topographic heterogeneity. intermountain Journal of sciences, 5(1-4): 23‒27. https://download.osgeo.org/qgis/doc/reference-docs/Terrain_Ruggedness_Index.pdf
Rosenblatt E., Becker M.S., Creel S., Droge E., Mweetwa T., Schuette P.A., et al. 2014. Detecting declines of apex carnivores and evaluating their causes: An example with Zambian lions. Biological Conservation, 180: 176‒186. https://doi.org/10.1016/j.biocon.2014.10.006
Rota C.T., Ferreira M.A., Kays R.W., Forrester T.D., Kalies E.L., McShea W.J., et al. 2016. A multispecies occupancy model for two or more interacting species. Methods in Ecology and Evolution, 7(10): 1164‒1173. https://doi.org/10.1111/2041-210X.12587
Rybicki J. and Hanski I. 2013. Species–area relationships and extinctions caused by habitat loss and fragmentation. Ecology letters, 16: 27‒38.https://doi.org/10.1111/ele.12065
Sanderson E.W., Miquelle D.G., Fisher K., Harihar A., Clark C., Moy J., et al. 2023. Range-wide trends in tiger conservation landscapes, 2001-2020. Frontiers in Conservation Science, 4: 1191280. https://doi.org/10.3389/fcosc.2023.1191280
Shah S.K., Karki J.B., Bhatta B., Subedi N., Bahadur K.R., Kadariya R., et al. 2025. Ecological factors at fine spatial scale associated with habitat use by tigers in Western Terai Arc Landscape, Nepal. Ecology and Evolution, ece3.71109.
https://doi.org/10.1002/ece3.71109
Shah S.K., Karki J.B., Bhatta B., Subedi N., Lamichhane B.R., KC R.B., et al. 2024. Recovery of tigers (Panthera tigris): Assessing ecological carrying capacity in The Bardia-Banke Complex, Nepal. Global Ecology and Conservation, 56: e03326. https://doi.org/10.1016/j.gecco.2024.e03326
Sleeman J.M. 2013. Has the time come for big science in wildlife health? Ecological Health, 10(4): 335‒338. https://doi.org/10.1007/s10393-013-0880-0
Srivathsa A, Karanth K.K., Jathanna D, Kumar N.S. and Karanth K.U. 2014. On a Dhole Trail: Examining Ecological and Anthropogenic Correlates of Dhole Habitat Occupancy in the Western Ghats of India. PLoS ONE 9(6): e98803. https://doi.org/10.1371/journal.pone.0098803
Sunarto S., Kelly M.J., Parakkasi K., Klenzendorf S., Septayuda E. and Kurniawan H. 2012. Tigers need cover: multi-scale occupancy study of the big cat in Sumatran forest and plantation landscapes. Plos one, 7(1), e30859. https://doi.org/10.1371/journal.pone.0030859
Thapa K. and Kelly M.J. 2016. Prey and tigers on the forgotten trail: high prey occupancy and tiger habitat use reveal the importance of the understudied Churia habitat of Nepal. Biodiversity and Conservation, 26: 593‒616. https://doi.org/10.1007/s10531-016-1260-1
Thapa K., Wikramanayake E., Malla S., Acharya K.P., Lamichhane B.R., Subedi N., et al. 2017. Tigers in the Terai: Strong evidence for meta-population dynamics contributing to tiger recovery and conservation in the Terai Arc Landscape. PLoS One, 12(6): e0177548. https://doi.org/10.1371/journal.pone.0177548
Thapa K. and Kelly M.J. 2017. Density and carrying capacity in the forgotten tigerland: Tigers in the understudied Nepalese Churia. Integrative Zoology, 12(3): 211‒227.
https://doi.org/10.1111/1749-4877.12240
Thapa K., Kelly M.J. and Pradhan N.M.B. 2019. Elephant (Elephas maximus) temporal activity, distribution, and habitat use patterns on the tiger’s forgotten trails across the seasonally dry, subtropical, hilly Churia forests of Nepal. PLoS One, 14(5): e0216504. https://doi.org/10.1371/journal.pone.0216504
Thapa K., Malla S., Subba S.A., Thapa G.J., Lamichhane B.R., Subedi N., et al. 2021. On the tiger trails: Leopard occupancy decline and leopard interaction with tigers in the forested habitat across the Terai Arc Landscape of Nepal. Global Ecology and Conservation, 25: p.e01412. https://doi.org/10.1016/j.gecco.2020.e01412
Thapa K., Subba S.A., Thapa G.J., Dewan K., Acharya B.P., Bohara D., et al. 2022. Wildlife in climate refugia: Mammalian diversity, occupancy, and tiger distribution in the Western Himalayas, Nepal. Ecology and Evolution, 12(12): e9600. https://doi.org/10.1002/ece3.9600
Van der Weyde L.K., Mbisana C. and Klein R. 2018. Multi-species occupancy modelling of a carnivore guild in wildlife management areas in the Kalahari. Biological Conservation, 220: 21‒28. https://doi.org/10.1016/j.biocon.2018.01.033
Walston J., Robinson J.G., Bennett E.L., Breitenmoser U., da Fonseca G.A.B., Goodrich J., et. al. 2010. Bringing the tiger back from the brink-the six percent solution. PLoS Biol. 8. https://doi.org/10.1371/journal.pbio.1000485.
Warrier R., Noon B.R. and Bailey L. 2020. Agricultural lands offer seasonal habitats to tigers in a human‐dominated and fragmented landscape in India. Ecosphere, 11(7): e03080. https://doi.org/10.1002/ecs2.3080
Wearn O.R., Reuman D.C. and Ewers R.M. 2012. Extinction debt and windows of conservation opportunity in the Brazilian Amazon. Science, 337(6091): 228‒232. https://doi.org/10.1126/science.1231438
Wegge P. and Storaas T. 2009. Sampling tiger ungulate prey by the distance method: lessons learned in Bardia National Park, Nepal. Animal Conservation, 12: 78–84. https:// doi.org/10.1111/j.1469-1795.2008.00230.x.
Wolf C. and Ripple W.J. 2017. Range contractions of the world's large carnivores. Royal Society open science, 4(7): 170052. https://doi.org/10.1098/rsos.170052. https://doi.org/10.1098/rsos.170052
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