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Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

Ensemble forecasting of Persian leopard (Panthera pardus saxicolor) distribution and habitat suitability in south-western Iran

Fatemeh Jamali A , Seyed Mehdi Amininasab https://orcid.org/0000-0002-4863-4606 B * , Hamid Taleshi https://orcid.org/0009-0007-0646-3785 A and Hossein Madadi https://orcid.org/0000-0002-8909-9427 A
+ Author Affiliations
- Author Affiliations

A Faculty of Natural Resources, Behbahan Khatam Alanbia University of Technology, Behbahan, Iran.

B Faculty of Natural Resources, Sari Agricultural Sciences and Natural Resources University, Sari, Iran.

* Correspondence to: smamininasab@yahoo.com

Handling Editor: Adam Stow

Wildlife Research 51, WR23010 https://doi.org/10.1071/WR23010
Submitted: 28 March 2022  Accepted: 25 February 2024  Published: 18 March 2024

© 2024 The Author(s) (or their employer(s)). Published by CSIRO Publishing

Abstract

Context

Habitat loss and fragmentation are major threats to global biodiversity, especially for wide-ranging apex carnivores. The Persian leopard (Panthera pardus saxicolor) is an endangered species. Although populations of Persian leopards are declining, the species remains widely distributed across large areas of Iran.

Aims

This study aimed to determine habitat suitability for Persian leopards in the Khaeiz and Sorkh protected area of south-western Iran and to identify the most significant factors influencing their habitat use/selection and spatial distribution.

Methods

We performed species distribution modelling in two stages: First, we ran the model with three abiotic predictors: slope; aspect; and distance from water resources. In the second stage, modelling was conducted using three ecological predictors: caracal distribution; wild goat distribution; and livestock distribution. Ensemble modelling was applied based on five replicates of eight SDMs (species distribution models; GLM, CTA, FDA, GBM, ANN, MARS, RF and MaxEnt).

Key results

We observed only minor differences in habitat suitability between the abiotic and ecological models. Habitat suitability for Persian leopards was higher in steeper areas, close to water resources and near the distribution of caracals, livestock and wild goats. The ecological model predicted 2.03% (329 ha) more suitable habitat than the abiotic model did.

Conclusions

Most habitat suitability models focus on abiotic variables, but we found that ecological variables offer similar predictive power for determining the habitat suitability of Persian leopards.

Implications

Habitat suitability models for Persian leopards can be used to guide conservation and management decisions. They are also useful indicating where conflicts between predators and humans may occur.

Keywords: caracal, conservation, ensemble forecasting, felids, human–wildlife conflict, livestock, Panthera pardus saxicolor, Persian leopard, SDM (species distribution model), south-western Iran, wild goat.

References

Abade L, Cusack J, Moll RJ, Strampelli P, Dickman AJ, Macdonald DW, Montgomery RA (2018) Spatial variation in leopard (Panthera pardus) site use across a gradient of anthropogenic pressure in Tanzania’s Ruaha landscape. PLoS ONE 13(10), e0204370.
| Crossref | Google Scholar | PubMed |

Acharya KP, Paudel PK, Jnawali SR, Neupane PR, Köhl M (2017) Can forest fragmentation and configuration work as indicators of human–wildlife conflict? Evidences from human death and injury by wildlife attacks in Nepal. Ecological Indicators 80, 74-83.
| Crossref | Google Scholar |

Adibi MA, Karami M, Kaboli M (2014) Study of seasonal changes in habitat suitability of Caracal caracal schmitzi (Maschie 1812) in the central desert of Iran. Journal of Biodiversity and Environmental Sciences 5(3), 95-106.
| Google Scholar |

Ahmadi M, Farhadinia MS, Cushman SA, Hemami M-R, Nezami Balouchi B, Jowkar H, Macdonald DW (2020) Species and space: a combined gap analysis to guide management planning of conservation areas. Landscape Ecology 35(7), 1505-1517.
| Crossref | Google Scholar |

Allouche O, Tsoar A, Kadmon R (2006) Assessing the accuracy of species distribution models: prevalence, kappa and the true skill statistic (TSS). Journal of Applied Ecology 43(6), 1223-1232.
| Google Scholar |

Amininasab SM, Zamani N, Taleshi H, Xu CCY (2023) Ensemble modelling the distribution and habitat suitability of wild goat Capra aegagrus in southwestern Iran. Biodiversity 24(3), 124-136.
| Crossref | Google Scholar |

Ansari M, Ghoddousi A (2018) Water availability limits brown bear distribution at the southern edge of its global range. Ursus 29(1), 13-24.
| Crossref | Google Scholar |

Arvin A, Ghangherme A, Hajipour D, Heidari M (2016) Investigating the trend of changes in some climatic elements in Chaharmahal and Bakhtiari Province. Journal of Geographical Sciences 16, 153-176 [In Persian with English abstract].
| Google Scholar |

Ashrafzadeh MR, Naghipour AA, Haidarian M, Khorozyan I (2019) Modeling the response of an endangered flagship predator to climate change in Iran. Mammal Research 64, 39-51.
| Crossref | Google Scholar |

Ashrafzadeh MR, Khosravi R, Adibi MA, Taktehrani A, Wan HY, Cushman SA (2020) A multi-scale, multi-species approach for assessing effectiveness of habitat and connectivity conservation for endangered felids. Biological Conservation 245, 108523.
| Crossref | Google Scholar |

Athreya V, Odden M, Linnell JDC, Krishnaswamy J, Karanth U (2013) Big cats in our backyards: persistence of large carnivores in a human dominated landscape in India. PLoS ONE 8(3), e57872.
| Crossref | Google Scholar | PubMed |

Babrgir S, Farhadinia MS, Moqanaki EM (2017) Socio-economic consequences of cattle predation by the endangered Persian leopard Panthera pardus saxicolor in a Caucasian conflict hotspot, northern Iran. Oryx 51, 124-130.
| Crossref | Google Scholar |

Baker AD, Leberg PL (2018) Impacts of human recreation on carnivores in protected areas. PLoS ONE 13, e0195436.
| Crossref | Google Scholar | PubMed |

Barbet-Massin M, Jiguet F, Albert CH, Thuiller W (2012) Selecting pseudo-absences for species distribution models: how, where and how many? Methods in Ecology and Evolution 3, 327-338.
| Crossref | Google Scholar |

Beier P, Majka D, Jenness J (2007) Conceptual steps for designing wildlife corridors. Available at https://www.corridordesign.org/ [accessed 10 June 2019]

Beier P, Majka DR, Spencer WD (2008) Forks in the road: choices in procedures for designing wildland linkages. Conservation Biology 22(4), 836-851.
| Crossref | Google Scholar | PubMed |

Breiman L (1996) Bagging predictors. Machine Learning 24(2), 123-140.
| Crossref | Google Scholar |

Breiman L, Friedman J, Stone CJ, Olshen RA (1984) ‘Classification and regression trees.’ (CRC press)

Breiner FT, Nobis MP, Bergamini A, Guisan A (2018) Optimizing ensembles of small models for predicting the distribution of species with few occurrences. Methods in Ecology and Evolution 9, 802-808.
| Crossref | Google Scholar |

Calvignac S, Hughes S, Hӓnni C (2009) Genetic diversity of endangered brown bear (Ursus arctos) populations at the crossroads of Europe, Asia and Africa. Diversity and Distributions 15, 742-750.
| Crossref | Google Scholar |

Carvalho SB, Brito JC, Crespo EG, Watts ME, Possingham HP (2011) Conservation planning under climate change: toward accounting for uncertainty in predicted species distributions to increase confidence in conservation investments in space and time. Biological Conservation 144, 2020-2030.
| Crossref | Google Scholar |

Carvalho EAR, Jr., Zarco-González MM, Monroy-Vilchis O, Morato RG (2015) Modeling the risk of livestock depredation by jaguar along the Transamazon highway, Brazil. Basic and Applied Ecology 16(5), 413-419.
| Crossref | Google Scholar |

Chala D, Brochmann C, Psomas A, Ehrich D, Gizaw A, Masao CA, Bakkestuen V, Zimmermann NE (2016) Good-bye to tropical alpine plant giants under warmer climates? Loss of range and genetic diversity in Lobelia rhynchopetalum. Ecology and Evolution 6, 8931-8941.
| Crossref | Google Scholar | PubMed |

Chame M (2003) Terrestrial mammal feces: a morphometric summary and description. Memórias do Instituto Oswaldo Cruz 98, 71-94.
| Crossref | Google Scholar | PubMed |

Cheraghi F, Delavar MR, Amiraslani F, Alavipanah K, Gurarie E, Jowkar H, Hunter L, Ostrowski S, Fagan WF (2019) Inter-dependent movements of Asiatic Cheetahs Acinonyx jubatus venaticus and a Persian Leopard Panthera pardus saxicolor in a desert environment in Iran (Mammalia: Felidae). Zoology in the Middle East 65, 283-292.
| Crossref | Google Scholar |

Cohen J (1960) A coefficient of agreement for nominal scales. Educational and Psychological Measurement 20, 37-46.
| Crossref | Google Scholar |

Connor T, Hull V, Viña A, Shortridge A, Tang Y, Zhang J, Wang F, Liu J (2018) Effects of grain size and niche breadth on species distribution modeling. Ecography 41(8), 1270-1282.
| Crossref | Google Scholar |

Crooks KR (2002) Relative sensitivities of mammalian carnivores to habitat fragmentation. Conservation Biology 16, 488-502.
| Crossref | Google Scholar |

Cruz P, Iezzi ME, De Angelo C, Varela D, Di Bitetti MS, Paviolo A (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.
| Crossref | Google Scholar |

Decœur H, Amir Z, Mendes CP, Moore JH, Luskin MS (2023) Mid-sized felids threatened by habitat degradation in Southeast Asia. Biological Conservation 283, 110103.
| Crossref | Google Scholar |

D’heygere T, Goethals PLM, De Pauw N (2006) Genetic algorithms for optimisation of predictive ecosystems models based on decision trees and neural networks. Ecological Modelling 195, 20-29.
| Crossref | Google Scholar |

Ebrahimi A, Farashi A, Rashki A (2017) Habitat suitability of Persian leopard (Panthera pardus saxicolor) in Iran in future. Environmental Earth Sciences 76, 697.
| Crossref | Google Scholar |

Ebrahimi E, Sayahnia R, Ranjbaran Y, Vaissi S, Ahmadzadeh F (2021) Dynamics of threatened mammalian distribution in Iran’s protected areas under climate change. Mammalian Biology - Zeitschrift fur Saugetierkunde 101, 759-774.
| Crossref | Google Scholar |

Elith J, Kearney M, Phillips S (2010) The art of modelling range-shifting species. Methods in Ecology and Evolution 1, 330-342.
| Crossref | Google Scholar |

Erfanian B, Mirkarimi SH, Mahini AS, Rezaei HR (2013) A presence-only habitat suitability model for Persian leopard Panthera pardus saxicolor in Golestan National Park, Iran. Wildlife Biology 19, 170-178.
| Crossref | Google Scholar |

Farashi A, Shariati M, Hosseini M (2017) Identifying biodiversity hotspots for threatened mammal species in Iran. Mammalian Biology 87, 71-88.
| Crossref | Google Scholar |

Farhadinia MS, Ahmadi M, Sharbafi E, Khosravi S, Alinezhad H, Macdonald DW (2015) Leveraging trans-boundary conservation partnerships: persistence of Persian leopard (Panthera pardus saxicolor) in the Iranian Caucasus. Biological Conservation 191, 770-778.
| Crossref | Google Scholar |

Farhadinia MS, Johnson PJ, Hunter LTB, Macdonald DW (2018a) Persian leopard predation patterns and kill rates in the Iran-Turkmenistan borderland. Journal of Mammalogy 99, 713-723.
| Crossref | Google Scholar |

Farhadinia MS, Johnson PJ, Macdonald DW, Hunter LTB (2018b) Anchoring and adjusting amidst humans: ranging behavior of Persian leopards along the Iran-Turkmenistan borderland. PLoS ONE 13, e0196602.
| Crossref | Google Scholar | PubMed |

Farhadinia MS, Moll RJ, Montgomery RA, Ashrafi S, Johnson PJ, Hunter LTB, MacDonald DW (2018c) Citizen science data facilitate monitoring of rare large carnivores in remote montane landscapes. Ecological Indicators 94, 283-291.
| Crossref | Google Scholar |

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 |

Friedman JH (1991) Multivariate adaptive regression splines. The Annals of Statistics 19(1), 1-67.
| Crossref | Google Scholar |

Friedman JH (2001) Greedy function approximation: a gradient boosting machine. The Annals of Statistics 29(5), 1189-1232.
| Crossref | Google Scholar |

Gavashelishvili A, Lukarevskiy V (2008) Modelling the habitat requirements of leopard Panthera pardus in west and central Asia. Journal of Applied Ecology 45, 579-588.
| Crossref | Google Scholar |

Ghoddousi A, Kh. Hamidi A, Ghadirian T, Ashayeri D, Khorozyan I (2010) The status of the endangered Persian leopard Panthera pardus saxicolor in Bamu National Park, Iran. Oryx 44, 551-557.
| Crossref | Google Scholar |

Ghoddousi A, Soofi M, Kh. Hamidi A, Lumetsberger T, Egli L, Khorozyan I, Kiabi BH, Waltert M (2016) Assessing the role of livestock in big cat prey choice using spatiotemporal availability patterns. PLoS ONE 11(4), e0153439.
| Crossref | Google Scholar | PubMed |

Ghoddousi A, Bleyhl B, Sichau C, Ashayeri D, Moghadas P, Sepahvand P, Kh Hamidi A, Soofi M, Kuemmerle T (2020) Mapping connectivity and conflict risk to identify safe corridors for the Persian leopard. Landscape Ecology 35, 1809-1825.
| Crossref | Google Scholar |

Guisan A, Zimmermann NE (2000) Predictive habitat distribution models in ecology. Ecological Modelling 135, 147-186.
| Crossref | Google Scholar |

Hallman TA, Robinson WD (2020) Deciphering ecology from statistical artefacts: competing influence of sample size, prevalence and habitat specialization on species distribution models and how small evaluation datasets can inflate metrics of performance. Diversity and Distributions 26, 315-328.
| Crossref | Google Scholar |

Hamidi AK, Ghoddousi A, Soufi M, Ghadirian T, Jowkar H, Ashayeri S (2014) Camera trap study of Persian leopard in Golestan National Park, Iran. Cat News 60, 12-14.
| Google Scholar |

Harrell FE, Jr., Lee KL, Califf RM, Pryor DB, Rosati RA (1984) Regression modelling strategies for improved prognostic prediction. Statistics in Medicine 3, 143-152.
| Crossref | Google Scholar | PubMed |

Hastie T, Tibshirani R, Buja A (1994) Flexible discriminant analysis by optimal scoring. Journal of the American Statistical Association 89(428), 1255-1270.
| Crossref | Google Scholar |

Hearn AJ, Cushman SA, Ross J, Goossens B, Hunter LTB, Macdonald DW (2018) Spatio-temporal ecology of sympatric felids on Borneo. Evidence for resource partitioning? PLoS ONE 13, e0200828.
| Crossref | Google Scholar | PubMed |

Hebblewhite M, Miquelle DG, Murzin AA, Aramilev VV, Pikunov DG (2011) Predicting potential habitat and population size for reintroduction of the Far Eastern leopards in the Russian Far East. Biological Conservation 144, 2403-2413.
| Crossref | Google Scholar |

Hemami M-R, Esmaeili S, Brito JC, Ahmadi M, Omidi M, Martínez-Freiría F (2018) Using ecological models to explore niche partitioning within a guild of desert felids. Hystrix 29, 216-222.
| Crossref | Google Scholar |

IUCN (2012) IUCN red list of threatened species. Version 2013. Available at www.iucnredlist.org

Jackson CR, Power RJ, Groom RJ, Masenga EH, Mjingo EE, Fyumagwa RD, Røskaft E, Davies-Mostert H (2014) Heading for the hills: risk avoidance drives den site selection in African wild dogs. PLoS ONE 9(6), e99686.
| Crossref | Google Scholar | PubMed |

Jacobson AP, Gerngross P, Lemeris JR, Jr., Schoonover RF, Anco C, Breitenmoser-Würsten C, Durant SM, Farhadinia MS, Henschel P, Kamler JF, Laguardia A, Rostro-García S, Stein AB, Dollar L (2016) Leopard (Panthera pardus) status, distribution, and the research efforts across its range. PeerJ 4, e1974.
| Crossref | Google Scholar | PubMed |

Jafari A, Zamani-Ahmadmahmoodi R, Mirzaei R (2018) Persian leopard and wild sheep distribution modeling using the Maxent model in the Tang-e-Sayad protected area, Iran. Mammalia 83(1), 84-96.
| Crossref | Google Scholar |

Jooste EC (2020) Ecology and diet of the caracal (Caracal caracal) on lethal and non-lethal control farms in the Karoo. Master thesis, Department of Biodiversity and Conservation Biology, Faculty of Science, University of the Western Cape.

Kaboodvandpour S, Almasieh K, Zamani N (2021) Habitat suitability and connectivity implications for the conservation of the Persian leopard along the Iran–Iraq border. Ecology and Evolution 11, 13464-13474.
| Crossref | Google Scholar | PubMed |

Kadmon R, Farber O, Danin A (2003) A systematic analysis of factors affecting the performance of climatic envelope models. Ecological Applications 13, 853-867.
| Crossref | Google Scholar |

Karami M, Ghadirian T, Faizolahi K (2016) ‘The atlas of mammals of Iran.’ (Jahad Daneshgahi, kharazmi Branch: Iran)

Khorozyan IG, Malkhasyan AG, Asmaryan SG, Abramov AV (2010) Using geographical mapping and occupancy modeling to study the distribution of the critically endangered leopard (Panthera pardus) population in Armenia. In ‘Spatial complexity, informatics, and wildlife conservation’. (Eds SA Cushman, F Huettmann) pp. 331–347. (Springer: Tokyo)

Khorozyan I, Soofi M, Khaleghi Hamidi A, Ghoddousi A, Waltert M (2015a) Dissatisfaction with veterinary services is associated with leopard (Panthera pardus) predation on domestic animals. PLoS ONE 10(6), e0129221.
| Crossref | Google Scholar | PubMed |

Khorozyan I, Ghoddousi A, Soofi M, Waltert M (2015b) Big cats kill more livestock when wild prey reaches a minimum threshold. Biological Conservation 192, 268-275.
| Crossref | Google Scholar |

Khosravi R, Hemami M-R, Cushman SA (2018) Multispecies assessment of core areas and connectivity of desert carnivores in central Iran. Diversity and Distributions 24, 193-207.
| Crossref | Google Scholar |

Khosravi R, Hemami M-R, Cushman SA (2019) Multi-scale niche modeling of three sympatric felids of conservation importance in central Iran. Landscape Ecology 34, 2451-2467.
| Crossref | Google Scholar |

Khosravi R, Hemami M-R, Malakoutikhah S, Ashrafzadeh MR, Cushman SA (2021) Prey availability modulates predicted range contraction of two large felids in response to changing climate. Biological Conservation 255, 109018.
| Crossref | Google Scholar |

Kie JG (1999) Optimal foraging and risk of predation: effects on behavior and social structure in ungulates. Journal of Mammalogy 80, 1114-1129.
| Crossref | Google Scholar |

Krüger L, Ramos JA, Xavier JC, Grémillet D, González-Solís J, Petry MV, Phillips RA, Wanless RM, Paiva VH (2018) Projected distributions of Southern Ocean albatrosses, petrels and fisheries as a consequence of climatic change. Ecography 41, 195-208.
| Crossref | Google Scholar |

Kuijper DPJ, Sahlén E, Elmhagen B, Chamaillé-Jammes S, Sand H, Lone K, Cromsigt JPGM (2016) Paws without claws? Ecological effects of large carnivores in anthropogenic landscapes. Proceedings of the Royal Society B: Biological Sciences 283, 20161625.
| Crossref | Google Scholar |

Kunin WE, Gaston KJ (1993) The biology of rarity: patterns, causes and consequences. Trends in Ecology & Evolution 8, 298-301.
| Crossref | Google Scholar | PubMed |

Lewis JS, Farnsworth ML, Burdett CL, Theobald DM, Gray M, Miller RS (2017) Biotic and abiotic factors predicting the global distribution and population density of an invasive large mammal. Scientific Reports 7, 44152.
| Crossref | Google Scholar | PubMed |

Malakoutikhah S, Fakheran S, Hemami M-R, Tarkesh M, Senn J (2020) Assessing future distribution, suitability of corridors and efficiency of protected areas to conserve vulnerable ungulates under climate change. Diversity and Distributions 26, 1383-1396.
| Crossref | Google Scholar |

Mateo Sánchez MC, Cushman SA, Saura S (2014) Scale dependence in habitat selection: the case of the endangered brown bear (Ursus arctos) in the Cantabrian Range (NW Spain). International Journal of Geographical Information Science 28, 1531-1546.
| Crossref | Google Scholar |

Mazzolli M, Haag T, Lippert BG, Eizirik E, Hammer MLA, Al Hikmani K (2017) Multiple methods increase detection of large and medium-sized mammals: working with volunteers in south-eastern Oman. Oryx 51(2), 290-297.
| Crossref | Google Scholar |

McCullagh P, Nelder JA (1989) ‘Generalized linear models.’ (CRC Press)

Meller L, Cabeza M, Pironon S, Barbet-Massin M, Maiorano L, Georges D, Thuiller W (2014) Ensemble distribution models in conservation prioritization: from consensus predictions to consensus reserve networks. Diversity and Distributions 20(3), 309-321.
| Crossref | Google Scholar | PubMed |

Mi C, Huettmann F, Guo Y, Han X, Wen L (2017) Why choose Random Forest to predict rare species distribution with few samples in large undersampled areas? Three Asian crane species models provide supporting evidence. PeerJ 5, e2849.
| Crossref | Google Scholar | PubMed |

Michalski F, Boulhosa RLP, Faria A, Peres CA (2006) Human–wildlife conflicts in a fragmented Amazonian forest landscape: determinants of large felid depredation on livestock. Animal Conservation 9(2), 179-188.
| Crossref | Google Scholar |

Mohammadi A, Almasieh K, Nayeri D, Ataei F, Khani A, López-Bao JV, Penteriani V, Cushman SA (2021) Identifying priority core habitats and corridors for effective conservation of brown bears in Iran. Scientific Reports 11, 1044.
| Crossref | Google Scholar | PubMed |

Momeni S, Malekian M, Hemami M-R (2019) Molecular versus morphological approaches to diet analysis of the caracal (Caracal caracal). Mammalia 83(6), 586-592.
| Crossref | Google Scholar |

Mondal K (2011) Ecology of leopard (Panthera pardus) in Sariska tiger reserve, Rajashtan. PhD thesis, Saurashtra University, India. p. 200.

Moradi AS, Ahmadi A, Toranjzar H, Shams-Esfandabad B (2022) Modeling the Habitat Suitability of Persian Leopard (Panthera Pardus Saxicolor) in the Conservation Areas of Kohgiluyeh and Boyer-Ahmad Province, Iran. Ecopersia 10(2), 109-119.
| Google Scholar |

Naderi GR, Riazi B, Aref N, Khalatbari M, Mohammadi S, Lahoot M, Kamran M (2013) Habitat preferences of Bezoar wild goats (Capra aegagrus) in Agh-Dagh protected area, Iran. North-Western Journal of Zoology 9(1), 99-102.
| Google Scholar |

Naderi M, Farashi A, Erdi MA (2018) Persian leopard’s (Panthera pardus saxicolor) unnatural mortality factors analysis in Iran. PLoS ONE 13, e0195387.
| Crossref | Google Scholar | PubMed |

Nezer O, Bar-David S, Gueta T, Carmel Y (2017) High-resolution species-distribution model based on systematic sampling and indirect observations. Biodiversity and Conservation 26, 421-437.
| Crossref | Google Scholar |

Nickel BA, Suraci JP, Nisi AC, Wilmers CC (2021) Energetics and fear of humans constrain the spatial ecology of pumas. Proceedings of the National Academy of Sciences of the United States of America 118(5), e2004592118.
| Crossref | Google Scholar |

Nowell K, Jackson P (1996) ‘Wild cats – status survey and conservation action plan.’ (IUCN/SSC Cat Specialist Group, IUCN: Gland, Switzerland). p. 383.

Obunga G, Siljander M, Maghenda M, Pellikka PKE (2022) Habitat suitability modelling to improve conservation status of two critically endangered endemic Afromontane forest bird species in Taita Hills, Kenya. Journal for Nature Conservation 65, 126111.
| Crossref | Google Scholar |

Odden M, Wegge P (2005) Spacing and activity patterns of leopards Panthera pardus in the Royal Bardia National Park, Nepal. Wildlife Biology 11, 145-152.
| Crossref | Google Scholar |

Omidi F, Kaboli M, Karami M, Salman-Mahini A, Hassanzadeh-Kiabi B (2010) Habitat suitability modeling for Iranian leopard (Panthera pardus saxicolor) using ENFA in Kolah-Ghazi National Park, Isfahan Province. Journal of Environmental Science and Technology 12, 138-148 [In Persian with English abstract].
| Google Scholar |

Parchizadeh J, Adibi MA (2019) Distribution and human-caused mortality of Persian leopards Panthera pardus saxicolor in Iran, based on unpublished data and Farsi gray literature. Ecology and Evolution 9, 11972-11978.
| Crossref | Google Scholar | PubMed |

Parchizadeh J, Schooler SL, Adibi MA, Arias MG, Rezaei S, Belant JL (2023) A review of caracal and jungle cat diets across their geographical ranges during 1842–2021. Ecology and Evolution 13, e10130.
| Crossref | Google Scholar | PubMed |

Pellissier L, Rohr RP, Ndiribe C, Pradervand J-N, Salamin N, Guisan A, Wisz M (2013) Combining food web and species distribution models for improved community projections. Ecology and Evolution 3, 4572-4583.
| Crossref | Google Scholar | PubMed |

Phillips SJ, Dudík M, Schapire RE (2004) A maximum entropy approach to species distribution modeling. In ‘Proceedings of the twenty-first international conference on Machine learning’. pp. 655–662, Banff, Canada.

Phillips ND, Reid N, Thys T, Harrod C, Payne NL, Morgan CA, White HJ, Porter S, Houghton JDR (2017) Applying species distribution modelling to a data poor, pelagic fish complex: the ocean sunfishes. Journal of Biogeography 44, 2176-2187.
| Crossref | Google Scholar |

Pinti J, Shatley M, Carlisle A, Block BA, Oliver MJ (2022) Using pseudo-absence models to test for environmental selection in marine movement ecology: the importance of sample size and selection strength. Movement Ecology 10, 60.
| Crossref | Google Scholar |

Poursalem S, Amininasab SM, Zamani N, Almasieh K, Mardani M (2021) Modeling the distribution and habitat suitability of Persian Leopard Panthera pardus saxicolor in Southwestern Iran. Biology Bulletin 48, 319-330.
| Crossref | Google Scholar |

Rajula HSR, Verlato G, Manchia M, Antonucci N, Fanos V (2020) Comparison of conventional statistical methods with machine learning in medicine: diagnosis, drug development, and treatment. Medicina 56, 455.
| Crossref | Google Scholar | PubMed |

R Development Core Team (2020) ‘R: a language and environment for statistical computing.’ (R Foundation for Statistical Computing: Vienna)

Rew J, Cho Y, Moon J, Hwang E (2020) Habitat suitability estimation using a two-stage ensemble approach. Remote Sensing 12, 1475.
| Crossref | Google Scholar |

Riordan P, Cushman SA, Mallon D, Shi K, Hughes J (2016) Predicting global population connectivity and targeting conservation action for snow leopard across its range. Ecography 39, 419-426.
| Crossref | Google Scholar |

Ripley BD (2007) ‘Pattern recognition and neural networks.’ (Cambridge University Press)

Ripple WJ, Estes JA, Beschta RL, Wilmers CC, Ritchie EG, Hebblewhite M, Berger J, Elmhagen B, Letnic M, Nelson MP, Schmitz OJ, Smith DW, Wallach AD, Wirsing AJ (2014) Status and ecological effects of the world’s largest carnivores. Science 343(6167), 1241484.
| Crossref | Google Scholar | PubMed |

Rogan MS, Balme GA, Distiller G, Pitman RT, Broadfield J, Mann GKH, Whittington-Jones GM, Thomas LH, O’Riain MJ (2019) The influence of movement on the occupancy–density relationship at small spatial scales. Ecosphere 10, e02807.
| Crossref | Google Scholar |

Rosenblatt E, Creel S, Becker MS, Merkle J, Mwape H, Schuette P, Simpamba T (2016) Effects of a protection gradient on carnivore density and survival: an example with leopards in the Luangwa valley, Zambia. Ecology and Evolution 6, 3772-3785.
| Crossref | Google Scholar | PubMed |

Rozhnov VV, Chistopolova MD, Lukarevskii VS, Hernandez-Blanco JA, Naidenko SV, Sorokin PA (2015) Home range structure and space use of a female Amur leopard, Panthera pardus orientalis (Carnivora, Felidae). Biology Bulletin 42, 821-830.
| Crossref | Google Scholar |

Salas EAL, Valdez R, Michel S (2017) Summer and winter habitat suitability of Marco Polo argali in southeastern Tajikistan: a modeling approach. Heliyon 3(11), e00445.
| Crossref | Google Scholar |

Sampson AM (2013) A habitat suitability analysis for cougar (Puma concolor) in Minnesota. MSc Thesis. University of Minnesota.

Sanei A, Zakaria M, Hermidas S (2011) Prey composition in the Persian leopard distribution range in Iran. Asia Life Sciences Supplement 7, 19-30.
| Google Scholar |

Sanei A, Mousavi M, Kiabi B, Masoud MR, Gord Mardi E, Mohamadi H, Shakiba M, Zehi A, Teimouri M, Raeesi T (2016) Status assessment of the Persian leopard in Iran. Cat News Special Issue 10, 43-50.
| Google Scholar |

Selvan KM, Krishnakumar BM, Ramasamy P, Thinesh T (2019) Diel activity pattern of meso-carnivores in the suburban tropical dry evergreen forest of the Coromandel Coast, India. Journal of Threatened Taxa 11, 13960-13966.
| Crossref | Google Scholar |

Senay SD, Worner SP, Ikeda T (2013) Novel three-step pseudo-absence selection technique for improved species distribution modelling. PLoS ONE 8, e71218.
| Crossref | Google Scholar | PubMed |

Shafaeipour A, Siverio M, Siverio F (2016) Data on habitat and breeding biology of the Barbary Falcon, Falco peregrinus pelegrinoides Temminck, 1829, from South-western Iran. Acta Zoologica Bulgarica 68, 85-88.
| Google Scholar |

Sharbafi E, Farhadinia MS, Rezaie HR, Braczkowski AR (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, 1-8.
| Crossref | Google Scholar |

Shirley SM, Yang Z, Hutchinson RA, Alexander JD, McGarigal K, Betts MG (2013) Species distribution modelling for the people: unclassified landsat TM imagery predicts bird occurrence at fine resolutions. Diversity and Distributions 19(7), 855-866.
| Crossref | Google Scholar |

Soofi M, Ghoddousi A, Zeppenfeld T, Shokri S, Soufi M, Egli L, Jafari A, Ahmadpour M, Qashqaei A, Ghadirian T, Filla M, Kiabi B, Balkenhol N, Waltert M, Khorozyan I (2019) Assessing the relationship between illegal hunting of ungulates, wild prey occurrence and livestock depredation rate by large carnivores. Journal of Applied Ecology 56, 365-374.
| Crossref | Google Scholar |

Stein AB, Hayssen V (2013) Panthera pardus (Carnivora: Felidae). Mammalian Species 45, 30-48.
| Crossref | Google Scholar |

Stein AB, Athreya V, Gerngross P, Balme G, Henschel P, Karanth U, Miquelle D, Rostro-Garcia S, Kamler JF, Laguardia A, Khorozyan I, Ghoddousi A (2020) Panthera pardus. (amended version of 2019 assessment) The IUCN red list of threatened species e. T15954A163991139. doi:10.2305/IUCN.UK.2020-1.RLTS.T15954A163991139.en

Stuart C, Stuart T (2000) ‘A field guide to the tracks and signs of Southern and East African Wildlife.’ p. 310. (Penguin Random House South Africa)

Taghdisi M, Mohammadi A, Nourani E, Shokri S, Rezaei A, Kaboli M (2013) Diet and habitat use of the endangered Persian leopard (Panthera pardus saxicolor) in northeastern Iran. Turkish Journal of Zoology 37, 4.
| Crossref | Google Scholar |

Taleshi H, Jalali SG, Alavi SJ, Hosseini SM, Naimi B, Zimmermann NE (2019) Climate change impacts on the distribution and diversity of major tree species in the temperate forests of Northern Iran. Regional Environmental Change 19(8), 2711-2728.
| Crossref | Google Scholar |

Thomaes A, Kervyn T, Maes D (2008) Applying species distribution modelling for the conservation of the threatened saproxylic Stag Beetle (Lucanus cervus). Biological Conservation 141, 1400-1410.
| Crossref | Google Scholar |

Thuiller W, Lafourcade B, Engler R, Araújo MB (2009) BIOMOD – a platform for ensemble forecasting of species distributions. Ecography 32(3), 369-373.
| Crossref | Google Scholar |

Thuiller W, Georges D, Engler R, Breiner F, (2020) ‘biomod2’: Ensemble platform for species distribution modeling. (R package version 3.5.0). Available at https://CRAN.R-project.org/package=biomod2

Torres LG, Read AJ, Halpin P (2008) Fine-scale habitat modeling of a top marine predator: do prey data improve predictive capacity. Ecological Applications 18, 1702-1717.
| Crossref | Google Scholar | PubMed |

Trainor AM, Schmitz OJ, Ivan JS, Shenk TM (2014) Enhancing species distribution modeling by characterizing predator-prey interactions. Ecological Applications 24, 204-216.
| Crossref | Google Scholar | PubMed |

van Proosdij ASJ, Sosef MSM, Wieringa JJ, Raes N (2016) Minimum required number of specimen records to develop accurate species distribution models. Ecography 39, 542-552.
| Crossref | Google Scholar |

Walker C (1996) ‘Signs of the wild.’ p. 215. (Struik Publish: Cape Town)

Wallach AD, Izhaki I, Toms JD, Ripple WJ, Shanas U (2015) What is an apex predator? Oikos 124, 1453-1461.
| Crossref | Google Scholar |

Wilmers CC, Isbell LA, Suraci JP, Williams TM (2017) Energetics-informed behavioral states reveal the drive to kill in African leopards. Ecosphere 8, e01850.
| Crossref | Google Scholar |

Wisz MS, Hijmans RJ, Li J, Peterson AT, Graham CH, Guisan A, NCEAS predicting species distributions working group (2008) Effects of sample size on the performance of species distribution models. Diversity and Distributions 14, 763-773.
| Crossref | Google Scholar |

Yusefi GH, Faizolahi K, Darvish J, Safi K, Brito JC (2019) The species diversity, distribution, and conservation status of the terrestrial mammals of Iran. Journal of Mammalogy 100, 55-71.
| Crossref | Google Scholar |

Zanaga D, Van De Kerchove R, Daems D, De Keersmaecker W, Brockmann C, Kirches G, Wevers J, Cartus O, Santoro M, Fritz S, Lesiv M, Herold M, Tsendbazar NE, Xu P, Ramoino F, Arino O (2022) ESA WorldCover 10 m 2021 v200. Available at https://doi.org/10.5281/zenodo.7254221

Zanin M, Palomares F, Brito D (2015) What we (don’t) know about the effects of habitat loss and fragmentation on felids. Oryx 49, 96-106.
| Crossref | Google Scholar |

Zhang L, Liu S, Sun P, Wang T, Wang G, Zhang X, Wang L (2015) Consensus forecasting of species distributions: the effects of niche model performance and niche properties. PLoS ONE 10(3), e0120056.
| Crossref | Google Scholar | PubMed |

Zimmermann F, Lukarevski VS, Beruchashvili G, Breitenmoser-Wursten C, Breitenmoser U (2007) Mapping the vision – potential living space for the leopard in the Caucasus. Cat News Special Issue 2, 28-33.
| Google Scholar |

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 |