Conservation deficits and climate-driven habitat shifts of threatened plants in South Asia

Maroof Ali; Zhongde Huang; Zishan Ahmad Wani; Jianhua Song; Bingxiao Wu; Khateeb Hussain; Abdul Rahim PP; Jie Wang; Zhou Fang; Rainer W. Bussmann; David Y.P. Tng; Jia-Qi Zhang; Yang Bai

doi:10.1016/j.geosus.2026.100477

Geography and Sustainability ›› 2026, Vol. 7 ›› Issue (4) :100477 DOI: 10.1016/j.geosus.2026.100477

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research-article

Conservation deficits and climate-driven habitat shifts of threatened plants in South Asia

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^a Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences , Menglun, Mengla, Yunnan 666303, China

^b University of Chinese Academy of Sciences , Beijing 100049, China

^c Terrestrial Ecology and Modelling (TEaM) Lab, Department of Environmental Science and Engineering , SRM University-AP, Amaravati 522 240, Andhra Pradesh, India

^d Institute of Facility Agriculture, Guangdong Academy of Agriculture Sciences , Guangzhou 510640, China

^e School of Ecology and Environment, Anhui Normal University , Wuhu 241000, China

^f Department of Ethnobotany, Institute of Botany, Ilia State University , Tbilisi, Georgia

^g Centre for Rainforest Studies, School for Field Studies , Yungaburra Qld 4872, Australia

^h Ministry of Education Key Laboratory for Ecology of Tropical Islands, Key Laboratory of Tropical Animal and Plant Ecology of Hainan Province, College of Life Sciences, Hainan Normal University , Haikou 571158, China

ⁱ Biodiversity Museum of Hainan Normal University , Haikou 571158, China

^j Yunnan Key Laboratory for Conservation of Tropical Rainforests and Asian Elephants Chinese Academy of Sciences , Mengla 666303, China

^k Yunnan International Joint Laboratory of Southeast Asia Biodiversity Conservation , Menglun 666303, China

^* Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun, Mengla, Yunnan 666303, PR China. E-mail addresses: maroof@xtbg.ac.cn (M. Ali), huangzhongde@xtbg.ac.cn (Z. Huang), zishanwani786@gmail.com (Z.A. Wani), songjianhua@xtbg.ac.cn (J.Song), wbx19911026@outlook.com (B. Wu), khateeb@ahnu.edu.cn (K. Hussain), abdulrahim_p@srmap.edu.in (A.R. PP), wangjie2021@xtbg.ac.cn (J. Wang), fangzhou91@xtbg.ac.cn (Z. Fang), Rainer.Bussmann@iliauni.edu.ge (R.W. Bussmann), dtng@fieldstudies.org (D.Y.P. Tng), zhangjiaqi_solana@163.com (J.-Q. Zhang), baiyang@xtbg.ac.cn (Y. Bai) .

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Abstract

South Asia’s rich biodiversity, shaped by its complex biogeographic history, faces increasing threats from climate change, land-use change, and habitat fragmentation. Impacts of these drivers of biodiversity loss on species distributions, habitat dynamics, and protected area effectiveness remain poorly understood. We integrated MaxEnt-based species distribution modelling, habitat fragmentation analysis, and spatial conservation prioritization to assess current and future conservation priorities for 127 threatened plant species (33 Near Threatened, 41 Vulnerable, 38 Endangered, and 15 Critically Endangered) across South Asia. We compiled 8,503 georeferenced occurrence records and 24 environmental predictors to model current distributions and projected suitability for the years 2050 and 2070 under SSP126, SSP245, and SSP585 scenarios using the GISS-E2-1-G climate model. Habitat fragmentation and connectivity were quantified using FRAGSTATS, and zonation was applied to identify high-priority conservation areas. Overlay analysis was performed to quantify the spatial overlap between high-priority conservation areas identified through zonation and the existing protected area network. MaxEnt revealed that under current climatic scenarios, only 6.05 % of the total area of interest was predicted to be highly suitable for these species. The most suitable habitats were identified in the Himalayan mountain range, including its western, central, and eastern parts, and the Western Ghats of southern India, with moderate patch density (0.0037 patches/km²), and high connectivity (proximity index 69,473; aggregation index 89.92). Species’ habitat suitability was predicted to undergo significant changes under future climatic scenarios. Under most of the climatic scenarios across time periods, highly suitable habitats were found to decline considerably with only 9.39 % to 14.33 % of highly suitable habitat overlapping with existing protected area networks. Species-wise analysis revealed that a considerable percentage of species ranging from 44.40 % under SSP126 (2050) to 83.3 % under SSP585 (2070) are predicted to lose suitable habitats. Further, future climate projections indicate increased fragmentation from 29.66 % to 36.39 % and declining connectivity across all scenarios. Spatial prioritization identified 912,037 km² as high-priority zone; 85.01 % of which lies outside the existing protected area networks. These results demonstrate the need to expand and connect protected areas and implement adaptive, climate-informed conservation strategies to safeguard South Asia’s threatened flora under rapid environmental change.

Keywords

Biodiversity hotspots / Threatened flora / Climate drivers / Biodiversity conservation / Conservation priorities

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Maroof Ali, Zhongde Huang, Zishan Ahmad Wani, Jianhua Song, Bingxiao Wu, Khateeb Hussain, Abdul Rahim PP, Jie Wang, Zhou Fang, Rainer W. Bussmann, David Y.P. Tng, Jia-Qi Zhang, Yang Bai. Conservation deficits and climate-driven habitat shifts of threatened plants in South Asia. Geography and Sustainability, 2026, 7 (4) : 100477 DOI:10.1016/j.geosus.2026.100477

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References

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[1]	Advani, N.K., 2023. Assessing species vulnerability to climate change, and implementing practical solutions. Biol. Conserv. 286, 110284. doi: 10.1016/j.biocon.2023.110284.

[2]	Ahmad, A., Gopi, G.V., 2025. Understanding multi-scale and multi-species habitat selection by mammals in the eastern Himalayan biodiversity hotspot. Ecol. Evol. 15 (4), e71247. doi: 10.1002/ece3.71247.

[3]	Ali, M., Huang, Z.D., Bai, Y., Tng, D.Y.P., Qin, F., Fang, Z., 2024. A multifaceted approach to expanding conservation efforts in the Pan-Himalayan landscape. J. Clean. Prod. 476, 143783. doi: 10.1016/j.jclepro.2024.143783.

[4]	Amindin, A., Pourghasemi, H.R., Safaeian, R., Rahmanian, S., Tiefenbacher, J.P., Naimi, B., 2024. Predicting current and future habitat suitability of an endemic species using data-fusion approach: responses to climate change. Rangel. Ecol. Manag. 94, 149-162. doi: 10.1016/j.rama.2024.03.002.

[5]	Banks-Leite, C., Ewers, R.M., Folkard-Tapp, H., Fraser, A., 2020. Countering the effects of habitat loss, fragmentation, and degradation through habitat restoration. One Earth 3 (6), 672-676. doi: 10.1016/j.oneear.2020.11.016.

[6]	Baral, P., Allen, S., Steiner, J.F., Gurung, T.R., McDowell, G., 2023. Climate change impacts and adaptation to permafrost change in High Mountain Asia: a comprehensive review. Environ. Res. Lett. 18 (9), 093005. doi: 10.1088/1748-9326/acf1b4.

[7]	Bergstrom, D.M., Wienecke, B.C., Van den Hoff, J., Hughes, L., Lindenmayer, D.B., Ainsworth, T.D., Baker, C.M., Bland, L., Bowman, D.M., Brooks, S.T., 2021. Combating ecosystem collapse from the tropics to the antarctic. Glob. Change Biol. 27 (9), 1692-1703. doi: 10.1111/gcb.15539.

[8]	Cabernard, L., Pfister, S., Hellweg, S., 2024. Biodiversity impacts of recent land-use change driven by increases in agri-food imports. Nat. Sustain. 7 (11), 1512-1524. doi: 10.1038/s41893-024-01433-4.

[9]	Chaudhary, R.P., Uprety, Y., Acharya, H.R., Rimal, S.K., 2023. Deforestation in Nepal: status, causes, consequences, and responses. In: Sivanpillai, R. (Ed.), Biological and Environmental Hazards, Risks, and Disasters. Elsevier, pp. 277-318. doi: 10.1016/B978-0-12-820509-9.00003-4.

[10]	Chen, I.C., Hill, J.K., Ohlemüller, R., Roy, D.B., Thomas, C.D., 2011. Rapid range shifts of species associated with high levels of climate warming. Science 333 (6045), 1024-1026. doi: 10.1126/science.1206432.

[11]	Chen, I.C., Shen, S.-F., Chan, S.-F., 2024. Niche theory and species range limits along elevational gradients: perspectives and future directions. Annu. Rev. Ecol. Evol. Syst. 55 (1), 449-469. doi: 10.1146/annurev-ecolsys-102722-123834.

[12]	Chettri, N., Adhikari, B., Chaudhary, S., Wangchuk, K., 2023. Changing discourses in the third pole: a systematic review of climate change impact on biodiversity in the Hindu Kush Himalaya. Ecol. Indic. 155, 111046. doi: 10.1016/j.ecolind.2023.111046.

[13]	Choden, K., Nitschke, C.R., Stewart, S.B., Keenan, R.J., 2021. The potential impacts of climate change on the distribution of key tree species and cordyceps in Bhutan: implications for ecological functions and rural livelihoods. Ecol. Model. 455, 109650. doi: 10.1016/j.ecolmodel.2021.109650.

[14]	Clark, N.E., Boakes, E.H., McGowan, P.J., Mace, G.M., Fuller, R.A., 2013. Protected areas in South Asia have not prevented habitat loss: a study using historical models of land-use change. PLoS One 8 (5), e65298. doi: 10.1371/journal.pone.0065298.

[15]	Convention on Biological Diversity, 2022. Kunming-Montreal Global Biodiversity Framework. Decision Adopted by the Conference of the Parties to the Convention on Biological Diversity. CBD/COP/DEC/15/4. Montreal, Canada, 7-19 December 2022.

[16]

De Frenne, P., Lenoir, J., Luoto, M., Scheffers, B.R., Zellweger, F., Aalto, J., Ashcroft, M.B., Christiansen, D.M., Decocq, G., De Pauw, K., Govaert, S., Greiser, C., Gril, E., Hampe, A., Jucker, T., Klinges, D.H., Koelemeijer, I.A., Lembrechts, J.J., Marrec, R., Meeussen, C., Ogée, J., Tyystjärvi, V., Vangansbeke, P., Hylander, K., 2021. Forest microclimates and climate change: importance, drivers and future research agenda. Glob. Change Biol. 27 (11), 2279-42297. doi: 10.1111/gcb.15569.

[17]	Des Roches, S., Post, D.M., Turley, N.E., Bailey, J.K., Hendry, A.P., Kinnison, M.T., Schweitzer, J.A., Palkovacs, E.P., 2018. The ecological importance of intraspecific variation. Nat. Ecol. Evol. 2 (1), 57-64. doi: 10.1038/s41559-017-0402-5.

[18]	Dhyani, S., Sivadas, D., Basu, O., Karki, M., 2022. Ecosystem health and risk assessments for high conservation value mountain ecosystems of South Asia: a necessity to guide conservation policies. Anth. Sci. 1 (1), 211-225. doi: 10.1007/s44177-022-00010-8.

[19]	Dunn, J.C., Buchanan, G.M., Stein, R.W., Whittingham, M.J., McGowan, P.J.K., 2016. Optimising different types of biodiversity coverage of protected areas with a case study using Himalayan Galliformes. Biol. Conserv. 196, 22-30. doi: 10.1016/j.biocon.2016.01.015.

[20]

Elith, J., Burgman, M.A., Wall, W.A., Haufler, J ., 2002. Predictions and their validation: rare plants in the central highlands, Victoria, Australia. In: Scott, J.M., Heglund, P.J., Morrison, M.L. (Eds.), Predicting Species Occurrences: Issues of Accuracy and Scale. Island Press, Covelo, pp. 303-314.

[21]	Elith, J., Graham, C.H., Anderson, R.P., Dudík, M., Ferrier, S., Guisan, A., Williams, S., 2006. Novel methods improve prediction of species’ distributions from occurrence data. Ecography 29 (2), 129-151. doi: 10.1111/j.2006.0906-7590.04596.x.

[22]	Fick, S.E., Hijmans, R.J., 2017. WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37 (12), 4302-4315. doi: 10.1002/joc.5086.

[23]	Fitzpatrick, M.C., Gotelli, N.J., Ellison, A.M., 2013. MaxEnt versus MaxLike: empirical comparisons with ant species distributions. Ecosphere 4 (5), art55. doi: 10.1890/ES13-00066.1.

[24]

Friedman, K., Bridgewater, P., Agostini, V., Agardy, T., Arico, S., Biermann, F., Brown, K., Cresswell, I.D., Ellis, E.C., Failler, P., Kim, R.E., Pratt, C., Rice, J., Rivera, V.S., Teneva, L., 2022. The CBD post-2020 biodiversity framework: people’s place within the rest of nature. People Nat. 4 (6), 1475-1484. doi: 10.1002/pan3.10403.

[25]

González-Moreno, P., Lazzaro, L., Vilà, M., Preda, C., Adriaens, T., Bacher, S., Brundu, G., Copp, G.H., Essl, F., García-Berthou, E., Katsanevakis, S., Moen, T.L., Lucy, F.E., Nentwig, W., Roy, H.E., Srebalienė, G., Talgø, V., Vanderhoeven, S., Andjelković, A., Arbačiauskas, K., Auger-Rozenberg, M.A., Bae, M.J., Bariche, M., Boets, P., Boieiro, M., Borges, P.A., Canning-Clode, J., Cardigos, F., Chartosia, N., Cottier-Cook, E.J., Crocetta, F., D’Hondt, B., Foggi, B., Follak, S., Gallardo, B., Gammelmo, Ø., Giakoumi, S., Giuliani, C., Guillaume, F., Jelaska, L.Š., Jeschke, J.M., Jover, M., Juárez-Escario, A., Kalogirou, S., Kočić, A., Kytinou, E., Laverty, C., Lozano, V., Maceda-Veiga, A., Marchante, E., Marchante, H., Martinou, A.F., Meyer, S., Minchin, D., Montero-Castaño, A., Morais, M.C., Morales-Rodriguez, C., Muhthassim, N., Nagy, Z.Á., Ogris, N., Onen, H., Pergl, J., Puntila, R., Rabitsch, W., Ramburn, T.T., Rego, C., Reichenbach, F., Romeralo, C., Saul, W.C., Schrader, G., Sheehan, R., Simonović, P., Skolka, M., Soares, A.O., Sundheim, L., Tarkan, A.S., Tomov, R., Tricarico, E., Tsiamis, K., Uludağ, A., van Valkenburg, J., Verreycken, H., Vettraino, A.M., Vilar, L., Wiig, Ø., Witzell, J., Zanetta, A., Kenis, M., 2019. Consistency of impact assessment protocols for non-native species. NeoBiota 44, 1-25. doi: 10.3897/neobiota.44.31650.

[26]	Habibullah, M.S., Din, B.H., Tan, S.-H., Zahid, H., 2022. Impact of climate change on biodiversity loss: global evidence. Environ. Sci. Pollut. Res. 29 (1), 1073-1086. doi: 10.1007/s11356-021-15702-8.

[27]

Hamid, M., Gulzar, A., Dar, F.A., Singh, C.P., Malik, A.H., Kamili, A.N., AhmadKhuroo, A., 2023. Microclimate heterogeneity modulates fine-scale edaphic and vegetation patterns on the Himalayan treelines: implications under climate change. Agric. For. Meteorol. 341, 109688. doi: 10.1016/j.agrformet.2023.109688.

[28]	Huang, Z.D., Bai, Y., Alatalo, J.M., Yang, Z.Q., 2020. Mapping biodiversity conservation priorities for protected area: A case study in Xisihuangbanna Tropical Area, China. Biol. Conserv. 249, 108741. doi: 10.1016/j.biocon.2020.108741.

[29]	Hughes, A.C., 2017. Mapping priorities for conservation in Southeast Asia. Biol. Conserv. 209, 395-405. doi: 10.1016/j.biocon.2017.03.007.

[30]	Ioan, S., Roseo, F., Brambilla, M., 2025. Mountain ecosystem services under a changing climate: a global perspective. Ecosyst. Serv. 73, 101732. doi: 10.1016/j.ecoser.2025.101732.

[31]	Körner, C., 2007. The use of ‘altitude’ in ecological research. Trends Ecol. Evol. 22 (11), 569-574. doi: 10.1016/j.tree.2007.09.006.

[32]	Kattel, G.R., 2022. Climate warming in the Himalayas threatens biodiversity, ecosystem functioning and ecosystem services in the 21st century: is there a better solution? Biodivers. Conserv. 31 (8), 2017-2044. doi: 10.1007/s10531-022-02417-6.

[33]	Kearney, M., Porter, W., 2009. Mechanistic niche modelling: combining physiological and spatial data to predict species’ ranges. Ecol. Lett. 12 (4), 334-350. doi: 10.1111/j.1461-0248.2008.01277.x.

[34]	Kong, X.S., Zhou, Z.Z., Jiao, L.M., 2021. Hotspots of land-use change in global biodiversity hotspots. Resour. Conserv. Recycl. 174, 105770. doi: 10.1016/j.resconrec.2021.105770.

[35]	Lalande, M., Ménégoz, M., Krinner, G., 2021. Climate change in the high Mountain Asia simulated with CMIP6 models. In: Copernicus Meetings, pp. 1061-1098. doi: 10.5194/EGUSPHERE-EGU21-8365.

[36]	Lenoir, J., Gégout, J.C., Pierrat, J.C., Bontemps, J.D., Dhôte, J.F ., 2009. Differences between tree species seedling and adult altitudinal distribution in mountain forests during the recent warm period (1986-2006). Ecography 32 (5), 765-777. doi: 10.1111/j.1600-0587.2009.05791.

[37]	Li, C.Z., Huang, Z.D., Fang, Z., Wang, Y.T., Ali, M., Bai, Y., 2025. Deciphering biodiversity drivers in mainland Southeast Asia: sensitivity and threshold response hotspots. Environ. Sustain. Indic. 27, 100758. doi: 10.1016/j.indic.2025.100758.

[38]	Liu, F.-L., Mambo, W.W., Liu, J., Zhu, G.-F., Khan, R., Abdullah Khan, S.M., Lu, L., 2025. Spatiotemporal range dynamics and conservation optimization for endangered medicinal plants in the Himalaya. Glob. Ecol. Conserv. 57, e03390. doi: 10.1016/j.gecco.2024.e03390.

[39]	Ludovicy, S., Noroozi, J., Semenchuk, P., Moser, D., Wessely, J., Talebi, A., Dullinger, S., 2022. Protected area network insufficiently represents climatic niches of endemic plants in a Global Biodiversity Hotspot. Biol. Conserv. 275, 109768. doi: 10.1016/j.biocon.2022.109768.

[40]

Ma, B.R., Zeng, W.H., Xie, Y.X., Wang, Z.Z., Hu, G.Z., Li, Q., Cao, R.X., Zhuo, Y., Zhang, T.Z., 2022. Boundary delineation and grading functional zoning of Sanjiangyuan National Park based on biodiversity importance evaluations. Sci. Total Environ. 825, 154068. doi: 10.1016/j.scitotenv.2022.154068.

[41]	Maina, F.Z., Xue, Y., Kumar, S.V., Getirana, A., McLarty, S., Appana, R., Forman, B., Zaitchik, B., Loomis, B., Maggioni, V., Zhou, Y.F., 2024. Development of a multidecadal land reanalysis over high mountain asia. Sci. Data 11 (1), 827. doi: 10.1038/s41597-024-03643-z.

[42]	Majeed, K.A., Khwarahm, N.R., Ahmed, S.H., 2023. Predicting the geographical distribution of the Persian leopard, Panthera pardus tulliana, a rare and endangered species . J. Nat. Conserv. 76, 126505. doi: 10.1016/j.jnc.2023.126505.

[43]	Matawa, F., Murwira, A., Zengeya, F.M., Atkinson, P.M., 2016. Modelling the spatial-temporal distribution of tsetse (Glossina pallidipes) as a function of topography and vegetation greenness in the Zambezi Valley of Zimbabwe . Appl. Geogr. 76, 198-206. doi: 10.1016/j.apgeog.2016.09.008.

[44]	McCain, C.M., Colwell, R.K., 2011. Assessing the threat to montane biodiversity from discordant shifts in temperature and precipitation in a changing climate. Ecol. Lett. 14 (12), 1236-1245. doi: 10.1111/j.1461-0248.2011.01695.x.

[45]	McGrigal, K., Cushman, S.A., Ene, E.E. . FRAGSTATS v4: spatial patttern analyisis program for cateogical maps. Computer softawre. http://www.fragstats.org.

[46]

Miller, R.L., Schmidt, G.A., Nazarenko, L.S., Bauer, S.E., Kelley, M., Ruedy, R., Russell, G.L., Ackerman, A.S., Aleinov, I., Bauer, M., Bleck, R., Canuto, V., Cesana, G., Cheng, Y., Clune, T.L., Cook, B.I., Cruz, C.A., Del Genio, A.D., Elsaesser, G.S., Faluvegi, G., Kiang, N.Y., Kim, D., Lacis, A.A., Leboissetier, A., LeGrande, A.N., Lo, K.K., Marshall, J., Matthews, E.E., McDermid, S., Mezuman, K., Murray, L.T., Oinas, V., Orbe, C., Pérez García-Pando, C., Perlwitz, J.P., Puma, M.J., Rind, D., Romanou, A., Shindell, D.T., Sun, S., Tausnev, N., Tsigaridis, K., Tselioudis, G., Weng, E.S., Wu, J.B., Yao, M.S., 2021. CMIP6 historical simulations (1850-2014) with GISS-E2. 1. J. Adv. Model. Earth Syst. 13 (1), e2019MS002034. doi: 10.1029/2019MS002034.

[47]	Mirhashemi, H., Ahmadi, K., Heydari, M., Karami, O., Valkó, O., Khwarahm, N.R., 2024. Climatic variables are more effective on the spatial distribution of oak forests than land use change across their historical range. Environ. Monit. Assess. 196 (3), 289. doi: 10.1007/s10661-024-12438-z.

[48]	Moilanen, A., Lehtinen, P., Kohonen, I., Jalkanen, J., Virtanen, E.A., Kujala, H., 2022. Novel methods for spatial prioritization with applications in conservation, land use planning and ecological impact avoidance. Methods Ecol. Evol. 13 (5), 1062-1072. doi: 10.1111/2041-210X.13819.

[49]

Morelli, T.L., Daly, C., Dobrowski, S.Z., Dulen, D.M., Ebersole, J.L., Jackson, S.T., Lundquist, J.D., Millar, C.I., Maher, S.P., Monahan, W.B., Nydick, K.R., Redmond, K.T., Sawyer, S.C., Stock, S., Beissinger, S.R., 2016. Managing climate change refugia for climate adaptation. PLoS One 11 (8), e0159909. doi: 10.1371/journal.pone.0159909.

[50]	Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca, G.A.B., Kent, J., 2000. Biodiversity hotspots for conservation priorities. Nature 403 (6772), 853-858. doi: 10.1038/35002501.

[51]

Newbold, T., Hudson, L.N., Hill, S.L., Contu, S., Lysenko, I., Senior, R.A., Börger, L., Bennett, D.J., Choimes, A., Collen, B., Day, J., De Palma, A., Díaz, S., Echeverria-Londoño, S., Edgar, M.J., Feldman, A., Garon, M., Harrison, M.L., Alhusseini, T., Ingram, D.J., Itescu, Y., Kattge, J., Kemp, V., Kirkpatrick, L., Kleyer, M., Correia, D.L., Martin, C.D., Meiri, S., Novosolov, M., Pan, Y., Phillips, H.R., Purves, D.W., Robinson, A., Simpson, J., Tuck, S.L., Weiher, E., White, H.J., Ewers, R.M., Mace, G.M., Scharlemann, J.P., Purvis, A., 2015. Global effects of land use on local terrestrial biodiversity. Nature 520 (7545), 45-50. doi: 10.1038/nature14324.

[52]

O’Neill, B.C., Tebaldi, C., van Vuuren, D.P., Eyring, V., Friedlingstein, P., Hurtt, G., Knutti, R., Kriegler, E., Lamarque, J.-F., Lowe, J., Meehl, G.A., Moss, R., Riahi, K., Sanderson, B.M., 2016. The scenario model intercomparison project (ScenarioMIP) for CMIP6. Geosci. Model Dev. 9 (9), 3461-3482. doi: 10.5194/gmd-9-3461-2016.

[53]	Pang, S.E., Slik, J.W.F., Chisholm, R.A., Webb, E.L., 2024. Conserving Southeast Asian trees requires mitigating both climate and land-use change. Nat. Sustain. 7 (10), 1313-1323. doi: 10.1038/s41893-024-01417-4.

[54]	Parks, S.A., Holsinger, L.M., Abatzoglou, J.T., Littlefield, C.E., Zeller, K.A., 2023. Protected areas not likely to serve as steppingstones for species undergoing climate-induced range shifts. Glob. Change Biol. 29 (10), 2681-2696. doi: 10.1111/gcb.16629.

[55]	Parmesan, C., 2006. Ecological and evolutionary responses to recent climate change. Annu. Rev. Ecol. Evol. Syst. 37 (1), 637-669. doi: 10.1146/annurev.ecolsys.37.091305.110100.

[56]	Penjor, U., Wangdi, S., Tandin, T., MacDonald, D.W., 2021. Vulnerability of mammal communities to the combined impacts of anthropic land-use and climate change in the Himalayan conservation landscape of Bhutan. Ecol. Indic. 121, 107085. doi: 10.1016/j.ecolind.2020.107085.

[57]	Phillips, S.J., Anderson, R.P., Schapire, R.E., 2006. Maximum entropy modeling of species geographic distributions. Ecol. Model. 190 (3-4), 231-259. doi: 10.1016/j.ecolmodel.2005.03.026.

[58]	Phillips, S.J., Anderson, R.P., Dudík, M., Schapire, R.E., Blair, M.E., 2017. Opening the black box: an open-source release of MaxEnt. Ecography 40 (7), 887-893. doi: 10.1111/ecog.03049.

[59]	Powers, R.P., Jetz, W., 2019. Global habitat loss and extinction risk of terrestrial vertebrates under future land-use-change scenarios. Nat. Clim. Chang. 9 (4), 323-329. doi: 10.1038/s41558-019-0406-z.

[60]	Radha, K.O., Khwarahm, N.R., 2022. An integrated approach to map the impact of climate change on the distributions of crataegus azarolus and crataegus monogyna in Kurdistan Region, Iraq. Sustainability 14 (21), 14621. doi: 10.3390/su142114621.

[61]

Ramesha, B.T., Ravikanth, G., Nageswara Rao, M., Ganeshaiah, K.N., Uma Shaanker, R., 2007. Genetic structure of the rattan Calamus thwaitesii in core, buffer and peripheral regions of three protected areas in central Western Ghats, India: do protected areas serve as refugia for genetic resources of economically important plants? J. Genet. 86 (1), 9-18. doi: 10.1007/s12041-007-0002-2.

[62]	Rather, Z.A., Ahmad, R., Dar, T.U.H., Khuroo, A.A., 2022. Ensemble modelling enables identification of suitable sites for habitat restoration of threatened biodiversity under climate change: a case study of Himalayan Trillium. Ecol. Eng. 176, 106534. doi: 10.1016/j.ecoleng.2021.106534.

[63]

Riahi, K., van Vuuren, D.P., Kriegler, E., Edmonds, J., O’Neill, B.C., Fujimori, S., Bauer, N., Calvin, K., Dellink, R., Fricko, O., Lutz, W., Popp, A., Cuaresma, J.C., Kc, S., Leimbach, M., Jiang, L.W., Kram, T., Rao, S., Emmerling, J., Ebi, K., Hasegawa, T., Havlik, P., Humpenöder, F., Da Silva, L.A., Smith, S., Stehfest, E., Bosetti, V., Eom, J., Gernaat, D., Masui, T., Rogelj, J., Streffler, J., Drouet, L., Krey, V., Luderer, G., Harmsen, M., Takahashi, K., Baumstark, L., Doelman, J.C., Kainuma, M., Klimont, Z., Marangoni, G., Lotze-Campen, H., Obersteiner, M., Tabeau, A., Tavoni, M., 2017. The Shared Socioeconomic Pathways and their energy, land use, and greenhouse gas emissions implications: an overview. Glob. Environ. Change 42, 153-168. doi: 10.1016/j.gloenvcha.2016.05.009.

[64]	Ribeiro, R.C., Figueiredo, M.L.N., Picorelli, A., Silveira, F.A.O., 2023. Limited seed dispersal distance in endemic species from tropical mountaintop grasslands may restrict upward migration in response to climate change. Flora 298, 152203. doi: 10.1016/j.flora.2022.152203.

[65]	Roy, S., Suman, A., Ray, S., Saikia, S.K., 2022. Use of species distribution models to study habitat suitability for sustainable management and conservation in the Indian subcontinent: a decade’s retrospective. Front. Sustain. Resour. Manag. 1, 1031646. doi: 10.3389/fsrma.2022.1031646.

[66]	Sandel, B., Arge, L., Dalsgaard, B., Davies, R.G., Gaston, K.J., Sutherland, W.J., Svenning, J.-C., 2011. The influence of late quaternary climate-change velocity on species endemism. Science 334 (6056), 660-664. doi: 10.1126/science.1210173.

[67]	Sandhu, H., Sandhu, S., 2015. Poverty, development, and Himalayan ecosystems. Ambio 44 (4), 297-307. doi: 10.1007/s13280-014-0569-9.

[68]

Saqib, S., Ullah, F., Omollo, W.O., Liu, Y., Tao, H.-Y., Zaman, W., Temur, A., Liu, B., Lai, Y.J., Chen, Z.D., Fang, Y.P., Dong, H.J., Wang, W.Y., Zhao, X.M., Xiong, Y.C., 2025. Identifying hotspots and climate drivers of alien plant species for conservation prioritization across the Pan-Himalaya. Biol. Conserv. 302, 110994. doi: 10.1016/j.biocon.2025.110994.

[69]	Saunders, S.P., Grand, J., Bateman, B.L., Meek, M., Wilsey, C.B., Forstenhaeusler, N., Graham, E., Warren, R., Price, J., 2023. Integrating climate-change refugia into 30 by 30 conservation planning in North America. Front. Ecol. Environ. 21 (2), 77-84. doi: 10.1002/fee.2592.

[70]	Schillaci, M., Raio, A., Sillo, F., Zampieri, E., Mahmood, S., Anjum, M., Khalid, A., Centritto, M., 2022. Pseudomonas and Curtobacterium strains from olive rhizosphere characterized and evaluated for plant growth promoting traits . Plants 11 (17), 2245. doi: 10.3390/plants11172245.

[71]	Selwood, K.E., Zimmer, H.C., 2020. Refuges for biodiversity conservation: a review of the evidence. Biol. Conserv. 245, 108502. doi: 10.1016/j.biocon.2020.108502.

[72]	Sengupta, A., Ravikanth, G., Seshadri, K.S., Bunyan, M., Ganesh, T., Rajan, P.D., Devy, M.S., Aravind, N.A., 2025. The shifting paradigms of biodiversity conservation in South Asia. Biotropica 57 (2), e70013. doi: 10.1111/btp.70013.

[73]	Shtiliyanova, A., Bellocchi, G., Borras, D., Eza, U., Martin, R., Carrère, P., 2017. Kriging-based approach to predict missing air temperature data. Comput. Electron. Agric. 142 (PartA), 440-449. doi: 10.1016/j.compag.2017.09.033.

[74]	Sonne, J., Rahbek, C., 2024. Idiosyncratic patterns of local species richness and turnover define global biodiversity hotspots. Proc. Natl. Acad. Sci. U.S.A. 121 (3), e2313106121. doi: 10.1073/pnas.2313106121.

[75]

Struebig, M.J., Wilting, A., Gaveau, D.L.A., Meijaard, E., Smith, R.J., Fischer, M., Metcalfe, K., Kramer-Schadt, S., Abdullah, T., Abram, N., Alfred, R., Ancrenaz, M., Augeri, D.M., Belant, J.L., Bernard, H., Bezuijen, M., Boonman, A., Boonratana, R., Boorsma, T., Breitenmoser-Würsten, C., Brodie, J., Cheyne, S.M., Devens, C., Duckworth, J.W., Duplaix, N., Eaton, J., Francis, C., Fredriksson, G., Giordano, A.J., Gonner, C., Hall, J., Harrison, M.E., Hearn, A.J., Heckmann, I., Heydon, M., Hofer, H., Hon, J., Husson, S., Ali Anwarali Khan, F., Kingston, T., Kreb, D., Lammertink, M., Lane, D., Lasmana, F., Liat, L.B., Lim, N.T., Lindenborn, J., Loken, B., MacDon ald, D.W., Marshall, A.J., Maryanto, I., Mathai, J., McShea, W.J., Mohamed, A., Nakabayashi, M., Nakashima, Y., Niedballa, J., Noerfahmy, S., Persey, S., Peter, A., Pieterse, S., Pilgrim, J.D., Pollard, E., Purnama, S., Rafiastanto, A., Reinfelder, V., Reusch, C., Robson, C., Ross, J., Rustam, R., Sadikin, L., Samejima, H., Santosa, E., Sapari, I., Sasaki, H., Scharf, A.K., Semiadi, G., Shepherd, C.R., Sykes, R., van Berkel, T., Wells, K., Wielstra, B., Wong, A., 2015. Targeted conservation to safeguard a biodiversity hotspot from climate and land-cover change. Curr. Biol. 25 (3), 372-378. doi: 10.1016/j.cub.2014.11.067.

[76]	Sun, W.C., Zhang, E.T., Zhao, Y.J., Wu, Z.S., Chen, W.H., Wang, Y., Bai, Y.F., 2025. Conservation priority corridors enhance the effectiveness of protected area networks in China. Commun. Earth Environ. 6 (1), 275. doi: 10.1038/s43247-025-02227-y.

[77]	Tan, Y.-L., Chen, J.-E., Yiew, T.-H., Habibullah, M.S., 2022. Habitat change and biodiversity loss in South and Southeast Asian countries. Environ. Sci. Pollut. Res. 29 (42), 63260-63276. doi: 10.1007/s11356-022-20054-y.

[78]

Thackeray, S.J., Henrys, P.A., Hemming, D., Bell, J.R., Botham, M.S., Burthe, S., Helaouet, P., Johns, D.G., Jones, I.D., Leech, D.I., MacKay, E.B., Massimino, D., Atkinson, S., Bacon, P.J., Brereton, T.M., Carvalho, L., Clutton-Brock, T.H., Duck, C., Edwards, M., Elliott, J.M., Hall, S.J.G., Harrington, R., Pearce-Higgins, J.W., Høye, T.T., Kruuk, L.E.B., Pemberton, J.M., Sparks, T.H., Thompson, P.M., White, I., Winfield, I.J., Wanless, S., 2016. Phenological sensitivity to climate across taxa and trophic levels. Nature 535 (7611), 241-245. doi: 10.1038/nature18608.

[79]	Thuiller, W., Lavergne, S., Roquet, C., Boulangeat, I., Lafourcade, B., Araujo, M.B., 2011. Consequences of climate change on the tree of life in Europe. Nature 470 (7335), 531-534. doi: 10.1038/nature09705.

[80]	Trew, B.T., Maclean, I.M., 2021. Vulnerability of global biodiversity hotspots to climate change. Glob. Ecol. Biogeogr. 30 (4), 768-783. doi: 10.1111/geb.13272.

[81]	Urban, M.C., 2024. Climate change extinctions. Science 386 (6726), 1123-1128. doi: 10.1126/science.adp4461.

[82]	Wang, Q., Hong, D.Y., 2022. Understanding the plant diversity on the roof of the world-a brief review of flora of Pan-Himalaya. Innovation 3 (2), 100215. doi: 10.1016/j.xinn.2022.100215.

[83]	Wani, I.A., Verma, S., Kumari, P., Charles, B., Hashim, M.J., El-Serehy, H.A., 2021. Ecological assessment and environmental niche modelling of Himalayan rhubarb (Rheum webbianum Royle) in northwest Himalaya . PLoS One 16 (11), e0259345. doi: 10.1371/journal.pone.0259345.

[84]	Wani, Z.A., Abdul Rahim, P.P., Dar, J.A., Lone, A.N., Siddiqui, S., 2025. Predicting the potential distribution of Podophyllum hexandrum Royle in the Himalaya under CMIP6 climate projections . Sci. Rep. 15 (1), 25374. doi: 10.1038/s41598-025-10862-w.

[85]	Wisz, M.S., Guisan, A., 2009. Do pseudo-absence selection strategies influence species distribution models and their predictions? An information-theoretic approach based on simulated data. BMC Ecol. 9, 8. doi: 10.1186/1472-6785-9-8.

[86]	Xu, J.C., Grumbine, R.E., 2014. Building ecosystem resilience for climate change adaptation in the Asian Highlands. Wiley Clim. Change 5 (6), 709-718. doi: 10.1002/wcc.302.

[87]

Yao, Y.-J., Wei, J.-C., Zhuang, W.-Y., Wei, T.-Z., Li, Y., Wei, X.-L., Deng, H., Liu, D.-M., Cai, L., Li, J.-S., Wang, K., Wu, H.J., Li, B.B., Wang, Y.H., Wei, X.D., Wu, H.M., Zhao, M.J., Yang, L., Su, J.H., Zhong, X., 2020. Threatened species list of China’s macrofungi. Biodivers. Sci. 28 (1), 20-25. doi: 10.17520/biods.2019174.

[88]	Zhang, C.L., Chen, Y., Xu, B.D., Xue, Y., Ren, Y.P., 2020. Improving prediction of rare species’ distribution from community data. Sci. Rep. 10 (1), 12230. doi: 10.1038/s41598-020-69157-x.

[89]	Zhu, K., Zhu, Y.W., Zhao, Z.Y., Wang, Y., Guo, X., Du, Y.X., Fu, J., Gao, J., 2025. Ecological vulnerability and driving factors in the Himalayan transboundary landscape under global climate change. Sci. Rep. 15 (1), 4895. doi: 10.1038/s41598-025-86811-4.