{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,20]],"date-time":"2026-03-20T15:15:27Z","timestamp":1774019727424,"version":"3.50.1"},"reference-count":99,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,2,26]],"date-time":"2025-02-26T00:00:00Z","timestamp":1740528000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2025,2,26]],"date-time":"2025-02-26T00:00:00Z","timestamp":1740528000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["I 5825-B"],"award-info":[{"award-number":["I 5825-B"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["I 5825-B"],"award-info":[{"award-number":["I 5825-B"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["P 34688-B"],"award-info":[{"award-number":["P 34688-B"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["I 5825-B"],"award-info":[{"award-number":["I 5825-B"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002428","name":"Austrian Science Fund","doi-asserted-by":"publisher","award":["I 5825-B"],"award-info":[{"award-number":["I 5825-B"]}],"id":[{"id":"10.13039\/501100002428","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Programa de Investigaci\u00f3n en Cambio Clim\u00e1tico (PINCC), UNAM"},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["521529463"],"award-info":[{"award-number":["521529463"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"name":"European Union through the Life-SIP European Program"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Commun"],"abstract":"<jats:title>Abstract<\/jats:title>\n          <jats:p>The diverse biotas of the world\u2019s mountains face a challenging future due to increasing threats like climate change, land-use change, and biological invasions, the last being particularly understudied in these regions. Here we compile occurrence records for 717 alien vertebrate species distributed in 2984 mountains worldwide. We analyze their distribution, biogeographic origin, presence in protected areas, and the drivers\u2019 explaining alien vertebrate richness\u00a0in mountains. We find that the alien vertebrates most frequently recorded are birds (318 species) and mammals (161 species) reported in 2595 and 1518 mountains globally, respectively. The Palearctic, Nearctic, and Australasian realms are the most common recipients; the Nearctic, Indo-Malay, and Afrotropic realms are the most frequent donors. Almost 50% of the alien species studied also occur in protected areas. Proxies of anthropogenic impacts (e.g., higher road density or lower biodiversity intactness) and mountains\u2019 physical characteristics (e.g., elevation range and roughness) explain the distribution of alien vertebrates in mountains. Importantly, the magnitude of invasions in tropical mountains could be underestimated due to sampling bias towards the Northern Hemisphere and Australia. Our large-scale assessment reveals the advance of alien vertebrates in mountains worldwide and urges attention to minimize the impacts of biological invasions on the exceptional mountain biotas.<\/jats:p>","DOI":"10.1038\/s41467-025-57214-w","type":"journal-article","created":{"date-parts":[[2025,2,26]],"date-time":"2025-02-26T02:38:18Z","timestamp":1740537498000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["The global distribution patterns of alien vertebrate richness in mountains"],"prefix":"10.1038","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9831-2963","authenticated-orcid":false,"given":"Adri\u00e1n","family":"Garc\u00eda-Rodr\u00edguez","sequence":"first","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2616-3479","authenticated-orcid":false,"given":"Bernd","family":"Lenzner","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2183-5758","authenticated-orcid":false,"given":"Juli\u00e1n A.","family":"Velasco","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6682-7719","authenticated-orcid":false,"given":"Anna","family":"Schertler","sequence":"additional","affiliation":[]},{"given":"Ali","family":"Omer","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8993-6419","authenticated-orcid":false,"given":"Hanno","family":"Seebens","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0666-9755","authenticated-orcid":false,"given":"C\u00e9sar","family":"Capinha","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1552-8233","authenticated-orcid":false,"given":"Belinda","family":"Gallardo","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3919-0887","authenticated-orcid":false,"given":"Stefan","family":"Dullinger","sequence":"additional","affiliation":[]},{"given":"Franz","family":"Essl","sequence":"additional","affiliation":[]}],"member":"297","published-online":{"date-parts":[[2025,2,26]]},"reference":[{"key":"57214_CR1","doi-asserted-by":"publisher","first-page":"718","DOI":"10.1038\/s41561-018-0236-z","volume":"11","author":"A Antonelli","year":"2018","unstructured":"Antonelli, A. et al. Geological and climatic influences on mountain biodiversity. Nat. Geosci. 11, 718\u2013725 (2018).","journal-title":"Nat. Geosci."},{"key":"57214_CR2","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1007\/0044-7447-33.sp13.11","volume":"33","author":"C K\u00f6rner","year":"2004","unstructured":"K\u00f6rner, C. Mountain biodiversity, its causes and function. AMBIO: A J. Hum. Environ. 33, 11\u201317 (2004).","journal-title":"AMBIO: A J. Hum. Environ."},{"key":"57214_CR3","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1111\/jbi.13731","volume":"47","author":"A Perrigo","year":"2020","unstructured":"Perrigo, A., Hoorn, C. & Antonelli, A. Why mountains matter for biodiversity. J. Biogeogr. 47, 315\u2013325 (2020).","journal-title":"J. Biogeogr."},{"key":"57214_CR4","doi-asserted-by":"publisher","first-page":"711","DOI":"10.1111\/ecog.00578","volume":"37","author":"CH Graham","year":"2014","unstructured":"Graham, C. H. et al. The origin and maintenance of montane diversity: integrating evolutionary and ecological processes. Ecography 37, 711\u2013719 (2014).","journal-title":"Ecography"},{"key":"57214_CR5","doi-asserted-by":"publisher","first-page":"347","DOI":"10.1038\/nature14949","volume":"524","author":"VSFT Merckx","year":"2015","unstructured":"Merckx, V. S. F. T. et al. Evolution of endemism on a young tropical mountain. Nature 524, 347\u2013350 (2015).","journal-title":"Nature"},{"key":"57214_CR6","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1093\/aob\/mcr134","volume":"108","author":"SW Ansell","year":"2017","unstructured":"Ansell, S. W. et al. The importance of Anatolian mountains as the cradle of global diversity in Arabis alpina, a key arctic-alpine species. Ann. Bot. 108, 241\u2013252 (2017).","journal-title":"Ann. Bot."},{"key":"57214_CR7","doi-asserted-by":"publisher","first-page":"E68","DOI":"10.1086\/715500","volume":"198","author":"A Garcia-Rodriguez","year":"2021","unstructured":"Garcia-Rodriguez, A. et al. Amphibian Speciation Rates Support a General Role of Mountains as Biodiversity Pumps. Am. Natural. 198, E68\u2013E79 (2021).","journal-title":"Am. Natural."},{"key":"57214_CR8","doi-asserted-by":"publisher","first-page":"674","DOI":"10.1111\/geb.13249","volume":"30","author":"A Garc\u00eda\u2010Rodr\u00edguez","year":"2021","unstructured":"Garc\u00eda\u2010Rodr\u00edguez, A., Velasco, J. A., Villalobos, F. & Parra\u2010Olea, G. Effects of evolutionary time, speciation rates and local abiotic conditions on the origin and maintenance of amphibian montane diversity. Glob. Ecol. Biogeogr. 30, 674\u2013684 (2021).","journal-title":"Glob. Ecol. Biogeogr."},{"key":"57214_CR9","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1659\/MRD-JOURNAL-D-11-00058.1","volume":"31","author":"J L\u00f3pez-Pujol","year":"2011","unstructured":"L\u00f3pez-Pujol, J., Zhang, F.-M., Sun, H.-Q., Ying, T.-S. & Ge, S. Mountains of Southern China as \u2018Plant Museums\u2019 and \u2018Plant Cradles\u2019: evolutionary and conservation insights. Mt Res Dev. 31, 261\u2013269 (2011).","journal-title":"Mt Res Dev."},{"key":"57214_CR10","doi-asserted-by":"publisher","first-page":"e0271466","DOI":"10.1371\/journal.pone.0271466","volume":"17","author":"JM Thornton","year":"2022","unstructured":"Thornton, J. M. et al. Human populations in the world\u2019s mountains: Spatio-temporal patterns and potential controls. PLoS One 17, e0271466 (2022).","journal-title":"PLoS One"},{"key":"57214_CR11","doi-asserted-by":"crossref","unstructured":"Gr\u00eat-Regamey, A. & Weibel, B. Global assessment of mountain ecosystem services using earth observation data. Ecosyst. Serv. 46, (2020).","DOI":"10.1016\/j.ecoser.2020.101213"},{"key":"57214_CR12","doi-asserted-by":"publisher","first-page":"40","DOI":"10.1016\/j.cosust.2017.11.001","volume":"29","author":"D Payne","year":"2017","unstructured":"Payne, D., Spehn, E. M., Snethlage, M. & Fischer, M. Opportunities for research on mountain biodiversity under global change. Curr. Opin. Environ. Sustain 29, 40\u201347 (2017).","journal-title":"Curr. Opin. Environ. Sustain"},{"key":"57214_CR13","doi-asserted-by":"publisher","first-page":"1768","DOI":"10.1126\/science.1156831","volume":"320","author":"J Lenoir","year":"2008","unstructured":"Lenoir, J., G\u00e9gout, J. C., Marquet, P. A., De Ruffray, P. & Brisse, H. A significant upward shift in plant species optimum elevation during the 20th century.Science 320, 1768\u20131771 (2008).","journal-title":"Science"},{"key":"57214_CR14","doi-asserted-by":"publisher","first-page":"646","DOI":"10.1111\/geb.12142","volume":"23","author":"R Men\u00e9ndez","year":"2014","unstructured":"Men\u00e9ndez, R., Gonz\u00e1lez-Meg\u00edas, A., Jay-Robert, P. & Marqu\u00e9z-Ferrando, R. Climate change and elevational range shifts: Evidence from dung beetles in two European mountain ranges. Glob. Ecol. Biogeogr. 23, 646\u2013657 (2014).","journal-title":"Glob. Ecol. Biogeogr."},{"key":"57214_CR15","doi-asserted-by":"publisher","first-page":"1697","DOI":"10.1111\/ele.13762","volume":"24","author":"BG Freeman","year":"2021","unstructured":"Freeman, B. G., Song, Y., Feeley, K. J. & Zhu, K. Montane species track rising temperatures better in the tropics than in the temperate zone. Ecol. Lett. 24, 1697\u20131708 (2021).","journal-title":"Ecol. Lett."},{"key":"57214_CR16","doi-asserted-by":"publisher","first-page":"1675","DOI":"10.1038\/s41559-019-1042-8","volume":"3","author":"S Cauvy-Frauni\u00e9","year":"2019","unstructured":"Cauvy-Frauni\u00e9, S. & Dangles, O. A global synthesis of biodiversity responses to glacier retreat. Nat. Ecol. Evol. 3, 1675\u20131685 (2019).","journal-title":"Nat. Ecol. Evol."},{"key":"57214_CR17","doi-asserted-by":"publisher","first-page":"449","DOI":"10.1023\/A:1026033116193","volume":"18","author":"MG Turner","year":"2003","unstructured":"Turner, M. G., Pearson, S. M., Bolstad, P. & Wear, D. N. Effects of land-cover change on spatial pattern of forest communities in the Southern Appalachian Mountains (USA). Landsc. Ecol. 18, 449\u2013464 (2003).","journal-title":"Landsc. Ecol."},{"key":"57214_CR18","doi-asserted-by":"crossref","unstructured":"Tasser, E., Tappeiner, U. & Cernusca, A. Ecological effects of land-use changes in the European Alps. in Global change and Mountain Regions: An overview of Current knowledge (ed. Huber, U. M.) 409\u2013420 (2005).","DOI":"10.1007\/1-4020-3508-X_41"},{"key":"57214_CR19","doi-asserted-by":"publisher","first-page":"88","DOI":"10.1038\/s41586-019-1048-z","volume":"568","author":"MK Peters","year":"2019","unstructured":"Peters, M. K. et al. Climate\u2013land-use interactions shape tropical mountain biodiversity and ecosystem functions. Nature 568, 88\u201392 (2019).","journal-title":"Nature"},{"key":"57214_CR20","unstructured":"Seebens, H. et al. Trends and status of alien and invasive alien species. in Thematic Assessment Report on Invasive Alien Species and their Control of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (eds. Roy, H. E., Pauchard, A., Stoett, P. & R. Truong) (IPBES Secretariat, Bonn, Germany, 2023)."},{"key":"57214_CR21","doi-asserted-by":"publisher","first-page":"4199","DOI":"10.1007\/s10531-023-02697-6","volume":"32","author":"E Fuentes-Lillo","year":"2023","unstructured":"Fuentes-Lillo, E. et al. Going up the Andes: patterns and drivers of non-native plant invasions across latitudinal and elevational gradients. Biodivers. Conserv 32, 4199\u20134219 (2023).","journal-title":"Biodivers. Conserv"},{"key":"57214_CR22","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1038\/nclimate3337","volume":"7","author":"M Dainese","year":"2017","unstructured":"Dainese, M. et al. Human disturbance and upward expansion of plants in a warming climate. Nat. Clim. Chang 7, 577\u2013580 (2017).","journal-title":"Nat. Clim. Chang"},{"key":"57214_CR23","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1007\/s00035-016-0172-8","volume":"126","author":"JM Alexander","year":"2016","unstructured":"Alexander, J. M. et al. Plant invasions into mountains and alpine ecosystems: current status and future challenges. Alp. Bot. 126, 89\u2013103 (2016).","journal-title":"Alp. Bot."},{"key":"57214_CR24","doi-asserted-by":"publisher","first-page":"e2201911120","DOI":"10.1073\/pnas.2201911120","volume":"120","author":"C Capinha","year":"2023","unstructured":"Capinha, C., Essl, F., Porto, M. & Seebens, H. The worldwide networks of spread of recorded alien species. Proc. Natl Acad. Sci. USA 120, e2201911120 (2023).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"57214_CR25","doi-asserted-by":"publisher","first-page":"571","DOI":"10.1038\/s41586-021-03405-6","volume":"592","author":"C Diagne","year":"2021","unstructured":"Diagne, C. et al. High and rising economic costs of biological invasions worldwide. Nature 592, 571\u2013576 (2021).","journal-title":"Nature"},{"key":"57214_CR26","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3897\/neobiota.32.10199","volume":"32","author":"J Pergl","year":"2017","unstructured":"Pergl, J. et al. Troubling travellers: Are ecologically harmful alien species associated with particular introduction pathways? NeoBiota 32, 1\u201320 (2017).","journal-title":"NeoBiota"},{"key":"57214_CR27","doi-asserted-by":"publisher","first-page":"1511","DOI":"10.1111\/brv.12627","volume":"95","author":"P Py\u0161ek","year":"2020","unstructured":"Py\u0161ek, P. et al. Scientists\u2019 warning on invasive alien species. Biol. Rev. 95, 1511\u20131534 (2020).","journal-title":"Biol. Rev."},{"key":"57214_CR28","doi-asserted-by":"crossref","unstructured":"Rabitsch, W., Essl, F. & Schindler, S. The Rise of Non-native Vectors and Reservoirs of Human Diseases. in IMPACT OF BIOLOGICAL INVASIONS ON ECOSYSTEM SERVICES (eds Vila, M. & Hulme, P. E.)12 263\u2013275 (SPRINGER INTERNATIONAL PUBLISHING AG, GEWERBESTRASSE 11, CHAM, CH-6330, SWITZERLAND, 2017).","DOI":"10.1007\/978-3-319-45121-3_17"},{"key":"57214_CR29","doi-asserted-by":"crossref","unstructured":"Dawson, W. et al. Global hotspots and correlates of alien species richness across taxonomic groups. Nat. Ecol. Evol. 1, (2017).","DOI":"10.1038\/s41559-017-0186"},{"key":"57214_CR30","doi-asserted-by":"publisher","first-page":"405","DOI":"10.1038\/s41559-022-01979-6","volume":"7","author":"E Iseli","year":"2023","unstructured":"Iseli, E. et al. Rapid upwards spread of non-native plants in mountains across continents. Nat. Ecol. Evol. 7, 405\u2013413 (2023).","journal-title":"Nat. Ecol. Evol."},{"key":"57214_CR31","doi-asserted-by":"publisher","first-page":"211","DOI":"10.3391\/ai.2014.9.2.10","volume":"9","author":"AS Quan","year":"2014","unstructured":"Quan, A. S., Pease, K. M., Breinholt, J. W. & Wayne, R. K. Origins of the invasive red swamp crayfish (Procambarus clarkii) in the Santa Monica Mountains. Aquat. Invasions 9, 211\u2013219 (2014).","journal-title":"Aquat. Invasions"},{"key":"57214_CR32","doi-asserted-by":"publisher","first-page":"791","DOI":"10.1007\/s10530-014-0769-z","volume":"17","author":"A Mir\u00f3","year":"2015","unstructured":"Mir\u00f3, A. & Ventura, M. Evidence of exotic trout mediated minnow invasion in Pyrenean high mountain lakes. Biol. Invasions 17, 791\u2013803 (2015).","journal-title":"Biol. Invasions"},{"key":"57214_CR33","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1002\/wsb.636","volume":"40","author":"EP Flesch","year":"2016","unstructured":"Flesch, E. P. et al. Range expansion and population growth of non-native mountain goats in the Greater Yellowstone Area: Challenges for management. Wildl. Soc. Bull. 40, 241\u2013250 (2016).","journal-title":"Wildl. Soc. Bull."},{"key":"57214_CR34","doi-asserted-by":"publisher","first-page":"3205","DOI":"10.1007\/s10530-010-9712-0","volume":"12","author":"BC L\u00f3pez","year":"2010","unstructured":"L\u00f3pez, B. C., Pino, J. & L\u00f3pez, A. Explaining the successful introduction of the alpine marmot in the Pyrenees. Biol. Invasions 12, 3205\u20133217 (2010).","journal-title":"Biol. Invasions"},{"key":"57214_CR35","doi-asserted-by":"crossref","unstructured":"Joshi, S., Shrestha, B. B., Shrestha, L., Rashid, I. & Adkins, S. Plant Invasions in Mountains. in Global Plant Invasions (eds. Clements, D. R., Upadhyaya, M. K., Joshi, S. & Shrestha, A.) 279\u2013300 (2022).","DOI":"10.1007\/978-3-030-89684-3_13"},{"key":"57214_CR36","doi-asserted-by":"publisher","first-page":"1862","DOI":"10.1038\/s41559-017-0365-6","volume":"1","author":"C Bellard","year":"2017","unstructured":"Bellard, C., Rysman, J. F., Leroy, B., Claud, C. & Mace, G. M. A global picture of biological invasion threat on islands. Nat. Ecol. Evol. 1, 1862\u20131869 (2017).","journal-title":"Nat. Ecol. Evol."},{"key":"57214_CR37","unstructured":"Bacher, S. et al. Impacts of biological invasions on nature, nature\u2019s contributions to people, and good quality of life. in Thematic Assessment Report on Invasive Alien Species and their Control of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services. (eds Roy, H. et al.) (IPBES Secretariat, Bonn, Germany, 2023)."},{"key":"57214_CR38","doi-asserted-by":"crossref","unstructured":"Meyer, C., Kreft, H., Guralnick, R. & Jetz, W. Global priorities for an effective information basis of biodiversity distributions. Nat. Commun. 6, (2015).","DOI":"10.1038\/ncomms9221"},{"key":"57214_CR39","doi-asserted-by":"publisher","first-page":"1259","DOI":"10.1111\/ecog.05926","volume":"44","author":"AC Hughes","year":"2021","unstructured":"Hughes, A. C. et al. Sampling biases shape our view of the natural world. Ecography 44, 1259\u20131269 (2021).","journal-title":"Ecography"},{"key":"57214_CR40","doi-asserted-by":"publisher","first-page":"992","DOI":"10.1111\/ele.12624","volume":"19","author":"C Meyer","year":"2016","unstructured":"Meyer, C., Weigelt, P. & Kreft, H. Multidimensional biases, gaps and uncertainties in global plant occurrence information. Ecol. Lett. 19, 992\u20131006 (2016).","journal-title":"Ecol. Lett."},{"key":"57214_CR41","doi-asserted-by":"publisher","first-page":"e3001296","DOI":"10.1371\/journal.pbio.3001296","volume":"19","author":"T Amano","year":"2021","unstructured":"Amano, T. et al. Tapping into non-English-language science for the conservation of global biodiversity. PLoS Biol. 19, e3001296 (2021).","journal-title":"PLoS Biol."},{"key":"57214_CR42","doi-asserted-by":"publisher","first-page":"e0206373","DOI":"10.1371\/journal.pone.0206373","volume":"13","author":"LD Einoder","year":"2018","unstructured":"Einoder, L. D. et al. Occupancy and detectability modelling of vertebrates in northern Australia using multiple sampling methods. PLoS One 13, e0206373 (2018).","journal-title":"PLoS One"},{"key":"57214_CR43","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-021-98584-7","volume":"11","author":"CT Callaghan","year":"2021","unstructured":"Callaghan, C. T., Poore, A. G. B., Hofmann, M., Roberts, C. J. & Pereira, H. M. Large-bodied birds are over-represented in unstructured citizen science data. Sci. Rep. 11, 19073 (2021).","journal-title":"Sci. Rep."},{"key":"57214_CR44","doi-asserted-by":"crossref","unstructured":"van Kleunen, M. et al. Global exchange and accumulation of non-native plants. Nature 525, 100\u2013103 (2015).","DOI":"10.1038\/nature14910"},{"key":"57214_CR45","doi-asserted-by":"crossref","unstructured":"Bellard, C., Leroy, B., Thuiller, W., Rysman, J. F. & Courchamp, F. Major drivers of invasion risks throughout the world. Ecosphere 7, e01241 (2016).","DOI":"10.1002\/ecs2.1241"},{"key":"57214_CR46","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1002\/fee.2059","volume":"17","author":"JL Lockwood","year":"2019","unstructured":"Lockwood, J. L. et al. When pets become pests: the role of the exotic pet trade in producing invasive vertebrate animals. Front. Ecol. Environ. 17, 323\u2013330 (2019).","journal-title":"Front. Ecol. Environ."},{"key":"57214_CR47","doi-asserted-by":"publisher","first-page":"1313","DOI":"10.1111\/ddi.12617","volume":"23","author":"C Capinha","year":"2017","unstructured":"Capinha, C. et al. Diversity, biogeography and the global flows of alien amphibians and reptiles. Divers Distrib. 23, 1313\u20131322 (2017).","journal-title":"Divers Distrib."},{"key":"57214_CR48","doi-asserted-by":"publisher","first-page":"1632","DOI":"10.1111\/geb.13714","volume":"32","author":"R Mu\u00f1oz-Mas","year":"2023","unstructured":"Mu\u00f1oz-Mas, R. et al. Two centuries of spatial and temporal dynamics of freshwater fish introductions. Glob. Ecol. Biogeogr. 32, 1632\u20131644 (2023).","journal-title":"Glob. Ecol. Biogeogr."},{"key":"57214_CR49","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1017\/dsj.2016.16","volume":"3","author":"L Reino","year":"2017","unstructured":"Reino, L. et al. Networks of global bird invasion altered by regional trade ban. Science\u00a03, 1\u20138 (2017).","journal-title":"Science"},{"key":"57214_CR50","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1126\/science.aav5327","volume":"366","author":"BR Scheffers","year":"2019","unstructured":"Scheffers, B. R., Oliveira, B. F., Lamb, I. & Edwards, D. P. Global wildlife trade across the tree of life. Science\u00a0366, 71\u201376 (2019).","journal-title":"Science"},{"key":"57214_CR51","doi-asserted-by":"publisher","first-page":"977","DOI":"10.1093\/biosci\/biab056","volume":"71","author":"JS Sinclair","year":"2021","unstructured":"Sinclair, J. S. et al. The international vertebrate pet trade network and insights from US imports of exotic pets. Bioscience 71, 977\u2013990 (2021).","journal-title":"Bioscience"},{"key":"57214_CR52","doi-asserted-by":"crossref","unstructured":"Liu, X. et al. Animal invaders threaten protected areas worldwide. Nat. Commun. 11, 2892 (2020).","DOI":"10.1038\/s41467-020-16719-2"},{"key":"57214_CR53","doi-asserted-by":"publisher","first-page":"5331","DOI":"10.1111\/gcb.13798","volume":"23","author":"B Gallardo","year":"2017","unstructured":"Gallardo, B. et al. Protected areas offer refuge from invasive species spreading under climate change. Glob. Chang Biol. 23, 5331\u20135343 (2017).","journal-title":"Glob. Chang Biol."},{"key":"57214_CR54","doi-asserted-by":"crossref","unstructured":"Tiago, P., Ceia-Hasse, A., Marques, T. A., Capinha, C. & Pereira, H. M. Spatial distribution of citizen science casuistic observations for different taxonomic groups. Sci. Rep. 7, 12832 (2017).","DOI":"10.1038\/s41598-017-13130-8"},{"key":"57214_CR55","doi-asserted-by":"publisher","first-page":"1187","DOI":"10.1111\/ecog.02194","volume":"40","author":"T Mang","year":"2017","unstructured":"Mang, T. et al. Accounting for imperfect observation and estimating true species distributions in modelling biological invasions. Ecography 40, 1187\u20131197 (2017).","journal-title":"Ecography"},{"key":"57214_CR56","doi-asserted-by":"publisher","first-page":"2751","DOI":"10.1007\/s10530-017-1481-6","volume":"19","author":"ML Thomas","year":"2017","unstructured":"Thomas, M. L. et al. Many eyes on the ground: citizen science is an effective early detection tool for biosecurity. Biol. Invasions 19, 2751\u20132765 (2017).","journal-title":"Biol. Invasions"},{"key":"57214_CR57","doi-asserted-by":"publisher","first-page":"1248","DOI":"10.1126\/science.aaa8913","volume":"348","author":"C Capinha","year":"2015","unstructured":"Capinha, C., Essl, F., Seebens, H., Moser, D. & Pereira, H. M. The dispersal of alien species redefines biogeography in the Anthropocene. Science\u00a0348, 1248\u20131251 (2015).","journal-title":"Science"},{"key":"57214_CR58","doi-asserted-by":"publisher","first-page":"656","DOI":"10.1073\/pnas.1013136108","volume":"108","author":"JM Alexander","year":"2011","unstructured":"Alexander, J. M. et al. Assembly of nonnative floras along elevational gradients explained by directional ecological filtering. Proc. Natl Acad. Sci. USA 108, 656\u2013661 (2011).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"57214_CR59","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1086\/282487","volume":"101","author":"DH Janzen","year":"1967","unstructured":"Janzen, D. H. Why Mountain Passes are Higher in the Tropics. Am. Nat. 101, 233 (1967).","journal-title":"Am. Nat."},{"key":"57214_CR60","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1146\/annurev-ecolsys-102220-013423","volume":"54","author":"SL Chown","year":"2023","unstructured":"Chown, S. L. & Mcgeoch, M. A. Functional trait variation along animal invasion pathways. Annu Rev. Ecol. Evol. Syst. 54, 151\u2013170 (2023).","journal-title":"Annu Rev. Ecol. Evol. Syst."},{"key":"57214_CR61","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1146\/annurev-ecolsys-102722-013135","volume":"55","author":"BA Bradley","year":"2024","unstructured":"Bradley, B. A. et al. Observed and potential range shifts of native and nonnative species with climate change. Annu Rev. Ecol. Evol. Syst. 55, 23\u201340 (2024).","journal-title":"Annu Rev. Ecol. Evol. Syst."},{"key":"57214_CR62","doi-asserted-by":"publisher","first-page":"1097","DOI":"10.1111\/geb.12469","volume":"25","author":"MJ Steinbauer","year":"2016","unstructured":"Steinbauer, M. J. et al. Topography-driven isolation, speciation and a global increase of endemism with elevation. Glob. Ecol. Biogeogr. 25, 1097\u20131107 (2016).","journal-title":"Glob. Ecol. Biogeogr."},{"key":"57214_CR63","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41467-021-27603-y","volume":"12","author":"Q Yang","year":"2021","unstructured":"Yang, Q. et al. The global loss of floristic uniqueness. Nat. Commun. 12, 1\u201310 (2021).","journal-title":"Nat. Commun."},{"key":"57214_CR64","doi-asserted-by":"crossref","unstructured":"Haider, S. et al. Think globally, measure locally: The MIREN standardized protocol for monitoring plant species distributions along elevation gradients. Ecol. Evol. 12, e8590 (2022).","DOI":"10.1002\/ece3.8590"},{"key":"57214_CR65","unstructured":"Barros, A. et al. The role of roads and trails for facilitating mountain plant invasions. in Tourism, Recreation and Biological Invasions. 14\u201326 (GB: CABI, 2022)."},{"key":"57214_CR66","doi-asserted-by":"publisher","first-page":"420","DOI":"10.1016\/j.gloenvcha.2006.11.007","volume":"17","author":"D Nogu\u00e9s-Bravo","year":"2007","unstructured":"Nogu\u00e9s-Bravo, D., Ara\u00fajo, M. B., Errea, M. P. & Mart\u00ednez-Rica, J. P. Exposure of global mountain systems to climate warming during the 21st Century. Glob. Environ. Change 17, 420\u2013428 (2007).","journal-title":"Glob. Environ. Change"},{"key":"57214_CR67","doi-asserted-by":"publisher","first-page":"784","DOI":"10.1111\/geb.13246","volume":"30","author":"MA Mamantov","year":"2021","unstructured":"Mamantov, M. A., Gibson-Reinemer, D. K., Linck, E. B. & Sheldon, K. S. Climate-driven range shifts of montane species vary with elevation. Glob. Ecol. Biogeogr. 30, 784\u2013794 (2021).","journal-title":"Glob. Ecol. Biogeogr."},{"key":"57214_CR68","doi-asserted-by":"publisher","first-page":"3040","DOI":"10.1111\/gcb.15067","volume":"26","author":"JA Rehbein","year":"2020","unstructured":"Rehbein, J. A. et al. Renewable energy development threatens many globally important biodiversity areas. Glob. Chang Biol. 26, 3040\u20133051 (2020).","journal-title":"Glob. Chang Biol."},{"key":"57214_CR69","doi-asserted-by":"crossref","unstructured":"Vorstenbosch, T., Essl, F. & Lenzner, B. An uphill battle? The elevational distribution of alien plant species along rivers and roads in the Austrian Alps. Neobiota 1\u201324 (2020).","DOI":"10.3897\/neobiota.63.55096"},{"key":"57214_CR70","doi-asserted-by":"publisher","first-page":"1121","DOI":"10.1007\/s10530-019-02165-9","volume":"22","author":"R Liedtke","year":"2020","unstructured":"Liedtke, R. et al. Hiking trails as conduits for the spread of non-native species in mountain areas. Biol. Invasions 22, 1121\u20131134 (2020).","journal-title":"Biol. Invasions"},{"key":"57214_CR71","doi-asserted-by":"crossref","unstructured":"Rodrigues, A. S. L. et al. Global gap analysis: Priority regions for expanding the Global Protected-Area Network. Bioscience 54, 1092\u20131100 (2004).","DOI":"10.1641\/0006-3568(2004)054[1092:GGAPRF]2.0.CO;2"},{"key":"57214_CR72","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41597-022-01256-y","volume":"9","author":"MA Snethlage","year":"2022","unstructured":"Snethlage, M. A. et al. A hierarchical inventory of the world\u2019s mountains for global comparative mountain science. Sci. Data 9, 1\u201314 (2022).","journal-title":"Sci. Data"},{"key":"57214_CR73","doi-asserted-by":"publisher","unstructured":"Snethlage, M. A. et al. GMBA Mountain Inventory v2. GMBA-EarthEnv https:\/\/doi.org\/10.48601\/earthenv-t9k2-1407 (2022).","DOI":"10.48601\/earthenv-t9k2-1407"},{"key":"57214_CR74","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s00035-016-0182-6","volume":"127","author":"C K\u00f6rner","year":"2017","unstructured":"K\u00f6rner, C. et al. A global inventory of mountains for biogeographical applications. Alp. Bot. 127, 1\u201315 (2017).","journal-title":"Alp. Bot."},{"key":"57214_CR75","doi-asserted-by":"publisher","first-page":"75","DOI":"10.3897\/neobiota.74.81082","volume":"74","author":"H Seebens","year":"2022","unstructured":"Seebens, H. & Kaplan, E. DASCO: A workflow to downscale alien species checklists using occurrence records and to re-allocate species distributions across realms. NeoBiota 74, 75\u201391 (2022).","journal-title":"NeoBiota"},{"key":"57214_CR76","first-page":"9","volume":"5","author":"E Pebesma","year":"2005","unstructured":"Pebesma, E. & Bivand, R. Classes and methods for spatial data in. R. R. N. 5, 9\u201313 (2005).","journal-title":"R. R. N."},{"key":"57214_CR77","doi-asserted-by":"publisher","first-page":"439","DOI":"10.32614\/RJ-2018-009","volume":"10","author":"E Pebesma","year":"2018","unstructured":"Pebesma, E. Simple Features for R: Standardized Support for Spatial Vector Data. R. J. 10, 439 (2018).","journal-title":"R. J."},{"key":"57214_CR78","unstructured":"Hijmans, R. terra: Spatial Data Analysis. R package. CRAN (2024)."},{"key":"57214_CR79","doi-asserted-by":"publisher","first-page":"933","DOI":"10.1641\/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2","volume":"51","author":"DM Olson","year":"2001","unstructured":"Olson, D. M. et al. Terrestrial ecoregions of the world: A new map of life on Earth. Bioscience 51, 933\u2013938 (2001).","journal-title":"Bioscience"},{"key":"57214_CR80","doi-asserted-by":"publisher","first-page":"2811","DOI":"10.1093\/bioinformatics\/btu393","volume":"30","author":"Z Gu","year":"2014","unstructured":"Gu, Z., Gu, L., Eils, R., Schlesner, M. & Brors, B. Circlize implements and enhances circular visualization in. R. Bioinforma. 30, 2811\u20132812 (2014).","journal-title":"R. Bioinforma."},{"key":"57214_CR81","doi-asserted-by":"publisher","first-page":"788","DOI":"10.1126\/science.aap9565","volume":"360","author":"KR Jones","year":"2018","unstructured":"Jones, K. R. et al. One-third of global protected land is under intense human pressure. Science\u00a0360, 788\u2013791 (2018).","journal-title":"Science"},{"key":"57214_CR82","doi-asserted-by":"crossref","unstructured":"Meijer, J. R., Huijbregts, M. A. J., Schotten, K. C. G. J. & Schipper, A. M. Global patterns of current and future road infrastructure. Environ. Res. Lett. 13, 064006 (2018).","DOI":"10.1088\/1748-9326\/aabd42"},{"key":"57214_CR83","doi-asserted-by":"publisher","first-page":"288","DOI":"10.1126\/science.aaf2201","volume":"353","author":"T Newbold","year":"2016","unstructured":"Newbold, T. et al. Has land use pushed terrestrial biodiversity beyond the planetary boundary? A global assessment. Science\u00a0353, 288\u2013291 (2016).","journal-title":"Science"},{"key":"57214_CR84","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41597-019-0265-5","volume":"6","author":"A Nelson","year":"2019","unstructured":"Nelson, A. et al. A suite of global accessibility indicators. Sci. Data 6, 1\u20139 (2019).","journal-title":"Sci. Data"},{"key":"57214_CR85","doi-asserted-by":"publisher","first-page":"1052","DOI":"10.1038\/nature08649","volume":"462","author":"SR Loarie","year":"2009","unstructured":"Loarie, S. R. et al. The velocity of climate change. Nature 462, 1052\u20131055 (2009).","journal-title":"Nature"},{"key":"57214_CR86","doi-asserted-by":"publisher","DOI":"10.1038\/s41597-020-0453-3","volume":"7","author":"I Harris","year":"2020","unstructured":"Harris, I., Osborn, T. J., Jones, P. & Lister, D. Version 4 of the CRU TS monthly high-resolution gridded multivariate climate dataset. Sci. Data 7, 109 (2020).","journal-title":"Sci. Data"},{"key":"57214_CR87","doi-asserted-by":"publisher","first-page":"2195","DOI":"10.1111\/2041-210X.13295","volume":"10","author":"J Garc\u00eda Molinos","year":"2019","unstructured":"Garc\u00eda Molinos, J., Schoeman, D. S., Brown, C. J. & Burrows, M. T. VoCC: An r package for calculating the velocity of climate change and related climatic metrics. Methods Ecol. Evol. 10, 2195\u20132202 (2019).","journal-title":"Methods Ecol. Evol."},{"key":"57214_CR88","doi-asserted-by":"publisher","first-page":"3","DOI":"10.1080\/01490410701295962","volume":"30","author":"MFJ Wilson","year":"2007","unstructured":"Wilson, M. F. J., O\u2019Connell, B., Brown, C., Guinan, J. C. & Grehan, A. J. Multiscale terrain analysis of multibeam bathymetry data for habitat mapping on the continental slope. Mar. Geod. 30, 3\u201335 (2007).","journal-title":"Mar. Geod."},{"key":"57214_CR89","doi-asserted-by":"publisher","first-page":"583","DOI":"10.1029\/EO081i048p00583","volume":"81","author":"TG Farr","year":"2000","unstructured":"Farr, T. G. & Kobrick, M. Shuttle radar topography mission produces a wealth of data. Eos (Wash. DC) 81, 583\u2013585 (2000).","journal-title":"Eos (Wash. DC)"},{"key":"57214_CR90","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1038\/nclimate2656","volume":"5","author":"P Elsen","year":"2015","unstructured":"Elsen, P. & Tingley, M. W. Global mountain topography and the fate of montane species under climate change. Nat. Clim. Chang 5, 772\u2013776 (2015).","journal-title":"Nat. Clim. Chang"},{"key":"57214_CR91","unstructured":"Komsta, L. & Novomestky F. moments: Moments, Cumulants, Skewness, Kurtosis and Related Tests. R package. (2022)."},{"key":"57214_CR92","doi-asserted-by":"publisher","first-page":"2042","DOI":"10.1111\/ddi.13607","volume":"28","author":"M Monteiro","year":"2022","unstructured":"Monteiro, M. et al. Patterns and drivers of the global diversity of non-native macrofungi. Divers Distrib. 28, 2042\u20132055 (2022).","journal-title":"Divers Distrib."},{"key":"57214_CR93","doi-asserted-by":"crossref","unstructured":"Schertler, A. et al. Biogeography and global flows of 100 major alien fungal and fungus-like oomycete pathogens. J Biogeogr. 1\u201318 (2023).","DOI":"10.1111\/jbi.14755"},{"key":"57214_CR94","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1111\/j.1600-0587.2012.07348.x","volume":"36","author":"CF Dormann","year":"2012","unstructured":"Dormann, C. F. et al. Collinearity: a review of methods to deal with it and a simulation study evaluating their performance. Ecography 36, 27\u201346 (2012).","journal-title":"Ecography"},{"key":"57214_CR95","doi-asserted-by":"publisher","first-page":"791","DOI":"10.1086\/283438","volume":"113","author":"EF Connor","year":"1979","unstructured":"Connor, E. F. & McCoy, E. D. The statistics and biology of the species-area relationship. Am. Nat. 113, 791\u2013833 (1979).","journal-title":"Am. Nat."},{"key":"57214_CR96","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v067.i01","volume":"67","author":"D Bates","year":"2015","unstructured":"Bates, D., M\u00e4chler, M., Bolker, B. & Walker, S. Fitting linear mixed-effects models using lme4. J. Stat. Softw. 67, 1\u201348 (2015).","journal-title":"J. Stat. Softw."},{"key":"57214_CR97","unstructured":"Wood, S. & Wood, M. S. Package \u2018mgcv\u2019. R package Version 1, 729 (2015)."},{"key":"57214_CR98","doi-asserted-by":"publisher","unstructured":"Garcia-Rodriguez, A. The global distribution patterns of alien vertebrate richness in mountains. Zenodo https:\/\/doi.org\/10.5281\/zenodo.14751462 (2025).","DOI":"10.5281\/zenodo.14751462"},{"key":"57214_CR99","doi-asserted-by":"publisher","first-page":"1247579","DOI":"10.1126\/science.1247579","volume":"344","author":"RA Garcia","year":"2014","unstructured":"Garcia, R. A., Cabeza, M., Rahbek, C. & Ara\u00fajo, M. B. Multiple dimensions of climate change and their implications for biodiversity. Science\u00a0344, 1247579 (2014).","journal-title":"Science"}],"container-title":["Nature Communications"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41467-025-57214-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41467-025-57214-w","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41467-025-57214-w.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,2,26]],"date-time":"2025-02-26T03:17:14Z","timestamp":1740539834000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41467-025-57214-w"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,2,26]]},"references-count":99,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,12]]}},"alternative-id":["57214"],"URL":"https:\/\/doi.org\/10.1038\/s41467-025-57214-w","relation":{},"ISSN":["2041-1723"],"issn-type":[{"value":"2041-1723","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,2,26]]},"assertion":[{"value":"22 March 2024","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"14 February 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"26 February 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"1977"}}