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This picture was taken at night in the coastal community of Dakenggu in Yilan County, which is just southeast of Taipei in Taiwan. I’m on the left, working with two other researchers to measure the body size of a red-clawed crab (Chiromantes haematocheir).An old man from the local community told me that years ago, during the breeding season, you could barely cross the road because of all the crabs. He said nobody knows where they all went. They’re an important memory for the local people, and part of the culture here.Habitat loss — especially resulting from the widespread use of concrete — seems to be driving the decline. I’m working with local people to create rocky microhabitats and artificial wetlands for the red-clawed crabs to live in. They’re important scavengers — eating dead animals and other organic matter, breaking it down and playing a key part in the nutrient cycle.Small organisms need our help — they can’t stand up for themselves. But in Taiwan, a lot of people think a coastal villa is more important than a few crabs. Corporations want to build luxury developments in our national parks, and authorities often approve them. I’ve seen so many intact habitats destroyed or covered in concrete.Crabs caught my interest because they were frequent visitors to my dormitory. National Sun Yat-sen University in Kaohsiung sits in a coastal buffer zone between a mountain and the ocean, and land hermit crabs (Coenobita cavipes) have to scurry through it on their way to breed.After watching habitat after habitat destroyed by overdevelopment, I’ve realized that just doing the science is not enough. It doesn’t matter how many papers you publish: you need to connect with people through education and communication. That’s why I decided to do my PhD in social science. And it’s why I believe conservation will be my life’s work.

Nature 603, 962 (2022)
doi: https://doi.org/10.1038/d41586-022-00810-3

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Laboratory for Integrative Biodiversity Research (LIBRe), Finnish Museum of Natural History (LUOMUS), University of Helsinki, Helsinki, FinlandStefano Mammola, Jagoba Malumbres-Olarte, Pedro Cardoso, Caroline S. Fukushima, Tuuli Korhonen, Marija Miličić & Joni A. SaarinenMolecular Ecology Group (MEG), Water Research Institute, National Research Council of Italy (CNR-IRSA), Largo Tonolli 50, 28922, Verbania Pallanza, ItalyStefano Mammola & Alejandro MartínezCE3C – Centre for Ecology, Evolution and Environmental Changes / Azorean Biodiversity Group and Universidade dos Açores, Angra do Heroísmo, Azores, PortugalJagoba Malumbres-OlarteAlbert Katz International School for Desert Studies, Ben-Gurion University of the Negev, Sede Boqer Campus, Beersheba, IsraelValeria ArabeskyBlaustein Institutes for Desert Research, Ben-Gurion University of the Negev, Sede Boqer Campus, Beersheba, IsraelValeria Arabesky & Yael LubinColección Nacional de Arácnidos, Instituto de Biología, Universidad Nacional Autónoma de México (UNAM), Mexico City, MexicoDiego Alejandro Barrales-AlcaláEnvironmental Biology Division, Institute of Biological Sciences, College of Arts and Sciences and Museum of Natural History, University of the Philippines Los Banos, 4031, Los Baños, PhilippinesAimee Lynn Barrion-DupoCentro Universitario de Rivera, Universidad de la República, Montevideo, UruguayMarco Antonio BenamúLab. Ecotoxicología de Artrópodos Terrestres, Centro Univeritario de Rivera, Universidad de la República, Montevideo, UruguayMarco Antonio BenamúLaboratorio Ecología del Comportamiento, Instituto de Investigaciones Biológicas clemente Estable (IIBCE), Montevideo, UruguayMarco Antonio BenamúDitsong National Museum of Natural History, PO Box 4197, Pretoria, 0001, South AfricaTharina L. BirdDepartment of Zoology and Entomology, University of Pretoria, Private Bag X20, Hatfield, 0028, South AfricaTharina L. BirdFreelance translator, Verbania Pallanza, ItalyMaria BogomolovaDepartment of Molecular Biology and Genetics, Democritus University of Thrace, Komotini, GreeceMaria ChatzakiDepartment of Life sciences, National Chung Hsing University, No.145 Xingda Rd., South Dist., Taichung City, 402204, TaiwanRen-Chung Cheng & Tien-Ai ChuDepartment of Biology, Macelwane Hall, 3507 Laclede Avenue, Saint Louis University, St. Louis, MO, 63103, USALeticia M. Classen-RodríguezCroatian Biospeleological Society, Rooseveltov trg 6, Zagreb, CroatiaIva Čupić & Martina PavlekProgram Sarjana, Fakultas Biologi, Universitas Gadjah Mada, Yogyakarta, IndonesiaNaufal Urfi Dhiya’ulhaqInsectarium de Montréal, Espace pour la vie, 4101, rue Sherbrooke Est, Montréal, Québec, H1X 2B2, CanadaAndré-Philippe Drapeau PicardSerket, Arachnid Collection of Egypt (ACE), Cairo, EgyptHisham K. El-HennawyErzincan Binali Yıldırım University, Faculty of Science and Arts, Biology Department, 24002, Erzincan, TurkeyMert ElvericiThe National Natural History Collections, The Hebrew University of Jerusalem, Edmond J. Safra Campus, Givat Ram, Jerusalem, 9190401, IsraelZeana Ganem & Efrat Gavish-RegevThe Department of Ecology, Evolution and Behavior, The Hebrew University of Jerusalem, Edmond J. Safra Campus, Givat Ram, Jerusalem, 9190401, IsraelZeana GanemBotswana International University of Science and Technology, Palapye, BotswanaNaledi T. GonnyeUMR CNRS 6553 Ecobio, Université de Rennes, 263 Avenue du Gal Leclerc, CS 74205, 35042, Rennes Cedex, FranceAxel Hacala & Julien PétillonDepartment of Zoology and Entomology, University of the Free State, P.O. Box 339, Bloemfontein, 9300, South AfricaCharles R. Haddad & Zingisile MboDepartment of Zoology, University of Oxford, Oxford, OX1 3PS, United KingdomThomas HesselbergDepartment of Biological Sciences, National University of Singapore, 14 Science Drive 4, Singapore, 117543, SingaporeTammy Ai Tian HoDepartment of Biotechnology, Faculty of Science and Technology, Thammasat University, Rangsit, Pathum Thani, 12121, ThailandThanakorn Into & Booppa PetcharadDept. of Life Science and Systems Biology, University of Torino, Via Accademia Albertina, 13 – 10123, Torino, ItalyMarco Isaia & Veronica NanniUnit of Conservation Biology, Department of Zoology, Bharathiar University, Coimbatore, 641046, Tamilnadu, IndiaDharmaraj JayaramanNational Museum of Namibia, Windhoek, NamibiaNanguei Karuaera5A Sagar Sangeet, SBS Marg, Mumbai, 400005, IndiaRajashree Khalap & Kiran KhalapDepartment of Biological Sciences, Ajou University, Suwon, Republic of KoreaDongyoung KimResearch Centre of the Slovenian Academy of Sciences and Arts, Jovan Hadži Institute of Biology, Ljubljana, SloveniaSimona Kralj-FišerUniversity of Greifswald, Zoological Institute and Museum, General and Systematic Zoology, Loitzerstrasse 26, 17489, Greifswald, GermanyHeidi Land, Shou-Wang Lin & Gabriele UhlDepartment of Natural Resource Sciences, McGill University, 21 111 Lakeshore Road, Sainte-Anne-de-Bellevue, Quebec, H9X 3V9, CanadaSarah Loboda & Catherine ScottDepartment of Biological Science, Macquarie University, Sydney, NSW, 2122, AustraliaElizabeth LoweMitrani Department of Desert Ecology, University in Midreshet Ben-Gurion, Midreshet Ben-Gurion, IsraelYael LubinBioSense Institute – Research Institute for Information Technologies in Biosystems, University of Novi Sad, Dr Zorana Đinđića 1, 21000, Novi Sad, SerbiaMarija MiličićNational Museums of Kenya, Museum Hill, P.O. BOX 40658- 00100, Nairobi, KenyaGrace Mwende KiokoSchool for Advanced Studies IUSS, Science, Technology and Society Department, 25100, Pavia, ItalyVeronica NanniInstitute of Biological Sciences, Faculty of Science, University of Malaya, 50603, Kuala Lumpur, MalaysiaYusoff Norma-RashidDepartment of Animal and Environmental Biology, Federal University, Oye-Ekiti, Ekiti State, NigeriaDaniel NwankwoTe Aka Mātuatua School of Science, University of Waikato, Private Bag 3105, Hamilton, 3240, New ZealandChristina J. PaintingIndependent researcher, Toronto, CanadaAleck PangMuseo Civico di Scienze Naturali “E. Caffi”, Piazza Cittadella, 10, I-24129, Bergamo, ItalyPaolo PantiniRuđer Bošković Institute, Bijenička cesta 54, 10000, Zagreb, CroatiaMartina PavlekBiodiversity Research Laboratory, Moreton Morrell, Warwickshire College University Centre, Warwickshire, United KingdomRichard PearceInstitute for Coastal and Marine Research, Nelson Mandela University, Port Elizabeth, South AfricaJulien PétillonDepartment of Entomology, University of Antananrivo, Antananarivo, MadagascarOnjaherizo Christian RaberahonaSchool of Biological Sciences, University of Nebraska-Lincoln, Lincoln, Nebraska, 68588, United StatesLaura Segura-HernándezDepartment of Biological Sciences, University of Toronto Scarborough, 1265 Military Trail, Scarborough, Ontario, M1C 1A4, CanadaLenka SentenskáNatural Sciences, Auckland War Memorial Museum, Parnell, Auckland, 1010, New ZealandLeilani WalkerTe Pūnaha Matatini, University of Auckland, Auckland, New ZealandLeilani WalkerMurang’a University of Technology, Department of Physical & Biological Sciences, P.O.Box 75-10200, Murang’a, KenyaCharles M. WaruiInstitute of Biology and Earth Sciences, Pomeranian University in Słupsk, Arciszewskiego 22a, 76-200, Słupsk, PolandKonrad WiśniewskiZoological Museum, Biodiversity Unit, FI-20014, University of Turku, Turku, FinlandAlireza ZamaniDepartment of Psychology, University of Tennessee, Knoxville, Tennessee, USAAngela ChuangDepartment of Entomology and Nematology, Citrus Research and Education Center, University of Florida, Lake Alfred, Florida, USAAngela ChuangConceptualization: SM, JM-O, CS, AC; Data collection & validation: all authors; Data management: SM, VN, AC; Data analysis & visualization (Figs. 2–5): SM; Summary illustration (Fig. 1): JM-O; Writing (first draft): SM; Writing, contributions: JM-O, CS, AC; All authors read the text, provided comments, suggestions, and corrections, and approved the final version. More

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Direct habitat lossAccording to the global projections of urban expansion under five SSPs17 (Supplementary Note 3 and Supplementary Fig. 1), 36–74 million hectares (Mha) of land areas will be urbanized by 2100, representing a 54–111% increase compared with the baseline year of 2015. Among these, 11–33 Mha natural habitats (Supplementary Table 1) will become urban areas by 2100. Across SSP scenarios, the patterns of change in losses of total habitat, forest, shrubland, and grassland are consistent with the global projections of urban expansion (Fig. 1). In terms of urban encroachment on wetlands, wetland will undergo the largest loss under scenario SSP4 than under other scenarios. However, if the sustainable pathway of scenario SSP1 is properly implemented, this will enable us to conserve the global wetland. The greatest loss of other habitat will occur under scenario SSP3, but the minimal loss of other habitat will occur under scenario SSP1. Under the five different SSP scenarios, the United States, Nigeria, Australia, Germany, and the UK are consistently predicted to have greater habitat loss due to urban expansion (Supplementary Table 2).Fig. 1: Future direct habitat loss due to urban expansion under SSP scenarios.a The habitat loss by 2100 for each habitat type. Bars indicate the mean habitat loss area (five scenarios) for each habitat type. Error bars represent mean values ± 1 SEM for the loss of each habitat type under five scenarios, n = 5 scenarios. Points represent data in five scenarios. b The losses in total area, forest, shrubland, grassland, wetland, and other land.Full size imageThere are obvious disparities in the hot spots and cold spots of habitat loss under the five SSP scenarios (Fig. 2 and Supplementary Figs. 2–6). Potential hot spots of habitat loss are concentrated in regions such as the northeastern, southern, and western coasts of the United States, the Gulf of Guinea coastal areas, Sub-Saharan Africa, and the Persian Gulf coastal areas. Under scenario SSP5, parts of central and western Europe will also become hot spots. However, under other scenarios, the cold spots will be particularly concentrated in eastern and southern Europe. East Asia and South Asia, which are represented by China, India, and Japan, are dominated by cold spots (Supplementary Figs. 2–6), because these regions may experience a decline in urban land demand from 2050 to 2100 (for examples in China, see Supplementary Figs. 7–11), although they are currently the most populous regions in the world.Fig. 2: Future hot spots and cold spots of habitat loss due to urban expansion under SSP scenarios by 2100.Figures for the United States (a), Europe (b), Africa (c), and China (d) are presented separately. The Gi_Bin identifies statistically significant hot spots and cold spots. Statistical significance was based on the p-value and z-score (two-sided), and no adjustments were made for multiple comparisons.Full size imageOur scenario projections show that the largest natural habitat loss is expected to occur in the temperate broadleaf and mixed forests biome (except for scenario SSP3). In addition, many biomes will experience proportionate loss of natural habitat. These biomes include the tropical and subtropical coniferous forests biome, the temperate coniferous forests biome, the flooded grasslands and savannas biome, the Mediterranean forests, woodlands, and scrub biome, and the mangroves biome (Supplementary Table 3). Although the rate of future habitat loss is small at the global scale, it can be large in some areas. For example, the habitat in the temperate broadleaf and mixed forests may decrease by 1.4% under scenario SSP5. At the ecoregion scale, about 9% of 867 terrestrial ecoregions will lose more than 1% of habitat due to urban expansion (Supplementary Fig. 12). In the future, four ecoregions—the Atlantic coastal pine barrens, the coastal forests of the northeastern United States, and the Puerto Rican moist and dry forests—will experience more than 20% of habitat loss.Urban expansion threatens biodiversity prioritization schemesTo reflect the potential impact of urban expansion on protected areas (Supplementary Note 4), the analyses presented here were based on the assumption that urban expansion within protected areas is not strictly restricted and can even occur in the currently gazetted protected areas (Supplementary Note 5, Supplementary Figs. 13 and 14). In 2015, urban areas with a total area of 30,594 km2 were distributed in 28,152 protected areas, accounting for 12.6% of global protected areas (Supplementary Figs. 15 and 16). Moreover, 38% of the urban land-use changes within protected areas were due to the conversion of natural habitats into urban land between 1992 and 2015. If urban expansion continues without strict restrictions, 13.2–19.8% of the protected areas will be affected by urban land by 2100, and urban land will occur in 29,563–44,400 protected areas with a total urban land area of up to 46,705–89,901 km2 across the five SSP scenarios (the lowest and highest proportions of urban land in each protected area by 2100 under SSP3 and SSP5 scenarios are presented in Supplementary Figs. 17 and 18).We also found that 0.90% of all terrestrial biodiversity hotspots (Supplementary Note 6), which are the world’s most biologically rich yet threatened terrestrial regions24, were urbanized in 2015. And this proportion (0.90%) is higher than that located in the rest of the Earth’s surface (0.51%) in 2015. By 2100, the new urban expansion will additionally occupy 1.5–1.8% of hotspot areas under the five SSP scenarios (Supplementary Table 4). Five biodiversity hotspots are projected to suffer the largest proportion of urban land conversion: the California Floristic Province (6–11%), Japan (6–8%), the North American Coastal Plain (4–8%), the Guinean Forests of West Africa (4–8%), and the Forests of East Australia (2–6%). In contrast, the East Melanesian Islands and the New Caledonia are almost unaffected by urban expansion. Biodiversity hotspots (e.g., the Guinean Forests of West Africa, the Coastal Forests of Eastern Africa, Eastern Afromontane, and the Polynesia-Micronesia) with few human disturbances in 2015 are projected to experience the highest percentage of future urban growth. Compared with the urban areas in 2015, by 2100, the urban areas in these four biodiversity hotspots will experience a disproportionate increase of 281–708, 294–535, 169–305, and 33–337%, respectively.The World Wildlife Fund (WWF) selected the ecoregions that are most crucial to the conservation of global biodiversity as Global 20025 (Supplementary Note 7). However, about 93% of the Global 200 ecoregions will be affected by future urban expansion. Although the proportion of urban land in each ecoregion will be less than 1% in 2100, the urban area located in these ecoregions will experience an increase of 74–160% from 2015 to 2100 across the five SSP scenarios (Supplementary Table 4). Four ecologically vulnerable ecoregions that have the highest urban growth rates are the Sudd-Sahelian Flooded Grasslands and Savannas, the East African Acacia Savannas, the Hawaii Moist Forest, and the Congolian Coastal Forests. By 2100, the urban areas in these four ecoregions will increase by 877–9955, 527–646, 18–902, and 500–1037%, respectively.The five SSP scenarios showed that the urban area is expected to increase by only 73–213 km2 in the Last of the Wild areas26 (see Supplementary Note 8 for descriptions about the Last of the Wild areas) by 2100 (Supplementary Table 4).Impacts of urban expansion on habitat fragmentationThe increasing exposures of natural habitat to urbanized land use may cause long-term changes in the function and structure of the natural habitat that is adjacent to urban areas13. To examine this proximity effect, we investigated the impact of future urban expansion on the nearest distance between urban areas and natural habitat (i.e., the distance from patch edges of urban areas to patch edges of the nearest natural habitats) under different SSP scenarios. Although the global urban area is expected to increase by 36–74 Mha by 2100, the impacts of future urban expansion on adjacent natural habitat are disproportionately large. Future urban expansion will make urban areas much closer to patch edges of 34–40 Mha natural habitat, which will inevitably threaten the natural habitat and increase the risk of biodiversity decline. The effects of urban expansion on adjacent patch edges of natural habitats are remarkably different across different scenarios. Specifically, the area of affected adjacent natural habitat is expected to be 38.45, 34.24, 40.31, 37.84, and 39.42 Mha under SSP1 to SSP5 scenarios by 2100, with the smallest effect under scenario SSP2, and the largest effect under scenario SSP3. Moreover, the scale of urban expansion does not correspond directly with the size of the impact. Several countries, including Mauritania, Algeria, Saudi Arabia, Western Sahara, and the United States, will have a large change in the distance from future urban areas to natural habitats due to urban expansion (Supplementary Table 5). Such effects also varied across different natural habitat types. The distance from the patch edges of urban areas to patch edges of (a) wetland, other land, and forest, (b) grassland, and (c) shrubland will generally be shortened by ~2000, ~1500 and ~900 m, respectively.In addition to the effect on the distance to the habitat edge, urban-caused habitat fragmentation is also reflected in reducing mean patch size (MPS)13, increasing mean edge index (edge density (ED), i.e., edge length on a per-unit area)27, and enlarging isolation (mean Euclidean nearest neighbor distance, ENN_MN)28 (Fig. 3). Taking the global ecoregions as the analysis unit, we found that within a 5 km buffer of urban areas, the median of MPS of natural habitats tends to show an overall decline trend, and the segmentation and subdivision of habitats become more obvious as future urban land expands. The median of MPS is the largest under scenario SSP1, followed by SSP4, SPP2, and SSP3 with some fluctuations in between, and the smallest MPS is found with the most fragmented landscape under scenario SSP5. A smaller patch size indicates that the inner parts of the habitat are subject to higher risk of being influenced by external disturbance. Future urban expansion also tends to cause an increase in the ED of natural habitat, which is often linked with smaller patches or more irregular shapes, and therefore poses a threat to biodiversity that influences many ecological processes (e.g., the spread of dispersal and predation)13,27,28. Scenario SSP1 shows the best performance in maintaining a low habitat ED and a high level of biodiversity conservation. However, under scenario SSP5, ED will experience a rapid increase in the second half of the 21st century. Meanwhile, the ENN_MN will increase substantially in the future, suggesting that areas with the same habitat type will become increasingly isolated, irregular, dispersed, or unevenly distributed due to the barrier of urban land. This will affect the speed of dispersal and patch recolonization. Scenario SSP1 is also most conducive to maintaining the proximity of natural habitats with the same habitat type. Other scenarios show relatively similar performance.Fig. 3: Future urban expansion effects on habitat fragmentation under SSP scenarios.a Mean patch size (MPS), b edge density (ED), c mean Euclidean nearest-neighbor distance (ENN_MN).Full size imageImpacts of urban expansion on terrestrial biodiversityWe focus on biodiversity in three common vertebrate taxa (i.e., amphibians, mammals, and birds) in our analyses. Future land system conversion to urban land will cause an average of 34% loss in the overall relative species richness. Land conversion from dense forest, mosaic grassland and open forest, mosaic grassland, and bare and natural grassland to urban land will cause the highest overall relative biodiversity loss (48%, 95% confidence interval (CI): 34–59% on a 1 km grid). These land systems with a high risk of biodiversity loss are concentrated in the United States, Europe, and Sub-Saharan Africa (Supplementary Fig. 19). Overall, the negative effect of future urban expansion on the total abundance of species will be more pronounced than that on species richness. Urban land changes will result in an average of 52% overall loss in relative total abundance of species. In particular, the losses of dense forest, natural grassland, and mosaic grassland, due to conversion to urban land, will lead to a high risk of species loss (62%, 95% CI: 38–76%).In terms of the number of species (i.e., all amphibians, mammals, and birds), future urban expansion will cause an average loss of 7–9 species and a loss of up to ~197 species per 10 km grid cell by 2100 across the five SSP scenarios (Fig. 4 and Supplementary Fig. 20). Species loss is most likely to be concentrated in Sub-Saharan Africa (particularly the Gulf of Guinea coast), the United States, and Europe. In addition, southeastern Brazil, India, and the eastern coast of Australia are also relatively high-risk areas. However, the specific effects of urban expansion vary substantially across different SSP scenarios. For instance, under scenario SSP5, urban expansion will pose a fatal threat to the global species richness in areas with urban development potential (species richness loss will occur in ~740 Mha land areas), whereas under the divided pathway (SSP4) and regional rivalry pathway (SSP3) scenarios, urban expansion will threaten the richest biodiversity hotspots, such as Sub-Saharan Africa and Latin America (Supplementary Fig. 20).Fig. 4: Potential biodiversity loss due to future urban expansion under SSP scenarios.The biodiversity loss in terms of the number of terrestrial vertebrate species (amphibians, mammals, and birds) lost per 10 km grid cell in the North America (a), Europe (b), the Gulf of Guinea coast (c), and East Asia (d).Full size imageWe also found a loss of up to 12 species of threatened amphibians, mammals, and birds (including vulnerable, endangered, or critically endangered categories defined in the IUCN Red List), and a loss of up to 40 species of small-ranged amphibians, mammals, and birds (small-ranged species are species with a geographic range size smaller than the median range size for that taxon)29 due to future urban expansion by 2100. There are a few scattered areas that will be hotspots for the loss of threatened species, such as West Africa, East Africa, northern India, and the eastern coast of Australia (Supplementary Fig. 21). The loss of small-ranged species will concentrate in fewer areas (Supplementary Fig. 22). We have identified 30 conservation priority ecoregions with high risks of habitat loss and small-ranged species loss due to future urban expansion (Supplementary Table 6). These conservation priority ecoregions are all found in Latin America and Sub-Saharan Africa (Supplementary Fig. 23). However, some hotspots outside of these conservation priority regions, such as tropical Southeast Asia, the west coast of the United States, and northern New Zealand, will also be affected (Supplementary Fig. 23).The top 5% 10 km grid cells with the highest loss in species richness (28–38 species potentially being lost) scatter across adjacent urban areas. However, only 6.4–8.6% of these regions are covered by the current global network of protected areas. These areas are often overlooked, and thus receive relatively low conservation spending. Ecoregions in Sub-Saharan African, Central and South America, Southeast Asia, and Australia will be responsible for the top 43% of average species loss across the SSP scenarios (Fig. 5). Kenya, Swaziland, Brunei, Zambia, Republic of Congo, and Zimbabwe will face the largest potential species richness loss (approximately > 29 species lost per 10 km grid cell) under all five SSP scenarios (Supplementary Fig. 24 and Supplementary Table 7).Fig. 5: Average potential biodiversity loss per 10 km grid cell in ecoregions due to future urban expansion under SSP scenarios.The mean potential biodiversity loss represents the average number of terrestrial vertebrate species (amphibians, mammals, and birds) lost per 10 km grid cell.Full size image More