AbstractHabitat fragmentation generally reduces biodiversity at the patch scale (α diversity)1. However, there is ongoing debate about whether such negative effects can be alleviated at the landscape scale (γ diversity) if among-patch diversity (β diversity) increases as a result of fragmentation2,3,4,5,6. This controversial view has not been rigorously tested. Here we use a dataset of 4,006 taxa across 37 studies from 6 continents to test the effects of fragmentation on biodiversity across scales by explicitly comparing continuous and fragmented landscapes. We find that fragmented landscapes consistently have both lower α diversity and lower γ diversity. Although fragmented landscapes did tend to have higher β diversity, this did not translate into higher γ diversity. Our findings refute claims that habitat fragmentation can increase biodiversity at landscape scales, and emphasize the need to restore habitat and increase connectivity to minimize biodiversity loss at ever-increasing scales.
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Fig. 1: Habitat amount and distance decay predict different patterns of α, β and γ diversity.Fig. 2: Differences in α, β and γ diversity in continuous and fragmented landscapes.Fig. 3: Differences in α, β and γ diversity in continuous and fragmented landscapes controlling for differences in sampling effort and using all plot pairs.
Data availability
The datasets used in this paper are available at GitHub (https://github.com/thiago-goncalves-souza/ms-biodiversity-loss-fragmented-landscapes) and Zenodo (https://zenodo.org/records/14885581)61.
Code availability
The code used in this paper is available at GitHub (https://github.com/thiago-goncalves-souza/ms-biodiversity-loss-fragmented-landscapes) and Zenodo (https://zenodo.org/records/14885581)61.
ReferencesHaddad, N. M. et al. Habitat fragmentation and its lasting impact on Earth’s ecosystems. Sci. Adv. 1, e1500052 (2015).Article
ADS
PubMed
PubMed Central
MATH
Google Scholar
Fahrig, L. Effects of habitat fragmentation on biodiversity. Annu. Rev. Ecol. Evol. Syst. 34, 487–515 (2003).Article
MATH
Google Scholar
Tscharntke, T. et al. Landscape moderation of biodiversity patterns and processes—eight hypotheses. Biol. Rev. 87, 661–685 (2012).Article
PubMed
MATH
Google Scholar
Fletcher, R. J. et al. Is habitat fragmentation good for biodiversity? Biol. Conserv. 226, 9–15 (2018).Article
MATH
Google Scholar
Simberloff, D. & Abele, L. G. Refuge design and island biogeographic theory: effects of fragmentation. Am. Nat. 120, 41–50 (1982).Article
MATH
Google Scholar
Fahrig, L. et al. Is habitat fragmentation bad for biodiversity? Biol. Conserv. 230, 179–186 (2019).Article
Google Scholar
Díaz, S. et al. Pervasive human-driven decline of life on Earth points to the need for transformative change. Science 366, eaax3100 (2019).Article
PubMed
Google Scholar
Jaureguiberry, P. et al. The direct drivers of recent global anthropogenic biodiversity loss. Sci. Adv. 8, eabm9982 (2022).Article
PubMed
PubMed Central
Google Scholar
Newbold, T. et al. Has land use pushed terrestrial biodiversity beyond the planetary boundary? A global assessment. Science 353, 288–291 (2016).Article
ADS
PubMed
MATH
Google Scholar
Betts, M. G. et al. Global forest loss disproportionately erodes biodiversity in intact landscapes. Nature 547, 441–444 (2017).Article
PubMed
MATH
Google Scholar
Johnson, C. N. et al. Biodiversity losses and conservation responses in the Anthropocene. Science 356, 270–275 (2017).Article
ADS
PubMed
MATH
Google Scholar
Watling, J. I. et al. Support for the habitat amount hypothesis from a global synthesis of species density studies. Ecol. Lett. 23, 674–681 (2020).Article
PubMed
MATH
Google Scholar
Wilson, E. O. & Willis, E. O. in Ecology and Evolution of Communities (eds Cody M. L. & Diamond J. M.) 522–534 (Harvard Univ. Press, 1975).Simberloff, D. & Abele, L. G. Island biogeography theory and conservation practice. Science 191, 285–286 (1976).Article
ADS
PubMed
Google Scholar
Ewers, R. M. & Didham, R. K. Confounding factors in the detection of species responses to habitat fragmentation. Biol. Rev. 81, 117–142 (2006).Article
PubMed
MATH
Google Scholar
Didham, R. K., Kapos, V. & Ewers, R. M. Rethinking the conceptual foundations of habitat fragmentation research. Oikos 121, 161–170 (2012).Article
ADS
MATH
Google Scholar
Lindenmayer, D. B. & Fischer, J. Tackling the habitat fragmentation panchreston. Trends Ecol. Evol. 22, 127–132 (2007).Article
PubMed
MATH
Google Scholar
Fahrig, L. Ecological responses to habitat fragmentation per se. Annu. Rev. Ecol. Evol. Syst. 48, 1–23 (2017).Article
Google Scholar
Riva, F. & Fahrig, L. Landscape‐scale habitat fragmentation is positively related to biodiversity, despite patch‐scale ecosystem decay. Ecol. Lett. 26, 268–277 (2023).Article
PubMed
MATH
Google Scholar
Martín-Queller, E., Albert, C. H., Dumas, P. J. & Saatkamp, A. Islands, mainland, and terrestrial fragments: how isolation shapes plant diversity. Ecol. Evol. 7, 6904–6917 (2017).Article
PubMed
PubMed Central
Google Scholar
de Castro Solar, R. R. et al. How pervasive is biotic homogenization in human-modified tropical forest landscapes? Ecol. Lett. 18, 1108–1118 (2015).Article
MATH
Google Scholar
Pardini, R., de Arruda Bueno, A., Gardner, T. A., Prado, P. I. & Metzger, J. P. Beyond the fragmentation threshold hypothesis: regime shifts in biodiversity across fragmented landscapes. PLoS One 5, e13666 (2010).Article
ADS
PubMed
PubMed Central
Google Scholar
Torrenta, R. & Villard, M.-A. A test of the habitat amount hypothesis as an explanation for the species richness of forest bird assemblages. J. Biogeogr. 44, 1791–1801 (2017).Article
MATH
Google Scholar
Vieira, M. V., Almeida-Gomes, M., Delciellos, A. C., Cerqueira, R. & Crouzeilles, R. Fair tests of the habitat amount hypothesis require appropriate metrics of patch isolation: an example with small mammals in the Brazilian Atlantic Forest. Biol. Conserv. 226, 264–270 (2018).Article
Google Scholar
Fletcher, R. J. Jr., Smith, T. A. H., Kortessis, N., Bruna, E. M. & Holt, R. D. Landscape experiments unlock relationships among habitat loss, fragmentation, and patch-size effects. Ecology 104, e4037 (2023).Article
PubMed
Google Scholar
Horváth, Z., Ptacnik, R., Vad, C. F. & Chase, J. M. Habitat loss over six decades accelerates regional and local biodiversity loss via changing landscape connectance. Ecol. Lett. 22, 1019–1027 (2019).Article
PubMed
PubMed Central
Google Scholar
Diekötter, T., Billeter, R. & Crist, T. O. Effects of landscape connectivity on the spatial distribution of insect diversity in agricultural mosaic landscapes. Basic Appl. Ecol. 9, 298–307 (2008).Article
Google Scholar
Fahrig, L. et al. Resolving the SLOSS dilemma for biodiversity conservation: a research agenda. Biol. Rev. 97, 99–114 (2022).Article
PubMed
Google Scholar
Morante-Filho, J. C., Arroyo-Rodríguez, V. & Faria, D. Patterns and predictors of β-diversity in the fragmented Brazilian Atlantic forest: a multiscale analysis of forest specialist and generalist birds. J. Anim. Ecol. 85, 240–250 (2016).Article
PubMed
Google Scholar
Quinn, J. F. & Harrison, S. P. Effects of habitat fragmentation and isolation on species richness: evidence from biogeographic patterns. Oecologia 75, 132–140 (1988).Article
ADS
PubMed
MATH
Google Scholar
May, F., Rosenbaum, B., Schurr, F. M. & Chase, J. M. The geometry of habitat fragmentation: effects of species distribution patterns on extinction risk due to habitat conversion. Ecol. Evol. 9, 2775–2790 (2019).Article
PubMed
PubMed Central
Google Scholar
Lôbo, D., Leão, T., Melo, F. P. L., Santos, A. M. M. & Tabarelli, M. Forest fragmentation drives Atlantic forest of northeastern Brazil to biotic homogenization. Divers. Distrib. 17, 287–296 (2011).Article
Google Scholar
Arce-Peña, N. P., Arroyo-Rodríguez, V., Avila-Cabadilla, L. D., Moreno, C. E. & Andresen, E. Homogenization of terrestrial mammals in fragmented rainforests: the loss of species turnover and its landscape drivers. Ecol. Appl. 32, e02476 (2022).Article
PubMed
Google Scholar
Tscharntke, T., Batáry, P. & Grass, I. Mixing on- and off-field measures for biodiversity conservation. Trends Ecol. Evol. 39, 726–733 (2024).Article
PubMed
Google Scholar
Fahrig, L. Why do several small patches hold more species than few large patches? Glob. Ecol. Biogeogr. 29, 615–628 (2020).Article
Google Scholar
Prugh, L. R., Hodges, K. E., Sinclair, A. R. E. & Brashares, J. S. Effect of habitat area and isolation on fragmented animal populations. Proc. Natl Acad. Sci. USA 105, 20770–20775 (2008).Article
ADS
PubMed
PubMed Central
Google Scholar
Jost, L. Partitioning diversity into independent alpha and beta components. Ecology 88, 2427–2439 (2007).Article
PubMed
MATH
Google Scholar
Jamoneau, A., Chabrerie, O., Closset-Kopp, D. & Decocq, G. Fragmentation alters beta-diversity patterns of habitat specialists within forest metacommunities. Ecography 35, 124–133 (2012).Article
ADS
Google Scholar
Dambros, C. S., Junqueira Izzo, T., Castuera de Oliveira, L., Eduardo Vicente, R. & Peres, C. A. Beta-diversity buffers fragmented landscapes against local species losses. Oikos 2024, e10401 (2024).Article
ADS
Google Scholar
Pfeifer, M. et al. Creation of forest edges has a global impact on forest vertebrates. Nature 551, 187–191 (2017).Article
ADS
PubMed
PubMed Central
MATH
Google Scholar
Phillips, H. R. P., Halley, J. M., Urbina-Cardona, J. N. & Purvis, A. The effect of fragment area on site‐level biodiversity. Ecography 41, 1220–1231 (2018).Article
ADS
Google Scholar
Nekola, J. C. & White, P. S. The distance decay of similarity in biogeography and ecology. J. Biogeogr. 26, 867–878 (1999).Article
MATH
Google Scholar
Graco-Roza, C. et al. Distance decay 2.0—a global synthesis of taxonomic and functional turnover in ecological communities. Glob. Ecol. Biogeogr. 31, 1399–1421 (2022).Article
PubMed
PubMed Central
MATH
Google Scholar
Arroyo-Rodríguez, V. et al. Plant β-diversity in fragmented rain forests: testing floristic homogenization and differentiation hypotheses. J. Ecol. 101, 1449–1458 (2013).Article
MATH
Google Scholar
Sfair, J. C., Arroyo-Rodríguez, V., Santos, B. A. & Tabarelli, M. Taxonomic and functional divergence of tree assemblages in a fragmented tropical forest. Ecol. Appl. 26, 1816–1826 (2016).Article
PubMed
Google Scholar
Simberloff, D. & Cox, J. Consequences and costs of conservation corridors. Conserv. Biol. 1, 63–71 (1987).Article
MATH
Google Scholar
Damschen, E. I. et al. Ongoing accumulation of plant diversity through habitat connectivity in an 18-year experiment. Science 365, 1478–1480 (2019).Article
ADS
PubMed
MATH
Google Scholar
Villard, M.-A. & Metzger, J. P. Beyond the fragmentation debate: a conceptual model to predict when habitat configuration really matters. J. Appl. Ecol. 51, 309–318 (2014).Article
MATH
Google Scholar
Banks-Leite, C., Ewers, R. M., Folkard-Tapp, H. & Fraser, A. Countering the effects of habitat loss, fragmentation, and degradation through habitat restoration. One Earth 3, 672–676 (2020).Article
ADS
Google Scholar
Chase, J. M., Blowes, S. A., Knight, T. M., Gerstner, K. & May, F. Ecosystem decay exacerbates biodiversity loss with habitat loss. Nature 584, 238–243 (2020).Article
ADS
PubMed
Google Scholar
Gonçalves-Souza, T. et al. LandFrag: a dataset to investigate the effects of forest loss and fragmentation on biodiversity. Glob. Ecol. Biogeogr. https://doi.org/10.1111/geb.70015 (2025).Arroyo-Rodríguez, V. & Fahrig, L. Why is a landscape perspective important in studies of primates? Am. J. Primatol. 76, 901–909 (2014).Article
PubMed
MATH
Google Scholar
Hesselbarth, M. H. K., Sciaini, M., With, K. A., Wiegand, K. & Nowosad, J. landscapemetrics: an open-source R tool to calculate landscape metrics. Ecography 42, 1648–1657 (2019).Article
ADS
Google Scholar
Chao, A. et al. Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies. Ecol. Monogr. 84, 45–67 (2014).Article
MATH
Google Scholar
Hsieh, T. C., Ma, K. H. & Chao, A. iNEXT: an R package for rarefaction and extrapolation of species diversity (Hill numbers). Methods Ecol. Evol. 7, 1451–1456 (2016).Article
MATH
Google Scholar
Marion, Z. H., Fordyce, J. A. & Fitzpatrick, B. M. Pairwise beta diversity resolves an underappreciated source of confusion in calculating species turnover. Ecology 98, 933–939 (2017).Article
PubMed
Google Scholar
Brooks, M. E. et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. R J. 9, 378 (2017).Article
MATH
Google Scholar
Hartig, F. DHARMa: residual diagnostics for hierarchical (multi-level / mixed) regression models. R package version 0.4.7 https://cran.r-project.org/web/packages/DHARMa/ (2022).Cade, B. S. Model averaging and muddled multimodel inferences. Ecology 96, 2370–2382 (2015).Article
PubMed
MATH
Google Scholar
Viechtbauer, W. Conducting meta-analyses in R with the metafor package. J. Stat. Softw. 36, 1–48 (2010).Article
MATH
Google Scholar
Gonçalves-Souza, T. & Vancine, M. Zenodo https://zenodo.org/records/14885581 (2025).Download referencesAcknowledgementsL.F.S.M was supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) by grant 307984/2022-2. M.H.V. was supported by grant 2022/01899-6 (São Paulo Research Foundation; FAPESP). J.M.C. was supported by the German Center for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig (DFG FZT 118-202548816) and an ERC Advanced Grant (MetaChange) funded by the European Union. Views and opinions expressed are those of the authors only and do not necessarily reflect those of the European Union or the European Research Council. Neither the European Union nor the granting authority can be held responsible for them. We thank N. Gotelli and G. Graves for feedback on the project.Author informationAuthors and AffiliationsInstitute for Global Change Biology, School for Environment and Sustainability, University of Michigan, Ann Arbor, MI, USAThiago Gonçalves-SouzaDepartment of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, MI, USAThiago Gonçalves-Souza & Nathan J. SandersGerman Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Leipzig, GermanyJonathan M. ChaseInstitute of Computer Science, Martin Luther University Halle-Wittenberg, Halle (Saale), GermanyJonathan M. ChaseKellogg Biological Station and Department of Integrative Biology, Michigan State University, Hickory Corners, MI, USANick M. HaddadLaboratório de Ecologia Espacial e Conservação Departamento de Biodiversidade, Instituto de Biociências, Universidade Estadual Paulista (Unesp), Rio Claro, BrazilMaurício H. VancineSchool of Biological Sciences, University of Western Australia, Perth, Western Australia, AustraliaRaphael K. Didham & Marcelo TabarelliCSIRO Health and Biosecurity, Centre for Environment and Life Sciences, Floreat, Western Australia, AustraliaRaphael K. DidhamCentro de Biociências, Universidade Federal de Pernambuco, Recife, BrazilFelipe L. P. MeloSchool of Animal, Rural and Environmental Sciences, Nottingham Trent University, Nottingham, UKFelipe L. P. MeloGrupo de Ecología de la Polinización (EcoPol), INIBIOMA (CONICET, Universidad Nacional del Comahue), San Carlos de Bariloche, ArgentinaMarcelo A. AizenLaboratório de Ciência Aplicada à Conservação da Biodiversidade, Departamento de Zoologia, Universidade Federal de Pernambuco, Recife, BrazilEnrico BernardLaboratório de Ecologia e Conservação, Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Ribeirão Preto, BrazilAdriano G. ChiarelloApplied Ecology & Conservation Lab, Universidade Estadual de Santa Cruz, Ilhéus, BrazilDeborah Faria & Larissa Rocha-SantosDepartment of Environment and Genetics, La Trobe University, Melbourne, Victoria, AustraliaHeloise GibbCenter for Large Landscape Conservation, Bozeman, MT, USAMarcelo G. de LimaIUCN WCPA Connectivity Conservation Specialist Group (CCPG), Cambridge, UKMarcelo G. de LimaCentro de Formação em Ciências Agroflorestais, Universidade Federal do Sul da Bahia, Ilhéus, BrazilLuiz F. S. Magnago & Nathalia Vieira Hissa SafarInstituto de Biologia, Universidade Federal da Bahia, Salvador, BrazilEduardo Mariano-NetoIndependent researcher, São Paulo, BrazilAndré A. NogueiraInstituto de Biologia, Universidade Federal de Uberlândia (UFU), Uberlândia, BrazilAndré Nemésio & Heraldo L. VasconcelosLaboratório de Ecologia e Conservação de Mamíferos, Departamento de Ecologia e Conservação, Instituto de Ciências Naturais, Universidade Federal de Lavras, Lavras, BrazilMarcelo PassamaniInstitute of Plant Sciences, University of Bern, Bern, SwitzerlandBruno X. PinhoCentro Nacional de Pesquisa e Conservação de Aves Silvestres (CEMAVE), Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio), Cabedelo, BrazilRodolpho C. RodriguesDepartamento de Sistemática e Ecologia, Universidade Federal da Paraíba, João Pessoa, BrazilBráulio A. SantosCuerpo Académico de Ecología y Diversidad Faunística. Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Querétaro, MéxicoAlejandra Soto-WerschitzIndependent researcher, Brasília, BrazilMarcio Uehara-PradoNúcleo de Estudos e Pesquisas Ambientais, Universidade Estadual de Campinas, Campinas, BrazilSimone VieiraAuthorsThiago Gonçalves-SouzaView author publicationsYou can also search for this author in
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PubMed Google ScholarContributionsT.G.-S., J.M.C., N.M.H. and N.J.S. conceived and designed the study. T.G.-S. performed the analyses with support from J.M.C., N.M.H., N.J.S., M.H.V. and F.L.P.M. T.G.-S., J.M.C., N.M.H. and N.J.S. wrote the first version of the paper. F.L.P.M., M.A.A., E.B., A.G.C., R.K.D., D.F., H.G., M.G.d.L., L.F.S.M., E.M.-N., A.A.N., A.N., M.P., B.X.P., L.R.-S., R.C.R., N.V.H.S., B.A.S., A.S.-W., M.T., M.U.-P., H.L.V. and S.V. collected data. J.M.C., T.G.-S. and M.H.V. curated and maintained the data. All authors contributed to the revisions of the manuscript.Corresponding authorCorrespondence to
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Additional informationPublisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.Extended data figures and tablesExtended Data Fig. 1 Study locations.A global map displaying the locations of the 37 studies, as well as the taxonomic groups sampled in each. Made with Natural Earth. Free vector and raster map data from naturalearthdata.com.Extended Data Fig. 2 Fragment size classes.Number (and percentage) of fragments or continuous forests in each fragment size class, which included fragments smaller than 100, 500 and 1,000 ha, as well as forest larger than 1,000 ha. The values were calculated using all 121 studies from the LandFrag dataset51.Extended Data Fig. 3 Habitat amounts in continuous and fragmented landscapes across all buffer sizes.The buffer size ranged from a radius of 200 m to 2,000 m, in 200 m increments. Large circles represent the mean habitat amount in continuous (blue circles) and fragmented (red circles) landscapes across studies (n = 37). In all panels, each study is represented by a small grey circle, with lines connecting the landscape types, and error bars represent standard deviations.Extended Data Fig. 4 Number of patches in continuous and fragmented landscapes across all buffer sizes.The buffer size ranged from a radius of 200 m to 2,000 m, in 200 m increments. Large circles represent the mean habitat amount in continuous (blue circles) and fragmented (red circles) landscapes across studies (n = 37). In all panels, each study is represented by a small grey circle, with lines connecting the landscape types, and error bars represent standard deviations.Extended Data Fig. 5 Variation in landscape variables related to fragmentation between continuous and fragmented landscapes across all studies.Dots represent the average value (error bars ± 1 s.e.) of the landscape variable (buffer radius = 2,000 m) in a given habitat amount class (n = 37 studies).Extended Data Fig. 6 Differences in α, β and γ diversity between fragmented and continuous landscapes using a meta-analytical approach.This orchard plot shows the effect size (log-response ratio, LRR) of the overall difference between continuous and fragmented landscapes across studies (n = 37). The values of α, β and γ diversity were computed using (a) all possible plot pairs or (b) only the nearest plot pairs (controlling for distance decay effects) in both continuous and fragmented landscapes. We also calculated α, β and γ diversity using the observed species richness without controlling for commonness or sampling effort, and with individual-based rarefaction giving greater relative weight to rare species (rarefied species richness; q = 0) and individual-based rarefaction giving greater relative weight to abundant species (effective number of species given Simpson diversity; q = 2). Solid points represent the LRR comparing α, β and γ diversity between continuous and fragmented landscapes, and the error bars represent 95% confidence intervals. Positive effect sizes indicate that continuous landscapes have higher diversity than fragmented landscapes, while negative effect sizes would indicate that fragmented landscapes have higher diversity. Transparent points indicate effect sizes from individual sites, and their sizes are proportional to the precision (inverse of the square root of the variance) of the individual effect size estimates.Extended Data Fig. 7 Individual contribution of landscape type and habitat amount to α, β and γ diversity.As described in the main text, this analysis represents the scaled importance of predictor variables in GLMMs. The most important variable (that is, the one with the highest unbiased AICc weight value) in a given model receives a value of 1, and the relative contribution of the other variables is calculated based on this benchmark. We performed these analyses using all species, as well as using rarefaction analyses to give weight to rare (order q = 0) and abundant (order q = 2) species.Extended Data Fig. 8 Illustration of the analytical pipeline used to standardize species diversity comparisons between continuous and fragmented landscapes.a, The grey squares represent one large forest in a continuous landscape and four small fragments in a fragmented landscape. The small black squares represent a sample and illustrate how differences in size generally affect the number of samples when comparing landscape types. Furthermore, when comparing these landscape types, there are at least four analytical challenges (1–4) that affect our ability to estimate and compare α, β, and γ diversity. b, Analytical approach used to estimate diversity by (1) standardizing α diversity while accounting for differences in study design and sampling effort, (2) standardizing β and γ diversity by calculating pairwise sample diversity, (3) controlling for distance decay effects to accurately estimate α, β, and γ diversity, and (4) standardizing α, β, and γ diversity through consistent sampling effort adjustments across landscapes. Silhouettes from PhyloPic (http://phylopic.org/), as a courtesy of Andy Wilson, Birgit Lang, Lauren Sumner-Rooney, Mattia Menchetti, Dorota Paczesniak, Birgit Lang, Wouter Koch, Guillaume Dera, Graham Montgomery and Gareth Monger.Extended Data Fig. 9 Method to calculate pairwise diversity for α, β and γ using a patch-landscape study design.a, The average α diversity for each pair is calculated as the mean number of species in pair i and pair ii, while γ diversity is the pooled ‘total’ number of distinct species in a given pair (accounting for the overlap of shared species occurring in both pairs). This pair is selected using two approaches. b, All possible pairs in fragmented or continuous landscapes in each study. c, Only the nearest pairs to control for the effect of distance decay on β and γ diversity. The black squares represent a plot, numbered between 1 and 10. Therefore, a pair can consist of either two plots within the same forest or fragment, or two plots in different fragments.Extended Data Fig. 10 Estimated mixed-effects meta-analysis coefficients using the ‘leave-one-out’ analysis to compare the effects of removing one study from South America versus other continents.Error bars represent 95% confidence intervals, shaded areas represent standard deviations, and points indicate the average coefficient by continent groups (South America in purple vs. other continents in green). The vertical dashed lines represent the observed coefficient value for the main model (n = 37 studies). We estimated α, β and γ diversity using all species, as well as giving greater relative weight for rare (q = 0) or abundant (q = 2) species in two scenarios: all plot pairs and the nearest plot pairs.Supplementary informationSupplementary InformationSupplementary Text 1–5, Supplementary Tables 1–6 and 8–13 and Supplementary ReferencesReporting SummaryPeer Review fileSupplementary Table 7Rights and permissionsSpringer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.Reprints and permissionsAbout this articleCite this articleGonçalves-Souza, T., Chase, J.M., Haddad, N.M. et al. Species turnover does not rescue biodiversity in fragmented landscapes.
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