Abstract
Species richness and ecological uniqueness (defined as local contributions of sites to beta diversity of a region) constitute two crucial components of biodiversity. Understanding the relationship between these two components is critical for effective conservation planning, yet global patterns and underlying drivers remain largely controversial. Here, we conduct a comprehensive global meta-analysis to investigate the patterns of the species richness-ecological uniqueness relationship across taxa groups and to evaluate the relative importance of four ecological hypotheses in driving this relationship (i.e. species pool, dispersal limitation, environmental filtering and spatial grain size). We find that negative richness-uniqueness relationships are prevalent across different taxa groups (e.g. terrestrial plants, freshwater macroinvertebrates, birds, and fishes), and such negative association is robust to different data types including presence absence and abundance data. Notably, relationships based on presence-absence data are primarily predicted by species pool attributes (i.e. the proportion of rare species and gamma diversity), whereas abundance-based relationships are more strongly associated with dispersal limitation. In summary, these findings reveal consistent global patterns and mechanistic underpinnings of the richness-uniqueness relationship, offering key insights for biodiversity conservation. We recommend that conservation strategies should prioritize both species-rich and ecologically unique communities to maximize biodiversity protection globally.
Data availability
The Supporting data and three appendices (Appendix 1, Appendix 2 and Appendix 3) that support the findings of this study are available in Figshare at https://doi.org/10.6084/m9.figshare.2927004894. The reference list of included and excluded studies is also provided as Supplementary Data 1. Source data are provided in this paper.
Code availability
R code that supports the findings of this study is available in Figshare at https://doi.org/10.6084/m9.figshare.2927004894.
References
Chesson, P. Mechanisms of Maintenance of Species Diversity. Annu. Rev. Ecol. Evol. Syst 31, 343–366 (2000).
Socolar, J. B., Gilroy, J. J., Kunin, W. E. & Edwards, D. P. How Should Beta-Diversity Inform Biodiversity Conservation?. Trends Ecol. Evol. 31, 67–80 (2016).
Kier, G. et al. A global assessment of endemism and species richness across island and mainland regions. Proc. Natl. Acad. Sci. USA 106, 9322–9327 (2009).
Myers, N., Mittermeier, R. A., Mittermeier, C. G., da Fonseca, G. A. B. & Kent, J. Biodiversity hotspots for conservation priorities. Nature 403, 853–858 (2000).
Legendre, P. & De Cáceres, M. Beta diversity as the variance of community data: dissimilarity coefficients and partitioning. Ecol. Lett. 16, 951–963 (2013).
Harper, L. M. et al. Blinded by the bright: How species-poor habitats contribute to regional biodiversity across a tropical seascape. Divers. Distrib. 28, 2272–2285 (2022).
Vilmi, A., Karjalainen, S. M. & Heino, J. Ecological uniqueness of stream and lake diatom communities shows different macroecological patterns. Divers. Distrib. 23, 1042–1053 (2017).
Violle, C. et al. Functional rarity: The ecology of outliers. Trends Ecol. Evol. 32, 356–367 (2017).
Casanelles-Abella, J., Fontana, S., Meier, E., Moretti, M. & Fournier, B. Spatial mismatch between wild bee diversity hotspots and protected areas. Conserv. Biol. 37, e14082 (2023).
Hill, M. J. et al. Local contributions to beta diversity in urban pond networks: Implications for biodiversity conservation and management. Divers. Distrib. 27, 887–900 (2021).
Niskanen, A. K. J., Heikkinen, R. K., Vare, H. & Luoto, M. Drivers of high-latitude plant diversity hotspots and their congruence. Biol. Conserv. 212, 288–299 (2017).
da Silva, P. G., Medina Hernandez, M. I. & Heino, J. Disentangling the correlates of species and site contributions to beta diversity in dung beetle assemblages. Divers. Distrib. 24, 1674–1686 (2018).
Yao, J. et al. Ecological uniqueness of species assemblages and their determinants in forest communities. Divers. Distrib. 27, 454–462 (2021).
Dansereau, G., Legendre, P. & Poisot, T. Evaluating ecological uniqueness over broad spatial extents using species distribution modelling. Oikos 2022, e09063 (2022).
Heino, J. & Gronroos, M. Exploring species and site contributions to beta diversity in stream insect assemblages. Oecologia 183, 151–160 (2017).
Chen, Y. et al. Multiple processes jointly determine ecological uniqueness across forest plant life-forms in Northeast China. J. Biogeogr. 51, 1133–1147 (2024).
Landeiro, V. L., Franz, B., Heino, J., Siqueira, T. & Bini, L. M. Species-poor and low-lying sites are more ecologically unique in a hyperdiverse Amazon region: Evidence from multiple taxonomic groups. Divers. Distrib. 24, 966–977 (2018).
Kong, H., Chevalier, M., Laffaille, P. & Lek, S. Spatio-temporal variation of fish taxonomic composition in a South-East Asian flood-pulse system. Plos ONE 12, e0174582 (2017).
Legendre, P. Interpreting the replacement and richness difference components of beta diversity. Glob. Ecol. Biogeogr. 23, 1324–1334 (2014).
Li, F., Tonkin, J. D. & Haase, P. Local contribution to beta diversity is negatively linked with community-wide dispersal capacity in stream invertebrate communities. Ecol. Indic. 108, 105715 (2019).
van der Plas, F., Hennecke, J., Chase, J. M., van Ruijven, J. & Barry, K. E. Universal beta-diversity–functioning relationships are neither observed nor expected. Trends Ecol. Evol. 38, 532–544 (2023).
Wang, X. et al. Spatial patterns of tree species richness in two temperate forests. J. Ecol. 99, 1382–1393 (2011).
Guo, Q., Qian, H. & Zhang, J. On the relationship between species diversity and range size. J. Biogeogr. 49, 1911–1919 (2022).
Jimenez-Alfaro, B. et al. History and environment shape species pools and community diversity in European beech forests. Nat. Ecol. Evol. 2, 483–490 (2018).
Luukkonen, S., Heino, J., Hjort, J., Helm, A. & Alahuhta, J. A macroecological analysis of ecological uniqueness of freshwater macrophyte assemblages across Europe and North America. J. Ecol. 112, 2944–2958 (2024).
Wang, S. et al. Differential responses of forest strata species richness to paleoclimate and forest structure. For. Ecol. Manag. 499, 119605 (2021).
Barton, P. S. et al. The spatial scaling of beta diversity. Glob. Ecol. Biogeogr. 22, 639–647 (2013).
Stein, A., Gerstner, K. & Kreft, H. Environmental heterogeneity as a universal driver of species richness across taxa, biomes and spatial scales. Ecol. Lett. 17, 866–880 (2014).
Anderson, M. J. et al. Navigating the multiple meanings of β diversity: a roadmap for the practicing ecologist. Ecol. Lett. 14, 19–28 (2011).
Chase, J. M. & Knight, T. M. Scale-dependent effect sizes of ecological drivers on biodiversity: why standardised sampling is not enough. Ecol. Lett. 16, 17–26 (2013).
Cai, L. et al. Environmental filtering, not dispersal history, explains global patterns of phylogenetic turnover in seed plants at deep evolutionary timescales. Nat. Ecol. Evol. 9, 314–324 (2025).
McGill, B. J. et al. Species abundance distributions: moving beyond single prediction theories to integration within an ecological framework. Ecol. Lett. 10, 995–1015 (2007).
Cornell, H. V. & Harrison, S. P. What Are Species Pools and When Are They Important?. Annu. Rev. Ecol. Evol. Syst. 45, 45–67 (2014).
Lessard, J.-P., Belmaker, J., Myers, J. A., Chase, J. M. & Rahbek, C. Inferring local ecological processes amid species pool influences. Trends Ecol. Evol. 27, 600–607 (2012).
Rabinowitz, D. Seven forms of rarity. in The biological aspects of rare plant conservation (ed. Synge, H.) 205–217 (1981).
Crisfield, V. E., Guillaume Blanchet, F., Raudsepp-Hearne, C. & Gravel, D. How and why species are rare: towards an understanding of the ecological causes of rarity. Ecography 2024, e07037 (2024).
Engel, T. et al. How does variation in total and relative abundance contribute to gradients of species diversity?. Ecol. Evol. 12, e9196 (2022).
Heino, J. The importance of metacommunity ecology for environmental assessment research in the freshwater realm. Biol. Rev. Camb. Philos. Soc 88, 166–178 (2013).
Brown, B. L. & Swan, C. M. Dendritic network structure constrains metacommunity properties in riverine ecosystems. J. Anim. Ecol. 79, 571–580 (2010).
Condit, R. et al. Spatial patterns in the distribution of tropical tree species. Science 288, 1414–1418 (2000).
Wang, X. et al. Spatial distributions of species in an old-growth temperate forest, northeastern China. Can. J. For. Res. 40, 1011–1019 (2010).
Rosenzweig, M. L. Species Diversity in Space and Time (Cambridge Univ. Press, Cambridge, 1995).
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).
Graham, L. J., Spake, R., Gillings, S., Watts, K. & Eigenbrod, F. Incorporating fine-scale environmental heterogeneity into broad-extent models. Methods Ecol. Evol. 10, 767–778 (2019).
Baselga, A. Separating the two components of abundance-based dissimilarity: balanced changes in abundance vs. abundance gradients. Methods Ecol. Evol. 4, 552–557 (2013).
Leibold, M. A. et al. The metacommunity concept: a framework for multi-scale community ecology. Ecol. Lett. 7, 601–613 (2004).
Murphy, S. J., Salpeter, K. & Comita, L. S. Higher beta-diversity observed for herbs over woody plants is driven by stronger habitat filtering in a tropical understory. Ecology 97, 2074–2084 (2016).
Chu, C. et al. Direct and indirect effects of climate on richness drive the latitudinal diversity gradient in forest trees. Ecol. Lett. 22, 245–255 (2019).
Rocha, M. P., Morris, T. J., Cottenie, K. & Schwalb, A. N. Limitations of beta diversity in conservation site selection. Ecol. Indic. 154, 110732 (2023).
Moi, D. A. et al. Non-native fishes homogenize native fish communities and reduce ecosystem multifunctionality in tropical lakes over 16 years. Sci. Total Environ. 769, 144524 (2021).
Shen, M. et al. FreshLanDiv: A global database of freshwater biodiversity across different land uses. Glob. Ecol. Biogeogr. 33, e13917 (2024).
Green, A. J., Baltzinger, C. & Lovas-Kiss, Á Plant dispersal syndromes are unreliable, especially for predicting zoochory and long-distance dispersal. Oikos 2022, e08327 (2022).
Alzate, A. & Onstein, R. E. Understanding the relationship between dispersal and range size. Ecol. Lett. 25, 2303–2323 (2022).
Violle, C. et al. Let the concept of trait be functional!. Oikos 116, 882–892 (2007).
Heino, J. et al. Assessing the conservation priority of freshwater lake sites based on taxonomic, functional and environmental uniqueness. Divers. Distrib. 28, 1966–1978 (2022).
Ascanio, A., Bracken, J. T., Stevens, M. H. H. & Jezkova, T. New theoretical and analytical framework for quantifying and classifying ecological niche differentiation. Ecol. Monogr. 94, e1622 (2024).
Page, M. J. et al. The PRISMA 2020 statement: an updated guideline for reporting systematic reviews. BMJ 372, n71 (2021).
Nakagawa, S. & Cuthill, I. C. Effect size, confidence interval and statistical significance: a practical guide for biologists. Biol. Rev. Camb. Philos. Soc. 82, 591–605 (2007).
Plotkin, J. B. & Muller-Landau, H. C. Sampling the species composition of a landscape. Ecology 83, 3344–3354 (2002).
Biurrun, I. et al. Benchmarking plant diversity of Palaearctic grasslands and other open habitats. J. Veg. Sci. 32, e13050 (2021).
Dengler, J. et al. GrassPlot – A database of multi-scale plant diversity in Palaearctic grasslands. Phytocoenologia 48, 331–347 (2018).
Bruelheide, H. et al. sPlot – A new tool for global vegetation analyses. J. Veg. Sci. 30, 161–186 (2019).
Jeliazkov, A. et al. A global database for metacommunity ecology, integrating species, traits, environment and space. Sci. Data 7, 6 (2020).
Burnham, K. P. & Anderson, D. R. Multimodel inference: Understanding AIC and BIC in model selection. Sociol. Methods Res. 33, 261–304 (2004).
Muruga, P., Siqueira, A. C. & Bellwood, D. R. Meta-analysis reveals weak associations between reef fishes and corals. Nat. Ecol. Evol. 8, 676–685 (2024).
Chang, L. W., Zeleny, D., Li, C. F., Chiu, S. T. & Hsieh, C. F. Better environmental data may reverse conclusions about niche- and dispersal-based processes in community assembly. Ecology 94, 2145–2151 (2013).
Kobayashi, T. & Sota, T. Distance decay of similarity in fungivorous insect communities: assessing dispersal limitation using genetic data. Ecosphere 7, e01358 (2016).
Marrec, R., Pontbriand-Paré, O., Legault, S. & James, P. M. A. Spatiotemporal variation in drivers of parasitoid metacommunity structure in continuous forest landscapes. Ecosphere 9, e02075 (2018).
Punchi-Manage, R. et al. Effect of spatial processes and topography on structuring species assemblages in a Sri Lankan dipterocarp forest. Ecology 95, 376–386 (2014).
Borcard, D., Legendre, P., Avois-Jacquet, C. & Tuomisto, H. Dissecting the spatial structure of ecological data at multiple scales. Ecology 85, 1826–1832 (2004).
Legendre, P. & Legendre, L. Numerical Ecology. (Elsevier, Amsterdam, 2012).
Legendre, P. et al. Partitioning beta diversity in a subtropical broad-leaved forest of China. Ecology 90, 663–674 (2009).
Cribari-Neto, F. & Zeileis, A. Beta Regression in R. J. Stat. Softw. 34, 1–24 (2010).
Myers, J. A. et al. Beta-diversity in temperate and tropical forests reflects dissimilar mechanisms of community assembly. Ecol. Lett. 16, 151–157 (2013).
Hijmans, R. J., Cameron, S. E., Parra, J. L., Jones, P. G. & Jarvis, A. Very high resolution interpolated climate surfaces for global land areas. Int. J. Climatol. 25, 1965–1978 (2005).
Hutchinson, G. E. Homage to Santa Rosalia or why are there so many kinds of animals?. Am. Nat. 93, 145–159 (1959).
Currie, D. J. et al. Predictions and tests of climate-based hypotheses of broad-scale variation in taxonomic richness. Ecol. Lett. 7, 1121–1134 (2004).
Chu, C. et al. Direct effects dominate responses to climate perturbations in grassland plant communities. Nat. Commun. 7, 11766 (2016).
Storch, D., Bohdalková, E. & Okie, J. The more-individuals hypothesis revisited: the role of community abundance in species richness regulation and the productivity–diversity relationship. Ecol. Lett. 21, 920–937 (2018).
Higgins, J., Thompson, S. G., Deeks, J. J. & Altman, D. G. Measuring inconsistency in meta-analyses. BMJ 327, 557–560 (2003).
Senior, A. M. et al. Heterogeneity in ecological and evolutionary meta-analyses: its magnitudes and implications. Ecology 97, 3293–3299 (2016).
Gotelli, N. J. Null model analysis of species co-occurrence patterns. Ecology 81, 2606–2621 (2000).
Kraft, N. J. B. et al. Disentangling the drivers of β diversity along latitudinal and elevational gradients. Science 333, 1755–1758 (2011).
Xing, D. L. & He, F. L. Environmental filtering explains a U-shape latitudinal pattern in regional β-deviation for eastern North American trees. Ecol. Lett. 22, 284–291 (2019).
Gelman, A. & Hill, J. Data Analysis Using Regression and Multilevel/Hierarchical Models (Cambridge Univ. Press, Cambridge, 2006).
García-Palacios, P., Gross, N., Gaitán, J. & Maestre, F. T. Climate mediates the biodiversity–ecosystem stability relationship globally. Proc. Natl. Acad. Sci. USA 115, 8400–8405 (2018).
Nakagawa, S. & Schielzeth, H. A general and simple method for obtaining R² from generalized linear mixed-effects models. Methods Ecol. Evol. 4, 133–142 (2013).
Viechtbauer, W. Conducting Meta-Analyses in R with the metafor Package. J. Stat. Softw. 36, 1–48 (2010).
Egger, M., Smith, G. D., Schneider, M. & Minder, C. Bias in meta-analysis detected by a simple, graphical test. BMJ 315, 629–634 (1997).
Rosenthal, R. The file drawer problem and tolerance for null results. Psychol. Bull. 86, 638–641 (1979).
Duval, S. J. & Tweedie, R. L. Trim and fill: A simple funnel-plot-based method of testing and adjusting for publication bias in meta-analysis. Biometrics 56, 455–463 (2000).
R Core Team. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria <https://www.rproject.org/> (2019).
Nakagawa, S. et al. orchaRd 2.0: An R package for visualizing meta-analyses with orchard plots. Methods Ecol. Evol. 14, 2003–2010 (2023).
Chen, Y. et al. Meta-analysis reveals widespread negative associations between species richness and ecological uniqueness. figshare. https://doi.org/10.6084/m9.figshare.29270048 (2026).
Acknowledgements
This work was supported by the National Natural Science Foundation of China (32525041), the CAS (Chinese Academy of Sciences) Project for Young Scientists in Basic Research (YSBR-108), and Liaoning Revitalization Talents Program (XLYC2402003). J.A.M. was supported by grants from the U.S. National Science Foundation (DEB 2240431) and the Seeding Projects for Enabling Excellence and Distinction (SPEED) program at Washington University in St. Louis.
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Conceptualization: X.W. and Y.C. Methodology (extraction): Y.C. Methodology (statistical analysis): Y.C. and J.S. Visualization: Y.C. Writing—original draft: Y.C. and X.W. Writing—review, and editing: J.S., J.A.M., Z.M., J.Y., and X.W.
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Chen, Y., Soininen, J., Myers, J.A. et al. Meta-analysis reveals widespread negative associations between species richness and ecological uniqueness.
Nat Commun (2026). https://doi.org/10.1038/s41467-026-70886-2
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DOI: https://doi.org/10.1038/s41467-026-70886-2
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