Abstract
Land-use intensification is a major driver of biodiversity loss1,2. Alongside reductions in local species diversity, biotic homogenization at larger spatial scales is of great concern for conservation. Biotic homogenization means a decrease in β-diversity (the compositional dissimilarity between sites). Most studies have investigated losses in local (α)-diversity1,3 and neglected biodiversity loss at larger spatial scales. Studies addressing β-diversity have focused on single or a few organism groups (for example, ref. 4), and it is thus unknown whether land-use intensification homogenizes communities at different trophic levels, above- and belowground. Here we show that even moderate increases in local land-use intensity (LUI) cause biotic homogenization across microbial, plant and animal groups, both above- and belowground, and that this is largely independent of changes in α-diversity. We analysed a unique grassland biodiversity dataset, with abundances of more than 4,000 species belonging to 12 trophic groups. LUI, and, in particular, high mowing intensity, had consistent effects on β-diversity across groups, causing a homogenization of soil microbial, fungal pathogen, plant and arthropod communities. These effects were nonlinear and the strongest declines in β-diversity occurred in the transition from extensively managed to intermediate intensity grassland. LUI tended to reduce local α-diversity in aboveground groups, whereas the α-diversity increased in belowground groups. Correlations between the β-diversity of different groups, particularly between plants and their consumers, became weaker at high LUI. This suggests a loss of specialist species and is further evidence for biotic homogenization. The consistently negative effects of LUI on landscape-scale biodiversity underscore the high value of extensively managed grasslands for conserving multitrophic biodiversity and ecosystem service provision. Indeed, biotic homogenization rather than local diversity loss could prove to be the most substantial consequence of land-use intensification.
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Acknowledgements
We are grateful to J. Chase and M. Fitzpatrick for their comments and suggestions on a previous version of the manuscript; B. Büche, R. Achtziger, T. Wagner, F. Köhler, T. Blick and M.-A. Fritze for arthropod species identification and U. Kern for creating the small icons of the 12 trophic groups used in the figures. We thank the managers of the three Exploratories, K. Hartwich, S. Gockel, K. Wiesner and M. Gorke for their work in maintaining the plot and project infrastructure; C. Fischer and S. Pfeiffer for giving support through the central office, M. Owonibi for managing the central data base; and E. Linsenmair, D. Hessenmöller, J. Nieschulze, I. Schöning and the late E. Kalko for their role in setting up the Biodiversity Exploratories project. We are also grateful to E. Kalko for her invaluable inspiration and for launching the studies on bats and birds. The work has been funded by the DFG Priority Program 1374 ‘Infrastructure-Biodiversity-Exploratories’. Field work permits were issued by the responsible state environmental offices of Baden-Württemberg, Thüringen and Brandenburg (according to §72 BbgNatSchG).
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M.M.G. and E.A. conceived the idea for the manuscript, and defined the final analysis, M.M.G., E.A., C.P., T.M.L. and T.K. analysed the data, M.M.G. and E.A. wrote the first manuscript draft and finalized the manuscript. A.M.K., C.B., C.N.W., C.W., D.J.P., D.P., E.P., F.B., H.A., I.S., J.K., J.M., J.S., J.O., K.J., K.B., M.Tü., M.Ts., M.F., M.L., M.M.G., M.W., N.B., P.C.V., S.Bl., S.Bo., S.A.S., S.C.R, S.K., S.W., T.D., T.W., V.B., V.W., and W.W.W. contributed data. T.M.L., F.G., S.Bo., D.P., L.R.J., K.B., S.C.R., A.C.K., O.P., P.S., T.T., W.W.W. and J.S. contributed substantially to revisions. All authors commented on the manuscript.
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Extended data figures and tables
Extended Data Figure 1 The effect of LUI on higher q-level α-diversity above- and belowground.
The partial effect of local LUI comes from a power law model fitted to the exponential Shannon diversity (q = 1) and reciprocal Simpson index (q = 2) of the seven aboveground (solid lines) and the five belowground trophic groups (dashed lines) (n = 105 plots; for more details see Methods). In the model, all parameters of the power law function depended on temporal variation in LUI (sdLUI) and isolation. LUI effects are plotted at the mean values of these two variables. α-diversity and land-use variables were corrected for differences due to region, pH and soil nutrients, by taking residuals, and were then scaled between 0 and 1. The models for protists (q = 1 and q = 2) and mycorrhizae (q = 2) failed to converge and are therefore not shown. Note that plant pathogens are missing because, for this group, no data on abundance was available.
Extended Data Figure 2 Effects of LUI on turnover of aboveground species.
Scatter plots showing the effects of mean LUI and ΔLUI, between plot pairs (n = 105 plots), on the species turnover component of β-diversity for seven aboveground groups. Regression lines show predictions from linear models.
Extended Data Figure 3 Effects of LUI on turnover of belowground species.
Scatter plots showing the effects of mean LUI and ΔLUI, between plot pairs (n = 105 plots), on the species turnover component of β-diversity for five belowground groups. Regression lines show predictions from linear models.
Extended Data Figure 4 Effects of LUI on total β-diversity above- and belowground.
a, c, e, Partial effects of mean LUI and ΔLUI, between plot pairs, on total β-diversity (a, Sørensen q = 0; c, Morisita q = 1; e, Morisita–Horn q = 2) for seven aboveground and five belowground groups from linear models. Negative effects of mean LUI indicate that land-use intensification reduces β-diversity. The bars show coefficients from the models. Numbers adjoining bars are the proportion of explained variance uniquely explained by mean LUI or ΔLUI. b, d, f, Results from the GDMs are shown for total β-diversity (b, Sørensen q = 0; d, Morisita q = 1; f, Morisita–Horn q = 2) for the same trophic groups. The figures show the effect of differences in LUI on β-diversity (calculated between all plot pairs). Effects of differences in LUI can vary nonlinearly along the gradient of LUI. Higher maximum curves indicate larger effects of differences in LUI on β-diversity. The values in the legend are the percentage of deviance that is explained uniquely by LUI. Effects of both linear models and GDMs are corrected for other drivers of β-diversity, and response and explanatory variables are scaled to allow comparisons across trophic groups (n = 105 plots; for details see Methods).
Extended Data Figure 5 Partial effects of geographic and environmental distances and temporal variation in LUI on β-diversity above- and belowground.
a, b, Results from GDMs are shown for seven aboveground and five belowground groups, with total β-diversity measured as the Sørensen index βsor (a) or as the species turnover component βsim (b). The figures show the effect of differences in each of the five variables on β-diversity (calculated between all plot pairs; n = 105 plots). Effects of differences in each explanatory variable can vary nonlinearly along the gradient of that variable and each is corrected for all other variables in the model. Higher maximum curves indicate larger effects of differences in a given variable on β-diversity. Soil nutrients refer to the scores of the first PCA axis. Temporal variation in LUI is shown as s.d. Geographic distance has to be multiplied by 100 km.
Extended Data Figure 6 Uncertainty of effects of LUI on β-diversity above (AG) and belowground (BG).
The uncertainty is calculated on the basis of 100 bootstraps for each model, each time removing 30% of the plot pairs, then fitting a GDM and extracting the predictions. Predictions are shown as fitted lines and s.d. Uncertainty is shown for all seven above- and five belowground trophic groups based on species turnover βsim (n = 105 plots). PriPro, primary producers; PlPa, plant pathogens; Herb, herbivores; Poll, pollinators; InvDec, invertebrate decomposers; SecCon, secondary consumers; TerCon, tertiary consumers; Myco, Mycorrhizae; MicDec, microbial decomposers; Bact, bacterivores.
Extended Data Figure 7 The relative importance of LUI as a driver of β-diversity.
The bar plot shows the importance of LUI (in terms of total effect size) relative to the most important variable in the GDM. Results are shown for each trophic group, for the species turnover component (βsim) and total β-diversity (Sørensen index) (n = 105 plots).
Extended Data Figure 8 Effects of single land-use components on β-diversity above- and belowground.
a, c, e, Partial effects of minimum LUI (min LUI) and ΔLUI between plot pairs (n = 105 plots), on the species turnover component of β-diversity (βsim) for seven aboveground and five belowground groups based on linear models. Negative effects of minimum LUI indicate that land-use intensification reduces β-diversity. The bars show coefficients from the models. Numbers adjoining bars are the proportion of the total explained variance that is uniquely explained by minimum LUI orΔLUI among plot pairs, on the basis of hierarchical partitioning. b, d, f, Results from GDMs are shown for the turnover component βsim for the same trophic groups. The figures show the effect of ΔLUI on β-diversity (calculated between all plot pairs). Effects of ΔLUI can vary nonlinearly along the gradient of LUI. Higher maximum curves indicate larger effects of ΔLUI on β-diversity. The values in the legend are the percentage of deviance that is explained uniquely by LUI. Effects of both linear models and GDMs are corrected for other drivers of β-diversity, and response and explanatory variables are scaled to allow comparisons across trophic levels (see Methods).
Extended Data Figure 9 Sample coverage of above- and belowground trophic groups based on species incidences.
Sample coverage was calculated for low (52 plots) and high (53) LUI plots based on refs 57, 58. Coverage is defined as the proportion of the total number of individuals in an assemblage that belong to species represented in the sample, and is therefore a measure of sampling completeness. Means and 95% confidence intervals based on 200 bootstraps are shown.
Extended Data Figure 10 The effect of LUI on the correlation between the β-diversities of different trophic groups.
Each dot represents the correlation (R2) between two trophic groups. Correlations are R2 values from matrix regressions between β-diversity values of different groups (corrected for effects of differences in LUI on β-diversity). Significant correlations (P < 0.05) are marked in red. Upward and downward triangles indicate significance under low or high LUI only. Interactions with R2 values higher than 0.2 in one of the LUI-categories are illustrated by icons. β-diversity was calculated as the Sørensen index (βsor, total β-diversity) and as the species turnover component (βsim) (n = 105 plots). For statistical details see Supplementary Information Section 5.
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Gossner, M., Lewinsohn, T., Kahl, T. et al. Land-use intensification causes multitrophic homogenization of grassland communities. Nature 540, 266–269 (2016). https://doi.org/10.1038/nature20575
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DOI: https://doi.org/10.1038/nature20575
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