R. D. Bardgett and W. H. Van-der-putten, Belowground biodiversity and ecosystem functioning. Nat. 515. 505, pp.505-511, 2014.

N. Eisenhauer, P. M. Antunes, A. E. Bennett, K. Birkhofer, A. Bissett et al., , p.455

S. Hart, J. Hättenschwiler, M. Haimi, N. Heethoff, L. C. Kaneko et al., Priorities for research in soil ecology, Pedobiologia (Jena), vol.63, pp.1-7, 2017.

L. Tedersoo, M. Bahram, S. Polme, U. Koljalg, N. S. Yorou et al., , p.460

P. Q. Vasco-palacios, A. Thu, M. E. Suija, C. Smith, E. Sharp et al.,

R. Dunk, J. Drenkhan, A. Dearnaley, T. De-kesel, X. Dang et al., Global diversity and geography of soil fungi, Science, vol.346, pp.1256688-1256688, 2014.

M. Delgado-baquerizo, A. M. Oliverio, T. E. Brewer, A. Benavent-gonzález, D. J. Eldridge et al., A global atlas of the dominant bacteria found in 470 soil. Science, vol.359, pp.320-325, 2018.

M. Bahram, F. Hildebrand, S. K. Forslund, J. L. Anderson, N. A. Soudzilovskaia et al., Structure and function of the global topsoil microbiome, Nature, vol.560, pp.233-237, 2018.

H. Hillebrand, On the Generality of the Latitudinal Diversity Gradient, Am. Nat, vol.163, pp.192-211, 2004.

E. K. Cameron, I. S. Martins, P. Lavelle, J. Mathieu, L. Tedersoo et al.,

. Eisenhauer, Global mismatches in aboveground and belowground biodiversity, Conserv. Biol, p.430, 2019.

N. Fierer, M. S. Strickland, D. Liptzin, M. A. Bradford, and C. C. Cleveland, Global patterns in belowground communities, Ecol. Lett, vol.12, pp.1238-1249, 2009.

J. Van-den-hoogen, S. Geisen, D. Routh, H. Ferris, W. Traunspurger et al., , p.485

B. J. Goede, W. Adams, W. S. Ahmad, R. D. Andriuzzi, M. Bardgett et al.,

T. A. Neilson, U. N. Nguyen, H. Nielsen, J. E. Okada, K. Rius et al., Soil nematode abundance and functional group composition at a global scale, Nature, vol.495, issue.572, pp.194-198, 2019.

T. Decaëns, Macroecological patterns in soil communities, Glob. Ecol. Biogeogr, vol.19, pp.287-302, 2010.

C. A. Edwards, Earthworm ecology, 2004.

M. Blouin, M. E. Hodson, E. A. Delgado, G. Baker, L. Brussaard et al.,

G. Dendooven, J. E. Peres, D. Tondoh, J. J. Cluzeau, and . Brun, A review of earthworm impact on soil function and ecosystem services, Eur. J. Soil Sci, vol.64, pp.161-182, 2013.
URL : https://hal.archives-ouvertes.fr/hal-00818329

D. Craven, M. P. Thakur, E. K. Cameron, L. E. Frelich, R. Beaus�jour et al., , p.505

M. Vellend, L. G. Umek, and N. Eisenhauer, The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta-analysis), Glob. Chang. Biol, vol.23, pp.1065-1074, 2017.

, Supplementary Materials and Methods

M. Rutgers, A. Orgiazzi, C. Gardi, J. Römbke, S. Jänsch et al.,

A. K. Schmidt, R. P. Murchie, G. Blackshaw, D. Pérès, M. Cluzeau et al., Europe. Appl. Soil Ecol, vol.97, pp.98-111, 2016.

P. F. Hendrix and P. J. Bohlen, Exotic earthworm invasions in North America: Ecological and policy 515 implications, Bioscience, vol.52, pp.801-811, 2002.

T. G. Piearce, The calcium relations of selected lumbricidae, J. Anim. Ecol, vol.41, p.167, 1972.

D. J. Spurgeon, A. M. Keith, O. Schmidt, D. R. Lammertsma, and J. H. Faber, Land-use and landmanagement change: relationships with earthworm and fungi communities and soil structural properties, BMC Ecol, vol.13, p.46, 2013.

J. Mathieu and T. J. Davies, Glaciation as an historical filter of below-ground biodiversity, J. Biogeogr, vol.41, pp.1204-1214, 2014.
URL : https://hal.archives-ouvertes.fr/hal-02148223

P. Lavelle, C. Lattaud, D. Trigo, and I. Barois, Mutualism and biodiversity in soils, Plant Soil, vol.170, pp.23-33, 1995.

R. R. Dunn, D. Agosti, A. N. Andersen, X. Arnan, C. A. Bruhl et al.,

M. C. Fisher, H. Fitzpatrick, N. J. Gibb, A. D. Gotelli, B. Gove et al., Climatic drivers of hemispheric asymmetry in global patterns of ant species richness, Ecol. Lett, vol.12, pp.324-333, 2009.

H. Kreft and W. Jetz, Global patterns and determinants of vascular plant diversity, Proc. Natl. Acad. Sci, vol.104, pp.5925-5930, 2007.

C. Fragoso and P. Lavelle, Earthworm communities of tropical rain forests, Soil Biol. Biochem, 1992.

K. J. Gaston and T. M. Blackburn, Pattern and process in macroecology, 2007.

D. Song, K. Pan, A. Tariq, F. Sun, Z. Li et al., Large-scale patterns of distribution and diversity of terrestrial nematodes, Appl. Soil Ecol, vol.114, pp.161-169, 2017.

M. Maraun, H. Schatz, and S. Scheu, Awesome or ordinary? Global diversity patterns of oribatid mites, Ecography (Cop.), vol.30, pp.209-216, 2007.

J. Davison, L. Ainsaar, S. Burla, G. Diedhiou, I. Hiiesalu et al., Global assessment of arbuscular mycorrhizal fungus diversity reveals very low endemism, Science, vol.545, pp.970-973, 2015.

T. Decaëns, D. Porco, S. W. James, G. G. Brown, V. Chassany et al., DNA barcoding reveals diversity patterns of earthworm communities in remote tropical forests of French Guiana, Soil Biol. Biochem, vol.92, pp.171-183, 2016.

D. C. Coleman, D. A. Crossley, and P. F. Hendrix, Fundamentals of Soil Ecology, vol.550, 2004.

J. W. Spaak, J. M. Baert, D. J. Baird, N. Eisenhauer, L. Maltby et al., Shifts of community composition and population density substantially affect ecosystem function despite invariant richness, Ecol. Lett, vol.20, pp.1315-1324, 2017.

N. Eisenhauer, J. Schlaghamerský, P. B. Reich, and L. E. Frelich, The wave towards a new steady state: Effects of earthworm invasion on soil microbial functions, Biol. Invasions, vol.13, pp.2191-2196, 2011.

M. Drumond, A. Guimarães, H. E. Bizri, L. Giovanetti, D. Sepúlveda et al., Life history, distribution and abundance of the giant earthworm Rhinodrilus alatus RIGHI 1971: conservation 560 and management implications, Brazilian J. Biol, vol.73, pp.699-708, 2013.

L. Santini, N. J. Isaac, L. Maiorano, G. F. Ficetola, M. A. Huijbregts et al., Global drivers of population density in terrestrial vertebrates, Glob. Ecol. Biogeogr, vol.27, pp.968-979, 2018.
URL : https://hal.archives-ouvertes.fr/hal-02346328

, Intergovernmental Panel on Climate Change, Climate Change, Synthesis Report Summary 565 Chapter, 2014.

D. K. Hackenberger and B. K. Hackenberger, Earthworm community structure in grassland habitats differentiated by climate type during two consecutive seasons, Eur. J. Soil Biol, vol.61, pp.27-34, 2014.

M. A. Bradford, G. F. Ciska, A. Bonis, E. M. Bradford, A. T. Classen et al.,

J. R. Crowther, G. T. Long, P. Freschet, M. Kardol, D. S. Manrubia-freixa et al.,

R. S. Newman, M. Logtestijn, D. A. Viketoft, W. R. Wardle, S. A. Wieder et al., A test of the hierarchical model of litter decomposition, Nat. Ecol. Evol, vol.1, pp.1836-1845, 2017.
URL : https://hal.archives-ouvertes.fr/hal-01659442

A. Rice, P. ?marda, M. Novosolov, M. Drori, L. Glick et al., The global biogeography of polyploid plants, Nat. Ecol. Evol, vol.3, p.38, 2019.

A. Shade, R. R. Dunn, S. A. Blowes, P. Keil, B. J. Bohannan et al.,

N. J. Lennon, D. Sanders, J. Storch, and . Chase, Macroecology to unite all life, large and small, Trends Ecol. Evol, vol.33, pp.731-744, 2018.

J. M. Anderson and J. S. Ingram, Tropical Soil Biology and Fertility: A handbook of methods, Trop. Soil Biol. Fertil. A Handb. methods. 2 Ed, pp.88-91, 1993.

. Iso, Soil quality -Sampling of soil invertebrates -Part 1: Hand-sorting and extraction of earthworms (ISO/FDIS, pp.23611-23612, 2018.

J. Schindelin, I. Arganda-carreras, E. Frise, V. Kaynig, M. Longair et al., an open-source platform for biological-image analysis, Nat. Methods, vol.9, pp.676-682, 2012.

J. Koricheva, J. Gurevitch, and K. Mengersen, Handbook of meta-analysis in ecology and evolution, 2013.

M. D. Bartlett, M. J. Briones, R. Neilson, O. Schmidt, D. Spurgeon et al., A critical review of current methods in earthworm ecology: From individuals to populations, Eur. J. Soil Biol, vol.46, pp.67-73, 2010.

M. J. Crawley, The R book, 2012.

M. B. Bouché, Strategies lombriciennes. Ecol. Bull, pp.122-132, 1977.

G. G. Brown, How do earthworms affect microfloral and faunal community diversity? Plant Soil, vol.170, pp.209-231, 1995.

J. Seeber, G. U. Seeber, R. Langel, S. Scheu, and E. Meyer, The effect of macro-invertebrates and plant litter of different quality on the release of N from litter to plant on alpine pastureland, Biol. Fertil. Soils, vol.44, pp.783-790, 2008.

M. Blouin, Y. Zuily-fodil, A. T. Pham-thi, D. Laffray, G. Reversat et al., Belowground organism activities affect plant aboveground phenotype, inducing plant tolerance to parasites, Ecol. Lett, vol.8, pp.202-208, 2005.
URL : https://hal.archives-ouvertes.fr/bioemco-00446710

J. Boyer, G. Reversat, P. Lavelle, and A. Chabanne, European Journal of Soil Biology Interactions between earthworms and plant-parasitic nematodes, Eur. J. Soil Biol, vol.59, p.50, 2013.

G. Loranger-merciris, Y. Cabidoche, B. Deloné, P. Quénéhervé, and H. Ozier-lafontaine, How earthworm activities affect banana plant response to nematodes parasitism, Appl. Soil Ecol, vol.52, pp.1-8, 2012.

G. G. Brown, E. Soja, C. A. Edwards, and L. Brussaard, Earthworm Ecology, vol.610, p.52, 2004.

M. B. Bouché and F. Al-addan, Earthworms, water infiltration and soil stability: Some new assessments, Soil Biol. Biochem, vol.29, pp.441-452, 1997.

M. Joschko, H. Diestel, and O. Larink, Assessment of earthworm burrowing efficiency in compacted soil with a combination of morphological and soil physical measurements, Biol. Fertil. Soils, vol.8, pp.191-196, 1989.

T. Hengl, J. Mendes-de-jesus, G. B. Heuvelink, M. Gonzalez, M. Kilibarda et al., SoilGrids250m: Global gridded soil information based on machine learning, PLoS One, vol.12, p.169748, 2017.

D. N. Karger, O. Conrad, J. Böhner, T. Kawohl, H. Kreft et al., Climatologies at high resolution for the earth's land surface areas, Sci. Data, vol.4, p.170122, 2017.

D. K. Hall and G. A. Riggs, MODIS/Terra Snow Cover Monthly L3 Global 0.05Deg CMG, Version 6, NASA National Snow and Ice Data Center Distributed Active Archive, vol.625

. Center and . Doi, , 2015.

R. J. Zomer, A. Trabucco, D. A. Bossio, and L. V. Verchot, Climate change mitigation: A spatial analysis of global land suitability for clean development mechanism afforestation and reforestation, Agric. Ecosyst. Environ, vol.126, pp.67-80, 2008.

R. J. Zomer, D. A. Bossio, A. Trabucco, L. Yuanjie, D. C. Gupta et al., Trees and Water, p.630

, Smallholder Agroforestry on Irrigated Lands in Northern India. IWMI Res. Rep, vol.122, p.45, 2007.

J. Danielson and D. Gesch, Global Multi-resolution Terrain Elevation Data 2010(GMTED2010), 2011.
URL : https://hal.archives-ouvertes.fr/hal-00747183

D. Bates, M. Mächler, B. Bolker, and S. Walker, Fitting Linear Mixed-Effects Models Using lme4, J. Stat. Softw, vol.67, pp.1-48, 2015.

A. F. Zuur, E. N. Ieno, and C. S. Elphick, A protocol for data exploration to avoid common statistical problems, Methods Ecol. Evol, vol.1, pp.3-14, 2010.

N. Eisenhauer, A. Stefanski, N. A. Fisichelli, K. Rice, R. Rich et al., Warming shifts "worming": Effects of experimental warming on invasive earthworms in northern, North America. Sci. Rep, vol.4, pp.4-10, 2014.

M. Nieminen, E. Ketoja, J. Mikola, J. Terhivuo, T. Siren et al., Local land use effects and regional environmental limits on earthworm communities in Finnish arable landscapes, Ecol. Appl, vol.21, pp.3162-3177, 2011.

A. F. Zuur, E. N. Ieno, and A. A. Saveliev, Mixed Effects Models and Extensions in Ecology with, 2009.

S. Dray and A. Dufour, The ade4 Package: Implementing the Duality Diagram for Ecologists, J. Stat. Softw, vol.22, pp.1-20, 2007.
URL : https://hal.archives-ouvertes.fr/hal-00434575

L. Breiman, Random forests, Mach. Learn, vol.45, pp.5-32, 2001.

A. Liaw and M. Wiener, Classification and regression by randomForest. R news, vol.2, pp.18-22, 2002.

U. Grömping, Variable importance assessment in regression: Linear regression versus random 650 forest, Am. Stat, vol.63, pp.308-319, 2009.

C. Strobl, A. Boulesteix, A. Zeileis, and T. Hothorn, Bias in random forest variable importance measures: Illustrations, sources and a solution, BMC Bioinformatics, vol.8, p.25, 2007.

B. H. Menze, B. M. Kelm, R. Masuch, U. Himmelreich, P. Bachert et al., A comparison of random forest and its Gini importance with standard chemometric methods for 655 the feature selection and classification of spectral data, BMC Bioinformatics, vol.10, p.213, 2009.

G. James, D. Witten, T. Hastie, and R. Tibshirani, An introduction to statistical learning, vol.112, 2013.

K. Barton, MuMIn: Multi-Model Inference. R Packag. version 1.42.1. https//CRAN.Rproject.org/package=MuMIn, 2018.

N. Gorelick, M. Hancher, M. Dixon, S. Ilyushchenko, D. Thau et al., Google Earth Engine: Planetary-scale geospatial analysis for everyone. Remote Sens. Environ, 2017.

. R-core-team, R: A language and environment for statistical computing, 2016.

P. Lavelle and A. Spain, Soil ecology, 2001.
URL : https://hal.archives-ouvertes.fr/bioemco-00455666

P. A. Sanchez, S. Ahamed, F. Carre, A. E. Hartemink, J. Hempel et al., Digital Soil Map of the World. Science, vol.325, pp.680-681, 2009.

, Royal Canadian Geographical Society, Environmental Protection Agency, p.2005

R. P. Helen, C. A. Phillips, M. L. Guerra, M. J. Bartz, G. Briones et al.,

M. B. Whalen, V. Wironen, I. V. Wolters, W. Zenkova, E. K. Zhang et al., Nico Eisenhauer Correspondence to: helen.phillips@idiv.de This PDF file includes: models when all other variables are at zero, i.e., the mean. Not all habitat cover categories had sampled estimates (i.e., species richness could not be estimated for

, Habitat cover at a sampled site was classified based on the 'Reclassified habitat cover' column. As not all categories of habitat were available in the data (i.e., due to too detailed categories, or in habitats typically devoid of sampling), some of the categories of the original habitat cover variable (left-hand column) were reclassified (right-hand column). Usually, this meant that categories were grouped together, The re-categorisation of the ESA habitat cover variable