Soil macroinvertebrate communities: A world-wide assessment

Aim Macroinvertebrates comprise a highly diverse set of taxa with great potential as indicators of soil quality. Communities were sampled at 3,694 sites distributed world-wide. We aimed to analyse the patterns of abundance, composition and network characteristics and their relationships to latitude, mean annual temperature and rainfall, land cover, soil texture and agricultural practices. Location Sites are distributed in 41 countries, ranging from 55 degrees S to 57 degrees N latitude, from 0 to 4,000 m in elevation, with annual rainfall ranging from 500 to >3,000 mm and mean temperatures of 5-32 degrees C. Time period 1980-2018. Major taxa studied All soil macroinvertebrates: Haplotaxida; Coleoptera; Formicidae; Arachnida; Chilopoda; Diplopoda; Diptera; Isoptera; Isopoda; Homoptera; Hemiptera; Gastropoda; Blattaria; Orthoptera; Lepidoptera; Dermaptera; and "others". Methods Standard ISO 23611-5 sampling protocol was applied at all sites. Data treatment used a set of multivariate analyses, principal components analysis (PCA) on macrofauna data transformed by Hellinger's method, multiple correspondence analysis for environmental data (latitude, elevation, temperature and average annual rainfall, type of vegetation cover) transformed into discrete classes, coinertia analysis to compare these two data sets, and bias-corrected and accelerated bootstrap tests to evaluate the part of the variance of the macrofauna data attributable to each of the environmental factors. Network analysis was performed. Each pairwise association of taxonomic units was tested against a null model considering local and regional scales, in order to avoid spurious correlations. Results Communities were separated into five clusters reflecting their densities and taxonomic richness. They were significantly influenced by climatic conditions, soil texture and vegetation cover. Abundance and diversity, highest in tropical forests (1,895 +/- 234 individuals/m(2)) and savannahs (1,796 +/- 72 individuals/m(2)), progressively decreased in tropical cropping systems (tree-associated crops, 1,358 +/- 120 individuals/m(2); pastures, 1,178 +/- 154 individuals/m(2); and annual crops, 867 +/- 62 individuals/m(2)), temperate grasslands (529 +/- 60 individuals/m(2)), forests (232 +/- 20 individuals/m(2)) and annual crops (231 +/- 24 individuals/m(2)) and temperate dry forests and shrubs (195 +/- 11 individuals/m(2)). Agricultural management decreased overall abundance by <= 54% in tropical areas and 64% in temperate areas. Connectivity varied with taxa, with dominant positive connections in litter transformers and negative connections with ecosystem engineers and Arachnida. Connectivity and modularity were higher in communities with low abundance and taxonomic richness. Main conclusions Soil macroinvertebrate communities respond to climatic, soil and land-cover conditions. All taxa, except termites, are found everywhere, and communities from the five clusters cover a wide range of geographical and environmental conditions. Agricultural practices significantly decrease abundance, although the presence of tree components alleviates this effect.