Linking species, traits and habitat characteristics of Collembola at European scale

Although much work has been done on factors which influence the patterning of species and species trait assemblages in a variety of groups such as plants, vertebrates and invertebrates, few studies have been realized at a broad geographic scale. We analyzed patterns of relationships between species, species trait distribution/assembly, and environmental variables from the west of Europe to Slovakia, Poland and Sweden. We created a database by compiling traits and occurrence data of European collembolan species, using literature and personal field studies embracing a large range of environmental gradients (vertical stratification, habitat closure, humus form, soil acidity and moisture, temperature, rainfall, altitude) over which Collembola are supposed to be distributed. Occurrences of the 58 best-documented species, environmental variables and species traits allowed us to (i) show which environmental variables impact the distribution of the 58 species at broad scale and (ii) document to what extent environmental variables and species trait assemblages are related and which trends could be found in trait/environment relationships. The impact of vertical stratification, habitat closure, humus form, soil acidity, soil moisture, temperature, and to a lesser extent rainfall and altitude on species distribution, firstly revealed by indirect gradient analysis (correspondence analysis, CA), was further shown to be significant by direct gradient analysis (canonical correspondence analysis, CCA). RLQ analyses were performed to find linear combination of variables of table R (environmental variables) and linear combinations of the variables of table Q (species traits) of maximum covariance weighted by species occurrence data contained in table L. RLQ followed by permutation tests showed that all tested environmental variables apparently contributed significantly to the assemblages of the twelve species traits studied. A convergence was observed between traits related to vertical stratification and those related to habitat closure/aperture. Well-developed locomotory organs (furcula, legs), presence of sensorial organs sensitive to air movements and light (e.g. trichobothria and eye spots), spherical body, large body size, pigmentation (UV protection and signaling) and sexual reproduction largely occur in epigeic and open habitats, while most of woodland and edaphic habitats are characterized by short locomotory appendages, small body size, high number of defense organs (pseudocelli), presence of post-antennal organs and parthenogenesis. Climate and especially temperature exert an effect on the assemblage of traits that are mostly present above-ground and in open habitats. The contribution of combinations of some environmental variables to the occurrence of each species trait was tested by linear, logistic or multinomial regression (Generalized Linear Models). Vertical stratification, followed by temperature, played a dominant role in the variation of the twelve studied traits. Relationships between traits and environment tested here shows that it is possible to use some traits as proxies to identify potential ecological preferences or tolerances of invertebrate species. However, a significant part of species distribution remained unexplained, probably partly because some traits, like ecophysiological ones, or traits involved in biotic interactions (e.g. competition) were unavailable. The present work is thus a first step towards the creation of models predicting changes in collembolan communities. Further studies are required to inform ecophysiological traits, in order to complete such models. Moreover the niche width of species will have to be determined. (C) 2014 Elsevier Ltd. All rights reserved.