Does trait-based joint species distribution modelling reveal the signature of competition in stream macroinvertebrate communities?

The occupancy and abundance of species are jointly driven by local factors, such as environmental characteristics and biotic interactions, and regional-scale factors, such as dispersal and climate. Recently, it has been shown that biotic interactions shape species occupancies and abundances beyond local extents. However, for small ectothermic animals, particularly for those occurring in freshwater environments, the importance of biotic interactions remains understudied. Species-to-species associations from joint species distribution models (i.e. species associations while controlling for environmental characteristics) are increasingly used to draw hypotheses of which species possibly show biotic interactions. We studied whether species-to-species associations from joint species distribution models show signs of competition using a hypothesis testing framework in stream macroinvertebrate communities at regional extent. We sampled aquatic macroinvertebrates from 105 stream sites in western Finland encompassing a latitudinal gradient of c. 500 kilometres. We hypothesized that if competition drives these associations (H1) functionally, similar species are mostly negatively associated, whereas functionally dissimilar species show random associations. We further hypothesized that the relationship between functional dissimilarity and the strength of association is more pronounced (H2) for abundances rather than occupancies, (H3) at small grain (i.e. stream site) rather than at large grain (i.e. river basin), and (H4) among species having weak dispersal ability than among species with high dispersal ability. Stream macroinvertebrates showed both negative and positive species-to-species associations while controlling for habitat characteristics. However, the negative associations were mostly at large grain (river basin) rather than at small grain (stream site), in occupancy rather than abundance, and not related to species functional dissimilarity or to their dispersal ability. Thus, all our hypotheses considering possible competition (H1-H4) were rejected. Competition does not appear to be a major driving force of stream macroinvertebrate communities at the spatial grain sizes considered. The observed positive associations in occupancy at small grain (stream site) may be attributed to species' similar microhabitat preferences, whereas at large grain (river basin), they may stem from metacommunity dynamics. Our results highlight that species traits were necessary to interpret whether or not species-to-species associations from joint species distribution models resulted from biotic interactions.

The effect of peatland drainage and restoration on Odonata species richness and abundance.

BACKGROUND: Restoration aims at reversing the trend of habitat degradation, the major threat to biodiversity. In Finland, more than half of the original peatland area has been drained, and during recent years, restoration of some of the drained peatlands has been accomplished. Short-term effects of the restoration on peatland hydrology, chemistry and vegetation are promising but little is known about how other species groups apart from vascular plants and bryophytes respond to restoration efforts. RESULTS: Here, we studied how abundance and species richness of Odonata (dragonflies and damselflies) respond to restoration. We sampled larvae in three sites (restored, drained, pristine) on each of 12 different study areas. We sampled Odonata larvae before restoration (n = 12), during the first (n = 10) and the third (n = 7) year after restoration and used generalized linear mixed models to analyze the effect of restoration. Drained sites had lower abundance and species richness than pristine sites. During the third year after restoration both abundance and species richness had risen in restored sites. CONCLUSIONS: Our results show that Odonata suffer from drainage, but seem to benefit from peatland restoration and are able to colonize newly formed water pools already within three years after restoration.

Titmice are a better indicator of bird density in Northern European than in Western European forests.

Population sizes of many birds are declining alarmingly and methods for estimating fluctuations in species' abundances at a large spatial scale are needed. The possibility to derive indicators from the tendency of specific species to co-occur with others has been overlooked. Here, we tested whether the abundance of resident titmice can act as a general ecological indicator of forest bird density in European forests. Titmice species are easily identifiable and have a wide distribution, which makes them potentially useful ecological indicators. Migratory birds often use information on the density of resident birds, such as titmice, as a cue for habitat selection. Thus, the density of residents may potentially affect community dynamics. We examined spatio-temporal variation in titmouse abundance and total bird abundance, each measured as biomass, by using long-term citizen science data on breeding forest birds in Finland and France. We analyzed the variation in observed forest bird density (excluding titmice) in relation to titmouse abundance. In Finland, forest bird density linearly increased with titmouse abundance. In France, forest bird density nonlinearly increased with titmouse abundance, the association weakening toward high titmouse abundance. We then analyzed whether the abundance (measured as biomass) of random species sets could predict forest bird density better than titmouse abundance. Random species sets outperformed titmice as an indicator of forest bird density only in 4.4% and 24.2% of the random draws, in Finland and France, respectively. Overall, the results suggest that titmice could act as an indicator of bird density in Northern European forest bird communities, encouraging the use of titmice observations by even less-experienced observers in citizen science monitoring of general forest bird density.

Environmental Characteristics and Anthropogenic Impact Jointly Modify Aquatic Macrophyte Species Diversity.

Species richness and spatial variation in community composition (i.e., beta diversity) are key measures of biodiversity. They are largely determined by natural factors, but also increasingly affected by anthropogenic factors. Thus, there is a need for a clear understanding of the human impact on species richness and beta diversity, the underlying mechanisms, and whether human-induced changes can override natural patterns. Here, we dissect the patterns of species richness, community composition and beta diversity in relation to different environmental factors as well as human impact in one framework: aquatic macrophytes in 66 boreal lakes in Eastern Finland. The lakes had been classified as having high, good or moderate status (according to ecological classification of surface waters in Finland) reflecting multifaceted human impact. We used generalized least square models to study the association between different environmental variables (Secchi depth, irregularity of the shoreline, total phosphorus, pH, alkalinity, conductivity) and species richness. We tested the null hypothesis that the observed community composition can be explained by random distribution of species. We used multivariate distance matrix regression to test the effect of each environmental variable on community composition, and distance-based test for homogeneity of multivariate dispersion to test whether lakes classified as high, good or moderate status have different beta diversity. We showed that environmental drivers of species richness and community composition were largely similar, although dependent on the particular life-form group studied. The most important ones were characteristics of water quality (pH, alkalinity, conductivity) and irregularity of the shoreline. Differences in community composition were related to environmental variables independently of species richness. Species richness was higher in lakes with higher levels of human impact. Lakes with different levels of human impact had different community composition. Between-lake beta diversity did not differ in high, good or moderate status groups. However, the variation in environmental variables shaping community composition was larger in lakes with moderate status compared to other lakes. Hence, beta diversity in lakes with moderate status was smaller than what could be expected on the basis of these environmental characteristics. This could be interpreted as homogenization.

Ecological response hides behind the species abundance distribution: Community response to low-intensity disturbance in managed grasslands.

Land-use and management are disturbance factors that have diverse effects on community composition and structure. In traditional rural grasslands, such as meadows and pastures, low-intensity management is maintained to enhance biodiversity. Maintenance of road verges, in turn, creates habitat, which may complement traditional rural grasslands. To evaluate the effect of low-intensity disturbance on insect communities, we characterized species abundance distributions (SAD) for Carabidae, Formicidae, and Heteroptera in three grassland types, which differed in management: meadows, pastures, and road verges. The shape of SAD was estimated with three parameters: abundance decay rate, dominance, and rarity. We compared the SAD shape among the grassland types and tested the effect of environmental heterogeneity (plant species richness) and disturbance intensity (trampling in pastures) on SADs. The shape of SADs did not differ among the grassland types but among the taxonomic groups instead. Abundance decay rate and dominance were larger for Formicidae, and rarity smaller, than for Carabidae and Heteroptera. For Carabidae and window-trapped Heteroptera, rarity increased with increasing plant species richness. For Formicidae, dominance increased with trampling intensity in pastures. Although the SAD shape remained largely unchanged, the identity of the dominant species tended to vary within and among grassland types. Our study shows that for a given taxonomic group, the SAD shape is similar across habitat types with low-intensity disturbances resulting from different management. This suggests that SADs respond primarily to the intensity of disturbance and thus could be best used in monitoring communities across strong disturbance and environmental gradients. Because taxonomic groups can inherently have different SADs, taxon-specific SADs for undisturbed communities must be empirically documented before the SAD shape can be used as an indicator of environmental change. Because the identity of the dominant species changes from management type to another, the SAD shape alone is not an adequate monitoring tool.