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Aim There has been a wide interest in the effect of biotic interactions on species' occurrences and abundances at large spatial scales, coupled with a vast development of the statistical methods to study them. Still, evidence for whether the effects of within-trophic-level biotic interactions (e.g. competition and heterospecific attraction) are discernible beyond local scales remains inconsistent. Here, we present a novel hypothesis-testing framework based on joint dynamic species distribution models and functional trait similarity to dissect between environmental filtering and biotic interactions. Location France and Finland. Taxon Birds. Methods We estimated species-to-species associations within a trophic level, independent of the main environmental variables (mean temperature and total precipitation) for common species at large spatial scale with joint dynamic species distribution (a multivariate spatiotemporal delta model) models. We created hypotheses based on species' functionality (morphological and/or diet dissimilarity) and habitat preferences about the sign and strength of the pairwise spatiotemporal associations to estimate the extent to which they result from biotic interactions (competition, heterospecific attraction) and/or environmental filtering. Results Spatiotemporal associations were mostly positive (80%), followed by random (15%), and only 5% were negative. Where detected, negative spatiotemporal associations in different communities were due to a few species. The relationship between spatiotemporal association and functional dissimilarity among species was negative, which fulfils the predictions of both environmental filtering and heterospecific attraction. Main conclusions We showed that processes leading to species aggregation (mixture between environmental filtering and heterospecific attraction) seem to dominate assembly rules, and we did not find evidence for competition. Altogether, our hypothesis-testing framework based on joint dynamic species distribution models and functional trait similarity is beneficial in ecological interpretation of species-to-species associations from data covering several decades and biogeographical regions. peerReviewed

Publisher Copyright: © 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. 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.

1. 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. 2. 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. 3. We sampled aquatic macroinvertebrates from 105 stream sites in western Finland encompassing a latitudinal gradient of ca. 500 kilometers. 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. 4. 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. 5. 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. peerReviewed

Grassland biodiversity, including traditional rural biotopes maintained by traditional agricultural practices, has become threatened worldwide. Road verges have been suggested to be complementary or compensatory habitats for species inhabiting grasslands. Species co-occurrence patterns linked with species traits can be used to separate between the different mechanisms (stochasticity, environmental filtering, biotic interactions) behind community structure. Here, we study species co-occurrence networks and underlying mechanisms of ground beetle species (Carabidae) in three different managed grassland types (meadows, pastures, road verges, n = 12 in each type) in Central Finland. We aimed to find out whether road verges can be considered as compensatory to traditional rural biotopes (meadows and pastures). We found that stochasticity explained over 90% of the pairwise co-occurrences, and the non-random co-occurrences were best explained by environmental filtering, regardless of the grassland type. However, the identities and traits of the species showing non-random co-occurrences differed among the habitat types. Thus, environmental factors behind environmental filtering differ among the habitat types and are related to the site-specific characteristics and variation therein. This poses challenges to habitat management since the species’ response to management action may depend on the site-specific characteristics. Although road verges are not fully compensatory to meadows and pastures, the high similarity of species richness and the high level of shared species suggest that for carabids road verges may be corridors connecting the sparse network of the remaining traditional rural biotopes.

Recently, Moreno-Mateos et al. (2017) coined the concept ‘recovery debt’, clearly a close relative of the ecosystem service debt (Isbell et al. 2015), and gave it significance as “the interim reduction of biodiversity and biogeochemical functions occurring during ecosystem recovery”. Using rather impressive dataset consisting 3,035 sampling plots worldwide as an example, they analysed the recovery debt for plant and animal species diversity and abundance as well as for carbon and nitrogen cycling. Based on their analysis Moreno-Mateos et al. conclude that “… recovering and restored ecosystems have less abundance, diversity and cycling of carbon and nitrogen than ‘undisturbed’ ecosystems …”. Here, we scrutinize the proposed new concept and point out problems in conclusions resulting from the operationalization of the concept.

Aim Understanding the relative contribution of different species interactions in shaping community assembly has been a pivotal aim in community ecology. Biotic interactions are acknowledged to be important at local scales, although their signal is assumed to weaken over longer distances. We examine the relationship between positive, neutral and negative pairwise bird abundance distributions and the phylogenetic and functional distance between these pairs after first controlling for habitat associations. Location France and Finland. Time period 1984 to 2011 (Finland), 2001 to 2012 (France). Major Taxa studied Birds. Methods We used results from French and Finnish land bird monitoring programmes, from which we created three independent datasets (French forests, French farmlands and Finnish forests). Separately for the three datasets, we fitted linear mixed-effects models for pairwise abundance values across years per point count station to infer the association between all common species pairs, while controlling for geographical distribution and habitat associations, and saved pairwise regression coefficients for further analyses. We used a null model approach to infer whether the observed associations (effect sizes) differ from random. Finally, using quantile regression, we analysed the relationships between functional dissimilarity/phylogenetic distance and effect sizes. Results Our results show both negative and positive species interactions, although negative interactions were twice as common as positive interactions. Closely related species were more likely to show strong associations, both negative and positive, than more distant species across broad spatial scales. For functional dissimilarity, the results varied across datasets. Main conclusions Our results emphasize the potential of functional and phylogenetic proximity in generating both negative and positive species associations, which can produce pervasive patterns from local to geographical scales. Future assembly studies should refrain from strict dichotomies, such as compensatory dynamics versus environmental forcing, and instead consider the possibility of positive interactions.

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. peerReviewed

Distribution of species across the Earth shows strong latitudinal and altitudinal gradients with the number of species decreasing with declining temperatures. While these patterns have been recognized for well over a century, the mechanisms generating and maintaining them have remained elusive. Here, we propose a mechanistic explanation for temperature-dependent rates of molecular evolution that can influence speciation rates and global biodiversity gradients. Our hypothesis is based on the effects of temperature and temperature-adaptation on stability of proteins and other catalytic biomolecules. First, due to the nature of physical forces between biomolecules and water, stability of biomolecules is maximal around + 20°C and decreases as temperature either decreases or increases. Second, organisms that have adapted to cold temperatures have evolved especially flexible (but unstable) proteins to facilitate catalytic reactions in cold, where molecular movements slow down. Both these effects should result in mutations being on average more detrimental at cold temperatures (i.e. lower mutational robustness in cold). At high temperatures, destabilizing water–biomolecule interactions, and the need to maintain structures that withstand heat denaturation, should decrease mutational robustness similarly. Decreased mutational robustness at extreme temperatures will slow down molecular evolution, as a larger fraction of new mutations will be removed by selection. Lower mutational robustness may also select for reduced mutation rates, further slowing down the rate of molecular evolution. As speciation requires the evolution of epistatic incompatibilities that prevent gene flow among incipient species, slow rate of molecular evolution at extreme temperatures will directly slow down the rate at which new species arise. The proposed mechanism can thus explain why molecular evolution is faster at warm temperatures, contributing to higher speciation rate and elevated species richness in environments characterized by stable and warm temperatures.

Anthropogenic disturbance often causes changes in communities. However, the mechanistic basis of these changes remains elusive. As all patterns in community ecology can be understood as a result of four processes (speciation, selection, drift, and dispersal), the effect of disturbance should depend on how disturbance disrupt these processes. We studied the effects of disturbance and productivity on species richness, community composition, and community dispersion (i.e., variation in community composition) in the vegetation of 120 boreal peatlands using null-model approach to determine whether community assembly processes differ between pristine and disturbed sites. Sites represented three peatland ecosystem types, each with two levels of productivity. Half of the sites were disturbed by drainage and half are pristine. Pristine and disturbed sites showed similar species richness. However, their community composition differed indicating a directional selection due to disturbance, whereas dispersion of disturbed and pristine communities did not differ suggesting no change in the relative strength of selection and drift. Our results suggest that understanding the combination of landscape level community changes and local selection pressures is important when restoration of degraded ecosystems is undertaken. peerReviewed

Aim An area’s ability to support species may be dependent not only on the total amount of available energy it contains but also on energy density (i.e. available energy per unit area). Acknowledging these two aspects of energy availability may increase mechanistic understanding of how increased energy availability results in increased species richness. We studied the relationship between energy density, its variation in space and boreal forest bird species richness and investigated two possible mechanisms: (1) metabolic constraints of organisms, and (2) increased resource availability for specialists. Location Protected areas in Finland’s boreal forest. Methods We tested whether bird species richness was best determined by total energy availability in an area or by energy density and its variation within the area, before and after including bird abundance in the models. We evaluated two main explanatory variables: tree growth reflecting the rate of energy production and tree volume as a measure of biomass. In addition, we modelled individual species’ responses to energy density and its variation, and evaluated the prediction of the metabolic constraints hypothesis that small species are limited by energy density whereas large species are limited by total energy availability in the area. Results Energy density and its variation were good predictors of species richness: together with abundance they explained 84% of variation in species richness (compared with 74% for abundance alone). Pure metabolic constraints were unlikely to explain this relationship. Instead, the mechanism probably involved increased habitat heterogeneity benefiting specialist species. Total energy availability was also an important factor determining species richness but its effect was indirect via abundance. Main conclusions Our results corroborate the importance of energy availability as a driver of species richness in forest bird communities, and they indicate that energy density and its variation in the landscape strongly influence species richness even after accounting for 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. Electronic supplementary material The online version of this article (doi:10.1186/s12898-015-0042-z) contains supplementary material, which is available to authorized users.