Accounting for temporal change in multiple biodiversity patterns improves the inference of metacommunity processes.

In metacommunity ecology, a major focus has been on combining observational and analytical approaches to identify the role of critical assembly processes, such as dispersal limitation and environmental filtering, but this work has largely ignored temporal community dynamics. Here, we develop a "virtual ecologist" approach to evaluate assembly processes by simulating metacommunities varying in three main processes: density-independent responses to abiotic conditions, density-dependent biotic interactions, and dispersal. We then calculate a number of commonly used summary statistics of community structure in space and time and use random forests to evaluate their utility for inferring the strength of these three processes. We find that (i) both spatial and temporal data are necessary to disentangle metacommunity processes based on the summary statistics we test, and including statistics that are measured through time increases the explanatory power of random forests by up to 59% compared to cases where only spatial variation is considered; (ii) the three studied processes can be distinguished with different descriptors; and (iii) each summary statistic is differently sensitive to temporal and spatial sampling effort. Including repeated observations of metacommunities over time was essential for inferring the metacommunity processes, particularly dispersal. Some of the most useful statistics include the coefficient of variation of species abundances through time and metrics that incorporate variation in the relative abundances (evenness) of species. We conclude that a combination of methods and summary statistics is probably necessary to understand the processes that underlie metacommunity assembly through space and time, but we recognize that these results will be modified when other processes or summary statistics are used.

Perils of life on the edge: Climatic threats to global diversity patterns of wetland macroinvertebrates.

Climate change is rapidly driving global biodiversity declines. How wetland macroinvertebrate assemblages are responding is unclear, a concern given their vital function in these ecosystems. Using a data set from 769 minimally impacted depressional wetlands across the globe (467 temporary and 302 permanent), we evaluated how temperature and precipitation (average, range, variability) affects the richness and beta diversity of 144 macroinvertebrate families. To test the effects of climatic predictors on macroinvertebrate diversity, we fitted generalized additive mixed-effects models (GAMM) for family richness and generalized dissimilarity models (GDMs) for total beta diversity. We found non-linear relationships between family richness, beta diversity, and climate. Maximum temperature was the main climatic driver of wetland macroinvertebrate richness and beta diversity, but precipitation seasonality was also important. Assemblage responses to climatic variables also depended on wetland water permanency. Permanent wetlands from warmer regions had higher family richness than temporary wetlands. Interestingly, wetlands in cooler and dry-warm regions had the lowest taxonomic richness, but both kinds of wetlands supported unique assemblages. Our study suggests that climate change will have multiple effects on wetlands and their macroinvertebrate diversity, mostly via increases in maximum temperature, but also through changes in patterns of precipitation. The most vulnerable wetlands to climate change are likely those located in warm-dry regions, where entire macroinvertebrate assemblages would be extirpated. Montane and high-latitude wetlands (i.e., cooler regions) are also vulnerable to climate change, but we do not expect entire extirpations at the family level.

Addressing trait selection patterns in temporary ponds in response to wildfire disturbance and seasonal succession.

Mediterranean ecosystems are increasingly threatened by disturbances such as wildfires. These disturbances are expected to shift the selective pressures that determine trait-dependent community assembly. In addition, the stochasticity in species assembly may decrease because of the introduction of strong selection regimes or may increase because of random variation in recruitment. However, these changes in the selection profile and stochasticity in disturbed communities have seldom been evaluated. We examined the relative roles of wildfire disturbance, local conditions and successional dynamics on the assembly of aquatic macroinvertebrate communities. We used the theory of community assembly by trait selection (CATS) to identify traits under selection and to estimate their dependence on wildfire disturbance and environmental gradients. We took advantage of a natural wildfire that partially burned a Mediterranean system of temporary ponds, which were surveyed before and after the wildfire, creating a natural before-after-control-impact design. Before the wildfire, the burned and unburned ponds did not show differences in the selected traits. After the wildfire event, species with larger body sizes and scrapers were favoured in the burned ponds, while collectors and active dispersers were underrepresented. Nonetheless, local environmental conditions and successional dynamics had greater relevance in the selection of traits than the wildfire. This suggests that assembly mechanisms were largely determined by seasonal successional changes regardless of wildfire disturbance. Finally, the relevance of the analysed traits diminished during the hydroperiod, suggesting more stochastic assemblages and/or a replacement in the set of selected traits. Despite the prominent role of seasonal succession over wildfire, this disturbance was associated with a change in the selection strength over specific traits related with feeding strategies and species life histories. Both hydroperiod and wildfire highlighted a strong role of trait-mediated processes (i.e. niche assembly). Therefore, the predicted increase in the frequency and intensity of wildfires is likely to result in community functional shifts. Furthermore, stochasticity was also important for community assembly, particularly from the middle towards the end of the hydroperiod. Our results evidenced the strong relevance of successional changes in trait-mediated assembly mechanisms and its interplay with wildfire disturbance in temporary pond communities.

Dispersal mode and spatial extent influence distance-decay patterns in pond metacommunities.

Assuming that dispersal modes or abilities can explain the different responses of organisms to geographic or environmental distances, the distance-decay relationship is a useful tool to evaluate the relative role of local environmental structuring versus regional control in community composition. Based on continuing the current theoretical framework on metacommunity dynamics and based on the predictive effect of distance on community similarity, we proposed a new framework that includes the effect of spatial extent. In addition, we tested the validity of our proposal by studying the community similarity among three biotic groups with different dispersal modes (macrofaunal active and passive dispersers and plants) from two pond networks, where one network had a small spatial extent, and the other network had an extent that was 4 times larger. Both pond networks have similar environmental variability. Overall, we found that environmental distance had larger effects than geographical distances in both pond networks. Moreover, our results suggested that species sorting is the main type of metacommunity dynamics shaping all biotic groups when the spatial extent is larger. In contrast, when the spatial extent is smaller, the observed distance-decay patterns suggested that different biotic groups were mainly governed by different metacommunity dynamics. While the distance-decay patterns of active dispersers better fit the trend that was expected when mass effects govern a metacommunity, passive dispersers showed a pattern that was expected when species sorting prevails. Finally, in the case of plants, it is difficult to associate their distance-decay patterns with one type of metacommunity dynamics.

Environmental filtering determines metacommunity structure in wetland microcrustaceans.

Metacommunity approaches are becoming popular when analyzing factors driving species distribution at the regional scale. However, until the popularization of the variation partitioning technique it was difficult to assess the main drivers of the observed patterns (spatial or environmental). Here we propose a new framework linking the emergence of different metacommunity structures (e.g., nested, Gleasonian, Clementsian) to spatial and environmental filters. This is a novel approach that provides a more profound analysis of how both drivers could lead to similar metacommunity structures. We tested this framework on 110 sites covering a strong environmental gradient (i.e., microcrustacean assemblages organized along a salinity gradient, from freshwater to brackish water wetlands). First we identified the metacommunity structure that better fitted these microcrustacean assemblages. Then, we used hierarchical variation partitioning to quantify the relative influences of environmental filters and the distance among wetlands on the identified structure. Our results showed that under strong environmental filtering metacommunity structures were non-random. We also noted that even passive dispersers, that are supposed to be poorly spatially filtered, showed spatial signals at a large geographical scale. However, some difficulties arose when inferring biotic interactions at finer-scale spatial signals. Overall, our study shows the potential of elements of metacommunity structure combined with variation partition techniques to detect environmental drivers and broadscale patterns of metacommunity structure, and that some caution is needed when interpreting finer-scale spatial signals.

Spatio-temporal variability of faunal and floral assemblages in Mediterranean temporary wetlands.

Six temporary wetlands in the region of Sejenane (Mogods, NW Tunisia) were studied in order to characterize the aquatic flora and fauna and to quantify their spatio-temporal variability. Samplings of aquatic fauna, phytosociological relevés, and measurements of the physicochemical parameters of water were taken during four different field visits carried out during the four seasons of the year (November 2009-July 2010). Despite the strong anthropic pressures on them, these temporary wetlands are home to rich and diversified biodiversity, including rare and endangered species. Spatial and temporal variations affect fauna and flora differently, as temporal variability influences the fauna rather more than the plants, which are relatively more dependent on spatial factors. These results demonstrate the interest of small water bodies for maintaining biodiversity at the regional level, and thus underscore the conservation issues of Mediterranean temporary wetlands that are declining on an ongoing basis currently.

Functional and phylogenetic relatedness in temporary wetland invertebrates: current macroecological patterns and implications for future climatic change scenarios.

In freshwater ecosystems, species compositions are known to be determined hierarchically by large to small­scale environmental factors, based on the biological traits of the organisms. However, in ephemeral habitats this heuristic framework remains largely untested. Although temporary wetland faunas are constrained by a local filter (i.e., desiccation), we propose its magnitude may still depend on large-scale climate characteristics. If this is true, climate should be related to the degree of functional and taxonomic relatedness of invertebrate communities inhabiting seasonal wetlands. We tested this hypothesis in two ways. First, based on 52 biological traits for invertebrates, we conducted a case study to explore functional trends among temperate seasonal wetlands differing in the harshness (i.e., dryness) of their dry season. After finding evidence of trait filtering, we addressed whether it could be generalized across a broader climatic scale. To this end, a meta-analysis (225 seasonal wetlands spread across broad climatic categories: Arid, Temperate, and Cold) allowed us to identify whether an equivalent climate-dependent pattern of trait richness was consistent between the Nearctic and the Western Palearctic. Functional overlap of invertebrates increased from mild (i.e., Temperate) to harsher climates (i.e., Arid and Cold), and phylogenetic clustering (using taxonomy as a surrogate) was highest in Arid and lowest in Temperate wetlands. We show that, (i) as has been described in streams, higher relatedness than would be expected by chance is generally observed in seasonal wetland invertebrate communities; and (ii) this relatedness is not constant but climate-dependent, with the climate under which a given seasonal wetland is located determining the functional overlap and the phylogenetic clustering of the community. Finally, using a space-for-time substitution approach we suggest our results may anticipate how the invertebrate biodiversity embedded in these vulnerable and often overlooked ecosystems will be affected by long-term climate change.

Nestedness and successional trajectories of macroinvertebrate assemblages in man-made wetlands.

Current successional models, primarily those based on floral succession, propose several distinct trajectories based on the integration of two key hypotheses from succession theory: convergence versus divergence in species composition among successional sites, and progression towards versus deviation from a desired reference state. We applied this framework to faunal succession, including differential colonization between active and passive dispersers, and the nested patterns generated as a consequence of this peculiarity. Nine man-made wetlands located in three different areas, from 0-3 years from wetland creation, were assessed. In addition, 91 wetlands distributed throughout the region were used as references for natural macroinvertebrate communities. We predicted the following: (1) highly nested structures in pioneering assemblages will decrease to lower mid-term values due to a shift from active pioneering taxa to passive disperser ones; (2) passive idiosyncratic taxa will elicit divergent successional trajectories among areas; (3) the divergent trajectories will provoke lower local and higher regional diversity values in the mid-term assemblages than in pioneer assemblages. Our results were largely congruent with hypotheses (1) and (2), diverging from the anticipated patterns only in the case of the temporary wetlands area. However, overall diversity trends based on hypothesis (3) did not follow the expected pattern. The divergent successional trajectories did not compensate for regional biodiversity losses that occurred as a consequence of pioneering colonizer decline over time. Consequently, we suggest reconsidering wetland construction for mitigation purposes within mid-term time frames (≤ 3 years). Wetlands may not offset, within this temporal scenario, regional biodiversity loss because the ecosystem may not support idiosyncratic taxa from natural wetlands.