Structural and functional responses of plant communities to climate change-mediated alterations in the hydrology of riparian areas in temperate Europe.

The hydrology of riparian areas changes rapidly these years because of climate change-mediated alterations in precipitation patterns. In this study, we used a large-scale in situ experimental approach to explore effects of drought and flooding on plant taxonomic diversity and functional trait composition in riparian areas in temperate Europe. We found significant effects of flooding and drought in all study areas, the effects being most pronounced under flooded conditions. In near-stream areas, taxonomic diversity initially declined in response to both drought and flooding (although not significantly so in all years) and remained stable under drought conditions, whereas the decline continued under flooded conditions. For most traits, we found clear indications that the functional diversity also declined under flooded conditions, particularly in near-stream areas, indicating that fewer strategies succeeded under flooded conditions. Consistent changes in community mean trait values were also identified, but fewer than expected. This can have several, not mutually exclusive, explanations. First, different adaptive strategies may coexist in a community. Second, intraspecific variability was not considered for any of the traits. For example, many species can elongate shoots and petioles that enable them to survive shallow, prolonged flooding but such abilities will not be captured when applying mean trait values. Third, we only followed the communities for 3 years. Flooding excludes species intolerant of the altered hydrology, whereas the establishment of new species relies on time-dependent processes, for instance the dispersal and establishment of species within the areas. We expect that altered precipitation patterns will have profound consequences for riparian vegetation in temperate Europe. Riparian areas will experience loss of taxonomic and functional diversity and, over time, possibly also alterations in community trait responses that may have cascading effects on ecosystem functioning.

Functional trait composition of aquatic plants can serve to disentangle multiple interacting stressors in lowland streams.

Historically, close attention has been paid to negative impacts associated with nutrient loads to streams and rivers, but today hydromorphological alterations are considered increasingly implicated when lowland streams do not achieve good ecological status. Here, we explore if trait-abundance patterns of aquatic plants change along gradients in hydromorphological degradation and eutrophication in lowland stream sites located in Denmark. Specifically, we hypothesised that: i) changes in trait-abundance patterns occur along gradients in hydromorphological degradation and ii) trait-abundance patterns can serve to disentangle effects of eutrophication and hydromorphological degradation in lowland streams reflecting that the mechanisms behind changes differ. We used monitoring data from a total of 147 stream reaches with combined data on aquatic plant species abundance, catchment land use, hydromorphological alterations (i.e. planform, cross section, weed cutting) and water chemistry parameters. Traits related to life form, dispersal, reproduction and survival together with ecological preference values for nutrients and light (Ellenberg N and L) were allocated to 41 species representing 79% of the total species pool. We found clear evidence that habitat degradation (hydromorphological alterations and eutrophication) mediated selective changes in the trait-abundance patterns of the plant community. Specific traits could distinguish hydromorphological degradation (free-floating, surface; anchored floating leaves; anchored heterophylly) from eutrophication (free-floating, submerged; leaf area). We provide a conceptual framework for interpretation of how eutrophication and hydromorphological degradation interact and how this is reflected in trait-abundance patterns in aquatic plant communities in lowland streams. Our findings support the merit of trait-based approaches in biomonitoring as they shed light on mechanisms controlling structural changes under environmental stress. The ability to disentangle several stressors is particularly important in lowland stream environments where several stressors act in concert since the impact of the most important stressor can be targeted first, which is essential to improve the ecological status.

A comparative analysis reveals weak relationships between ecological factors and beta diversity of stream insect metacommunities at two spatial levels.

The hypotheses that beta diversity should increase with decreasing latitude and increase with spatial extent of a region have rarely been tested based on a comparative analysis of multiple datasets, and no such study has focused on stream insects. We first assessed how well variability in beta diversity of stream insect metacommunities is predicted by insect group, latitude, spatial extent, altitudinal range, and dataset properties across multiple drainage basins throughout the world. Second, we assessed the relative roles of environmental and spatial factors in driving variation in assemblage composition within each drainage basin. Our analyses were based on a dataset of 95 stream insect metacommunities from 31 drainage basins distributed around the world. We used dissimilarity-based indices to quantify beta diversity for each metacommunity and, subsequently, regressed beta diversity on insect group, latitude, spatial extent, altitudinal range, and dataset properties (e.g., number of sites and percentage of presences). Within each metacommunity, we used a combination of spatial eigenfunction analyses and partial redundancy analysis to partition variation in assemblage structure into environmental, shared, spatial, and unexplained fractions. We found that dataset properties were more important predictors of beta diversity than ecological and geographical factors across multiple drainage basins. In the within-basin analyses, environmental and spatial variables were generally poor predictors of variation in assemblage composition. Our results revealed deviation from general biodiversity patterns because beta diversity did not show the expected decreasing trend with latitude. Our results also call for reconsideration of just how predictable stream assemblages are along ecological gradients, with implications for environmental assessment and conservation decisions. Our findings may also be applicable to other dynamic systems where predictability is low.

Plant trait characteristics vary with size and eutrophication in European lowland streams.

Previous studies investigating community-level relationships between plant functional trait characteristics and stream environmental characteristics remain scarce. Here, we used community-weighted means to identify how plant traits link to lowland stream typology and how agricultural intensity in the catchment affects trait composition.We analysed plant trait characteristics in 772 European lowland streams to test the following two hypotheses: (i) trait characteristics differ between plant communities in small and medium-sized streams, reflecting adaptations to different habitat characteristics, and (ii) trait characteristics vary with the intensity of agricultural land use in the stream catchment, mediated either directly by an increase in productive species or indirectly by an increase in species that efficiently intercept and utilize light.We found that the communities in small streams were characterized by a higher abundance of light-demanding species growing from single apical meristems, reproducing by seeds and rooted to the bottom with floating and/or heterophyllous leaves, whereas the community in medium-sized streams was characterized by a higher abundance of productive species growing from multi-apical and basal growth meristems forming large canopies.We also found indications that community trait characteristics were affected by eutrophication. We did not find enhanced abundance of productive species with an increasing proportion of agriculture in the catchments. Instead, we found an increase in the abundance of species growing from apical and multi-apical growth meristems as well as in the abundance of species tolerant of low light availability. The increase in the abundance of species possessing these traits likely reflects different strategies to obtain greater efficiency in light interception and utilization in nutrient-enriched environments. Synthesis and applications. Our findings challenge the general assumption of the EU Water Framework Directive compliant assessment systems that plant community patterns in streams reflect the nutrient preference of the community. Instead, light availability and the ability to improve interception and utilization appeared to be of key importance for community composition in agricultural lowland streams. We therefore suggest moving from existing approaches building on species-specific preference values for nutrients to determine the level of nutrient impairment to trait-based approaches that provide insight into the biological mechanisms underlying the changes. We recommend that existing systems are critically appraised in the context of the findings of this study.

The influence of environmental, biotic and spatial factors on diatom metacommunity structure in Swedish headwater streams.

Stream assemblages are structured by a combination of local (environmental filtering and biotic interactions) and regional factors (e.g., dispersal related processes). The relative importance of environmental and spatial (i.e., regional) factors structuring stream assemblages has been frequently assessed in previous large-scale studies, but biotic predictors (potentially reflecting local biotic interactions) have rarely been included. Diatoms may be useful for studying the effect of trophic interactions on community structure since: (1) a majority of experimental studies shows significant grazing effects on diatom species composition, and (2) assemblages can be divided into guilds that have different susceptibility to grazing. We used a dataset from boreal headwater streams in south-central Sweden (covering a spatial extent of ∼14000 km(2)), which included information about diatom taxonomic composition, abundance of invertebrate grazers (biotic factor), environmental (physicochemical) and spatial factors (obtained through spatial eigenfunction analyses). We assessed the relative importance of environmental, biotic, and spatial factors structuring diatom assemblages, and performed separate analyses on different diatom guilds. Our results showed that the diatom assemblages were mainly structured by environmental factors. However, unique spatial and biological gradients, specific to different guilds and unrelated to each other, were also evident. We conclude that biological predictors, in combination with environmental and spatial variables, can reveal a more complete picture of the local vs. regional control of species assemblages in lotic environments. Biotic factors should therefore not be overlooked in applied research since they can capture additional local control and therefore increase accuracy and performance of predictive models. The inclusion of biotic predictors did, however, not significantly influence the unique fraction explained by spatial factors, which suggests low bias in previous assessments of unique regional control of stream assemblages.

Hierarchical dynamics of ecological communities: do scales of space and time match?

Theory posits that community dynamics organize at distinct hierarchical scales of space and time, and that the spatial and temporal patterns at each scale are commensurate. Here we use time series modeling to investigate fluctuation frequencies of species groups within invertebrate metacommunities in 26 boreal lakes over a 20-year period, and variance partitioning analysis to study whether species groups with different fluctuation patterns show spatial signals that are commensurate with the scale-specific fluctuation patterns identified. We identified two groups of invertebrates representing hierarchically organized temporal dynamics: one species group showed temporal variability at decadal scales (slow patterns of change), whilst another group showed fluctuations at 3 to 5-year intervals (faster change). This pattern was consistently found across all lakes studied. A spatial signal was evident in the slow but not faster-changing species groups. As expected, the spatial signal for the slow-changing group coincided with broad-scale spatial patterns that could be explained with historical biogeography (ecoregion delineation, and dispersal limitation assessed through a dispersal trait analysis). In addition to spatial factors, the slow-changing groups correlated with environmental variables, supporting the conjecture that boreal lakes are undergoing environmental change. Taken together our results suggest that regionally distinct sets of taxa, separated by biogeographical boundaries, responded similarly to broad-scale environmental change. Not only does our approach allow testing theory about hierarchically structured space-time patterns; more generally, it allows assessing the relative role of the ability of communities to track environmental change and dispersal constraints limiting community structure and biodiversity at macroecological scales.

Metacommunity structure in a small boreal stream network.

Current ecological frameworks emphasize the relative importance of local and regional drivers for structuring species communities. However, most research has been carried out in systems with discrete habitat boundaries and a clear insular structure. Stream networks deviate from the insular structure and can serve as excellent model systems for studying hierarchical community dynamics over different temporal and spatial extents. We used benthic invertebrate data from streams in a small northern Swedish catchment to test whether metacommunity dynamics change between seasons, across spatial hierarchies (i.e. at the whole catchment scale vs. the scales of first-order and second/third-order sites within the catchment) and between stream-order groups. We assessed metacommunity structure as a function of three relevant dispersal dimensions (directional downstream processes, along-stream dispersal and overland dispersal). These dispersal dimensions were related to species groups with relevant dispersal traits (flying capacity, drift propensity) and dispersal capacities (weak vs. strong) to elucidate whether the observed spatial signals were due to dispersal limitation or mass effects. Results showed complex community organization that varied between seasons, with the scale of observation, and with stream order. The importance of spatial factors and specific dispersal dimensions was highly dependent on the time of sampling and the scale of observation. The importance of environmental factors was more consistent in our analyses, but their effect on species community structure peaked at first-order sites. Our analyses of species dispersal traits were not unequivocal, but indicated that both mass effects and dispersal limitation could simultaneously contribute to the spatial signal at the scale of the whole catchment through different dispersal pathways. We conclude that the study of hierarchically organized ecosystems uncovers complex patterns of metacommunity organization that may deviate substantially from those of systems with insular structure and discrete habitat boundaries. Moreover, we show that dispersal constraints imposed by the dendritic structure of stream networks and distinct dispersal mechanisms (e.g. dispersal limitation) may be evident also at very small spatial extents. Thus, even at this small scale, a landscape management approach that takes the dendritic nature of stream networks into account is needed to effectively conserve stream biodiversity.