Increasing climate-driven taxonomic homogenization but functional differentiation among river macroinvertebrate assemblages.

Global change is increasing biotic homogenization globally, which modifies the functioning of ecosystems. While tendencies towards taxonomic homogenization in biological communities have been extensively studied, functional homogenization remains an understudied facet of biodiversity. Here, we tested four hypotheses related to long-term changes (1991-2016) in the taxonomic and functional arrangement of freshwater macroinvertebrate assemblages across space and possible drivers of these changes. Using data collected annually at 64 river sites in mainland New Zealand, we related temporal changes in taxonomic and functional spatial ß-diversity, and the contribution of individual sites to ß-diversity, to a set of global, regional, catchment and reach-scale environmental descriptors. We observed long-term, mostly climate-induced, temporal trends towards taxonomic homogenization but functional differentiation among macroinvertebrate assemblages. These changes were mainly driven by replacements of species and functional traits among assemblages, rather than nested species loss. In addition, there was no difference between the mean rate of change in the taxonomic and functional facets of ß-diversity. Climatic processes governed overall population and community changes in these freshwater ecosystems, but were amplified by multiple anthropogenic, topographic and biotic drivers of environmental change, acting widely across the landscape. The functional diversification of communities could potentially provide communities with greater stability, resistance and resilience capacity to environmental change, despite ongoing taxonomic homogenization. Therefore, our study highlights a need to further understand temporal trajectories in both taxonomic and functional components of species communities, which could enable a clearer picture of how biodiversity and ecosystems will respond to future global changes.

Diverging response patterns of terrestrial and aquatic species to hydromorphological restoration.

Although experiences with ecological restoration continue to accumulate, the effectiveness of restoration for biota remains debated. We complemented a traditional taxonomic analysis approach with information on 56 species traits to uncover the responses of 3 aquatic (fish, macroinvertebrates, macrophytes) and 2 terrestrial (carabid beetles, floodplain vegetation) biotic groups to 43 hydromorphological river restoration projects in Germany. All taxonomic groups responded positively to restoration, as shown by increased taxonomic richness (10-164%) and trait diversity (habitat, dispersal and mobility, size, form, life history, and feeding groups) (15-120%). Responses, however, were stronger for terrestrial than aquatic biota, and, contrary to our expectation, taxonomic responses were stronger than those of traits. Nevertheless, trait analysis provided mechanistic insights into the drivers of community change following restoration. Trait analysis for terrestrial biota indicated restoration success was likely enhanced by lateral connectivity and reestablishment of dynamic processes in the floodplain. The weaker response of aquatic biota suggests recovery was hindered by the persistence of stressors in the aquatic environment, such as degraded water quality, dispersal constraints, and insufficient hydromorphological change. Therefore, river restoration requires combined local- and regional-scale approaches to maximize the response of both aquatic and terrestrial organisms. Due to the contrasting responses of aquatic and terrestrial biota, the planning and assessment of river restoration outcomes should consider effects on both components of riverine landscapes.

High mortality and enhanced recovery: modelling the countervailing effects of disturbance on population dynamics.

Disturbances cause high mortality in populations while simultaneously enhancing population growth by improving habitats. These countervailing effects make it difficult to predict population dynamics following disturbance events. To address this challenge, we derived a novel form of the logistic growth equation that permits time-varying carrying capacity and growth rate. We combined this equation with concepts drawn from disturbance ecology to create a general model for population dynamics in disturbance-prone systems. A river flooding example using three insect species (a fast life-cycle mayfly, a slow life-cycle dragonfly and an ostracod) found optimal tradeoffs between disturbance frequency vs. magnitude and a close fit to empirical data in 62% of cases. A savanna fire analysis identified fire frequencies of 3-4 years that maximised population size of a perennial grass. The model shows promise for predicting population dynamics after multiple disturbance events and for management of river flows and fire regimes.

Seasonality and predictability shape temporal species diversity.

Temporal environmental fluctuations, such as seasonality, exert strong controls on biodiversity. While the effects of seasonality are well known, the predictability of fluctuations across years may influence seasonality in ways that are less well understood. The ability of a habitat to support unique, non-nested assemblages of species at different times of the year should depend on both seasonality (occurrence of events at specific periods of the year) and predictability (the reliability of event recurrence) of characteristic ecological conditions. Drawing on tools from wavelet analysis and information theory, we developed a framework for quantifying both seasonality and predictability of habitats, and applied this using global long-term rainfall data. Our analysis predicted that temporal beta diversity should be maximized in highly predictable and highly seasonal climates, and that low degrees of seasonality, predictability, or both would lower diversity in characteristic ways. Using stream invertebrate communities as a case study, we demonstrated that temporal species diversity, as exhibited by community turnover, was determined by a balance between temporal environmental variability (seasonality) and the reliability of this variability (predictability). Communities in highly seasonal mediterranean environments exhibited strong oscillations in community structure, with turnover from one unique community type to another across seasons, whereas communities in aseasonal New Zealand environments fluctuated randomly. Understanding the influence of seasonal and other temporal scales of environmental oscillations on diversity is not complete without a clear understanding of their predictability, and our framework provides tools for examining these trends at a variety of temporal scales, seasonal and beyond. Given the uncertainty of future climates, seasonality and predictability are critical considerations for both basic science and management of ecosystems (e.g., dam operations, bioassessment) spanning gradients of climatic variability.

Linking river flow regimes to riparian plant guilds: a community-wide modeling approach.

Modeling riparian plant dynamics along rivers is complicated by the fact that plants have different edaphic and hydrologic requirements at different life stages. With intensifying human demands for water and continued human alteration of rivers, there is a growing need for predicting responses of vegetation to flow alteration, including responses related to climate change and river flow management. We developed a coupled structured population model that combines stage-specific responses of plant guilds with specific attributes of river hydrologic regime. The model uses information on the vital rates of guilds as they relate to different hydrologic conditions (flood, drought, and baseflow), but deliberately omits biotic interactions from the structure (interaction neutral). Our intent was to (1) consolidate key vital rates concerning plant population dynamics and to incorporate these data into a quantitative framework, (2) determine whether complex plant stand dynamics, including biotic interactions, can be predicted from basic vital rates and river hydrology, and (3) project how altered flow regimes might affect riparian communities. We illustrated the approach using five flow-response guilds that encompass much of the river floodplain community: hydroriparian tree, xeroriparian shrub, hydroriparian shrub, mesoriparian meadow, and desert shrub. We also developed novel network-based tools for predicting community-wide effects of climate-driven shifts and deliberately altered flow regimes. The model recovered known patterns of hydroriparian tree vs. xeroriparian shrub dominance, including the relative proportion of these two guilds as a function of river flow modification. By simulating flow alteration scenarios ranging from increased drought to shifts in flood timing, the model predicted that mature hydroriparian forest should be most abundant near the observed natural flow regime. Multiguild sensitivity analysis identified substantial network connectivity (many connected nodes) and biotic linkage (strong pairwise connections between nodes) under natural flow regime conditions. Both connectivity and linkage were substantially reduced under drought and other flow-alteration scenarios, suggesting that community structure is destabilized under such conditions. This structured population modeling approach provides a useful tool for understanding the community-wide effects of altered flow regimes due to climate change and management actions that influence river flow regime.

Characterizing fish responses to a river restoration over 21 years based on species' traits.

Understanding restoration effectiveness is often impaired by a lack of high-quality, long-term monitoring data and, to date, few researchers have used species' trait information to gain insight into the processes that drive the reaction of fish communities to restoration. We examined fish-community responses with a highly resolved data set from 21 consecutive years of electrofishing (4 years prerestoration and 17 years postrestoration) at multiple restored and unrestored reaches from a river restoration project on the Lippe River, Germany. Fish abundance peaked in the third year after the restoration; abundance was 6 times higher than before the restoration. After 5-7 years, species richness and abundance stabilized at 2 and 3.5 times higher levels relative to the prerestoration level, respectively. However, interannual variability of species richness and abundance remained considerable, illustrating the challenge of reliably assessing restoration outcomes based on data from individual samplings, especially in the first years following restoration. Life-history and reproduction-related traits best explained differences in species' responses to restoration. Opportunistic short-lived species with early female maturity and multiple spawning runs per year exhibited the strongest increase in abundance, which reflected their ability to rapidly colonize new habitats. These often small-bodied and fusiform fishes typically live in dynamic and ephemeral instream and floodplain areas that river-habitat restorations often aim to create, and in this case their increases in abundance indicated successful restoration. Our results suggest that a greater consideration of species' traits may enhance the causal understanding of community processes and the coupling of restoration to functional ecology. Trait-based assessments of restoration outcomes would furthermore allow for easier transfer of knowledge across biogeographic borders than studies based on taxonomy.

Species interactions drive continuous assembly of freshwater communities in stochastic environments.

Understanding the factors driving the maintenance of long-term biodiversity in changing environments is essential for improving restoration and sustainability strategies in the face of global environmental change. Biodiversity is shaped by both niche and stochastic processes, however the strength of deterministic processes in unpredictable environmental regimes is highly debated. Since communities continuously change over time and space-species persist, disappear or (re)appear-understanding the drivers of species gains and losses from communities should inform us about whether niche or stochastic processes dominate community dynamics. Applying a nonparametric causal discovery approach to a 30-year time series containing annual abundances of benthic invertebrates across 66 locations in New Zealand rivers, we found a strong negative causal relationship between species gains and losses directly driven by predation indicating that niche processes dominate community dynamics. Despite the unpredictable nature of these system, environmental noise was only indirectly related to species gains and losses through altering life history trait distribution. Using a stochastic birth-death framework, we demonstrate that the negative relationship between species gains and losses can not emerge without strong niche processes. Our results showed that even in systems that are dominated by unpredictable environmental variability, species interactions drive continuous community assembly.

Ecosystem-size relationships of river populations and communities.

Knowledge of ecosystem-size influences on river populations and communities is integral to the balancing of human and environmental needs for water. The multiple dimensions of dendritic river networks complicate understanding of ecosystem-size influences, but could be resolved by the development of scaling relationships. We highlight the importance of physical constraints limiting predator body sizes, movements, and population sizes in small rivers, and where river contraction limits space or creates stressful conditions affecting community stability and food webs. Investigations of the scaling and contingency of these processes will be insightful because of the underlying generality and scale independence of such relationships. Doing so will also pinpoint damaging water-management practices and identify which aspects of river size can be most usefully manipulated in river restoration.

Climate change impacts on Aotearoa New Zealand: a horizon scan approach.

Many of the implications of climate change for Aotearoa (New Zealand) remain unclear. To identify so-far unseen or understudied threats and opportunities related to climate change we applied a horizon-scanning process. First, we collated 171 threats and opportunities across our diverse fields of research. We then scored each item for novelty and potential impact and finally reduced the list to ten threats and ten opportunities through a prioritisation process. Within the 20 items presented in this paper, we uncover a range of climate-related costs and benefits. Unexpected opportunities evolve from economic reorganisation and changes to perspectives. The threats we highlight include the overall failure to interconnect siloed policy responses, as well as those relating to extreme events and feedbacks, as well as pressures that undermine the coherence of society. A major theme of our work is that climate change effects in Aotearoa are likely to transgress the boundaries of research disciplines, industry sectors and policy systems, emphasising the importance of developing transdisciplinary methods and approaches. We use this insight to connect potential responses to climate change with Aotearoa's culture and geography.

Matrix community models for ecology and evolution.

Ecological communities are shaped by biotic interactions as well as environmental forces, and both must be incorporated to obtain models capable of forecasting realistic community dynamics. Many community models first specify pairwise biotic interactions and then secondarily examine how extrinsic factors such as abiotic conditions affect species abundances. A disadvantage of this approach is that the species interactions themselves are often environment and context specific, making parameterization difficult. We propose an alternative approach, matrix community models (MCMs), which are sets of matrix population models linked by an assumption of aggregate density dependence. MCMs incorporate detailed species autecology but are neutral with respect to pairwise species interactions, instead allowing interactions to be revealed within the model structure. These model-revealed species interactions, including competitive exclusion, facilitation, and interference competition, shape the distribution and abundance of species within communities and generate empirically testable predictions about species interactions. We develop a framework for building MCMs using vital rates in a stochastic, multispecies framework. Single-species matrix population models are connected via an assumption of aggregate density dependence, pairwise species interactions are estimated with sensitivity analysis, and community trajectories are analyzed under different environmental regimes using standard statistical tools and network analysis. MCMs have the advantage that pairwise species interactions need not be specified a priori, and that mechanistic demographic-environment linkages permit forecasting of community dynamics under novel, non-stationary environmental regimes. A challenge is that species' autecological vital rates, such as fecundity, growth and survivorship, must be measured under a diverse range of environmental conditions to parameterize the models. We illustrate the approach with examples and discuss prospects for future theoretical and empirical developments.

Forecasting the future of life in Antarctica.

Antarctic ecosystems are under increasing anthropogenic pressure, but efforts to predict the responses of Antarctic biodiversity to environmental change are hindered by considerable data challenges. Here, we illustrate how novel data capture technologies provide exciting opportunities to sample Antarctic biodiversity at wider spatiotemporal scales. Data integration frameworks, such as point process and hierarchical models, can mitigate weaknesses in individual data sets, improving confidence in their predictions. Increasing process knowledge in models is imperative to achieving improved forecasts of Antarctic biodiversity, which can be attained for data-limited species using hybrid modelling frameworks. Leveraging these state-of-the-art tools will help to overcome many of the data scarcity challenges presented by the remoteness of Antarctica, enabling more robust forecasts both near- and long-term.

Taxonomic and functional reorganization in Central European stream macroinvertebrate communities over 25 years.

Climate warming can lead to a replacement of species that favour cold temperatures by species that favour warm temperatures. However, the implications of such thermic shifts for the functioning of ecosystems remain poorly understood. Here, we used stream macroinvertebrate biological and ecological traits to quantify the relative contribution of cold, intermediate and warm temperature-adapted taxa to changes in community functional diversity (FD) using a dataset of 3781 samples collected in Central Europe over 25 years, from 1990 to 2014. Our analyses indicated that functional diversity of stream macroinvertebrate communities increased over the study period. This gain was driven by a net 39 % increase in the richness of taxa that favour intermediate temperatures, which comprise the highest share in the community, and to a 97 % increase in the richness of taxa that favour warm temperatures. These warm temperature-adapted taxa displayed a distinct and more diverse suite of functional traits compared to the cold temperature-adapted group and thus contributed disproportionately to local FD on a per-taxon basis. At the same time, taxonomic beta-diversity declined significantly within each thermal group, in association with increasing local taxon richness. This study shows that over recent decades, small low-mountain streams in Central Europe have experienced a process of thermophilization and increasing functional diversity at local scales. However, a progressive homogenisation occurred at the regional scale, with communities converging towards similar taxonomic composition. As the reported increase in local functional diversity can be attributed mostly to the intermediate temperature-adapted taxa and a few expanding warm temperature-adapted taxa, these patterns could mask more subtle loss of sensitive cold temperature-adapted taxa with irreplaceable functional traits. In light of increasing climate warming, preservation of cold habitat refuges, should be considered a priority in river conservation.

Diversity-stability relationships across organism groups and ecosystem types become decoupled across spatial scales.

The relationship between biodiversity and stability, or its inverse, temporal variability, is multidimensional and complex. Temporal variability in aggregate properties, like total biomass or abundance, is typically lower in communities with higher species diversity (i.e., the diversity-stability relationship [DSR]). At broader spatial extents, regional-scale aggregate variability is also lower with higher regional diversity (in plant systems) and with lower spatial synchrony. However, focusing exclusively on aggregate properties of communities may overlook potentially destabilizing compositional shifts. It is not yet clear how diversity is related to different components of variability across spatial scales, nor whether regional DSRs emerge across a broad range of organisms and ecosystem types. To test these questions, we compiled a large collection of long-term metacommunity data spanning a wide range of taxonomic groups (e.g., birds, fish, plants, invertebrates) and ecosystem types (e.g., deserts, forests, oceans). We applied a newly developed quantitative framework for jointly analyzing aggregate and compositional variability across scales. We quantified DSRs for composition and aggregate variability in local communities and metacommunities. At the local scale, more diverse communities were less variable, but this effect was stronger for aggregate than compositional properties. We found no stabilizing effect of γ-diversity on metacommunity variability, but ß-diversity played a strong role in reducing compositional spatial synchrony, which reduced regional variability. Spatial synchrony differed among taxa, suggesting differences in stabilization by spatial processes. However, metacommunity variability was more strongly driven by local variability than by spatial synchrony. Across a broader range of taxa, our results suggest that high γ-diversity does not consistently stabilize aggregate properties at regional scales without sufficient spatial ß-diversity to reduce spatial synchrony.

Distance decay of benthic macroinvertebrate communities in a mountain river network: Do dispersal routes and dispersal ability matter?

Environmental heterogeneity and dispersal limitation are important drivers of beta diversity; however, their relative influence on the two fundamental components of beta diversity (i.e., species replacement and richness difference) has not been fully examined in montane streams. Here, we examined the relative importance of local environmental gradients and three physical distance matrices (i.e., overland, watercourse and cost distances) on beta diversity and its two components for a macroinvertebrate metacommunity in a stream network. To provide additional insights into community assembly, we also analysed variation in two deconstructed sub-communities based on dispersal ability (i.e., weak and strong dispersers). Both environmental filters and physical distances (dispersal limitation) drove patterns of overall beta diversity, with the former generally prevailing over the latter. Species replacement components showed stronger correlations with environmental gradients than physical distances, while the opposite is true for the richness difference components. Overland distances were generally more important than cost and watercourse distances for community dissimilarity of stream macroinvertebrates, implying that lateral dispersal out of stream corridors through flight was the major dispersal route in the studied steam network. As expected, community dissimilarity of strong dispersers was primarily shaped by environmental filtering, while community dissimilarity of weak dispersers was associated with the joint effects of environmental filtering and dispersal limitation. Our findings demonstrate that partitioning overall dissimilarity into species replacement and richness difference provides more insights into the processes driving spatial variability in biological communities compared with the utilization of total beta diversity alone. Our results support the notion that maintaining environmental heterogeneity and natural connectivity of stream networks should be effective measures to conserve regional biodiversity.

Physiology, niche characteristics and extreme events: Current and future habitat suitability of a rhodolith-forming species in the Southwestern Atlantic.

Given the ecological and biogeochemical importance of rhodolith beds, it is necessary to investigate how future environmental conditions will affect these organisms. We investigated the impacts of increased nutrient concentrations, acidification, and marine heatwaves on the performance of the rhodolith-forming species Lithothamnion crispatum in a short-term experiment, including the recovery of individuals after stressor removal. Furthermore, we developed an ecological niche model to establish which environmental conditions determine its current distribution along the Brazilian coast and to project responses to future climate scenarios. Although L. crispatum suffered a reduction in photosynthetic performance when exposed to stressors, they returned to pre-experiment values following the return of individuals to control conditions. The model showed that the most important variables in explaining the current distribution of L. crispatum on the Brazilian coast were maximum nitrate and temperature. In future ocean conditions, the model predicted a range expansion of habitat suitability for this species of approximately 58.5% under RCP 8.5. Physiological responses to experimental future environmental conditions corroborated model predictions of the expansion of this species' habitat suitability in the future. This study, therefore, demonstrates the benefits of applying combined approaches to examine potential species responses to climate-change drivers from multiple angles.

The application of metacommunity theory to the management of riverine ecosystems.

River managers strive to use the best available science to sustain biodiversity and ecosystem function. To achieve this goal requires consideration of processes at different scales. Metacommunity theory describes how multiple species from different communities potentially interact with local-scale environmental drivers to influence population dynamics and community structure. However, this body of knowledge has only rarely been used to inform management practices for river ecosystems. In this paper, we present a conceptual model outlining how the metacommunity processes of local niche sorting and dispersal can influence the outcomes of management interventions and provide a series of specific recommendations for applying these ideas as well as research needs. In all cases, we identify situations where traditional approaches to riverine management could be enhanced by incorporating an understanding of metacommunity dynamics. A common theme is developing guidelines for assessing the metacommunity context of a site or region, evaluating how that context may affect the desired outcome, and incorporating that understanding into the planning process and methods used. To maximize the effectiveness of management activities, scientists and resource managers should update the toolbox of approaches to riverine management to reflect theoretical advances in metacommunity ecology.

Moderate warming over the past 25 years has already reorganized stream invertebrate communities.

Climate warming often results in species range shifts, biodiversity loss and accumulated climatic debts of biota (i.e. slower changes in biota than in temperature). Here, we analyzed the changes in community composition and temperature signature of stream invertebrate communities over 25 years (1990-2014), based on a large set of samples (n = 3782) over large elevation, latitudinal and longitudinal gradients in central Europe. Although warming was moderate (average 0.5 °C), we found a strong reorganization of stream invertebrate communities. Total abundance (+35.9%) and richness (+39.2%) significantly increased. The share of abundance (TA) and taxonomic richness (TR) of warm-dwelling taxa (TA: +73.2%; TR: +60.2%) and medium-temperature-dwelling taxa (TA: +0.4%; TR: +5.8%) increased too, while cold-dwelling taxa declined (TA: -61.5%; TR: -47.3%). The community temperature index, representing the temperature signature of stream invertebrate communities, increased at a similar pace to physical temperature, indicating a thermophilization of the communities and, for the first time, no climatic debt. The strongest changes occurred along the altitudinal gradient, suggesting that stream invertebrates use the spatial configuration of river networks to track their temperature niche uphill. Yet, this may soon come to an end due to the summit trap effect. Our results indicate an ongoing process of replacement of cold-adapted species by thermophilic species at only 0.5 °C warming, which is particularly alarming in the light of the more drastic climate warming projected for coming decades.

Urbanization Impacts the Physicochemical Characteristics and Abundance of Fecal Markers and Bacterial Pathogens in Surface Water.

Urbanization is increasing worldwide and is happening at a rapid rate in China in line with economic development. Urbanization can lead to major changes in freshwater environments through multiple chemical and microbial contaminants. We assessed the impact of urbanization on physicochemical characteristics and microbial loading in canals in Suzhou, a city that has experienced rapid urbanization in recent decades. Nine sampling locations covering three urban intensity classes (high, medium and low) in Suzhou were selected for field studies and three locations in Huangshan (natural reserve) were included as pristine control locations. Water samples were collected for physicochemical, microbiological and molecular analyses. Compared to medium and low urbanization sites, there were statistically significant higher levels of nutrients and total and thermotolerant coliforms (or fecal coliforms) in highly urbanized locations. The effect of urbanization was also apparent in the abundances of human-associated fecal markers and bacterial pathogens in water samples from highly urbanized locations. These results correlated well with land use types and anthropogenic activities at the sampling sites. The overall results indicate that urbanization negatively impacts water quality, providing high levels of nutrients and a microbial load that includes fecal markers and pathogens.

Reintroduction of freshwater macroinvertebrates: challenges and opportunities.

Species reintroductions - the translocation of individuals to areas in which a species has been extirpated with the aim of re-establishing a self-sustaining population - have become a widespread practice in conservation biology. Reintroduction projects have tended to focus on terrestrial vertebrates and, to a lesser extent, fishes. Much less effort has been devoted to the reintroduction of invertebrates into restored freshwater habitats. Yet, reintroductions may improve restoration outcomes in regions where impoverished regional species pools limit the self-recolonisation of restored freshwaters. We review the available literature on macroinvertebrate reintroductions, focusing on identifying the intrinsic and extrinsic factors that determine their success or failure. Our study reveals that freshwater macroinvertebrate reintroductions remain rare, are often published in the grey literature and, of the attempts made, approximately one-third fail. We identify life-cycle complexity and remaining stressors as the two factors most likely to affect reintroduction success, illustrating the unique challenges of freshwater macroinvertebrate reintroductions. Consideration of these factors by managers during the planning process and proper documentation - even if a project fails - may increase the likelihood of successful outcomes in future reintroduction attempts of freshwater macroinvertebrates.