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ABSTRACT Emerging aquatic insects can be an important resource subsidy for a variety of terrestrial consumers, including spiders, birds, bats and lizards. Emergence flux is influenced by a variety of abiotic and biotic variables, such as temperature, drying, and predators and these variables can also control the body size of emergent insects. Despite their importance, these variables can change rapidly during drought conditions as water temperatures rise, surface area decreases and predator densities increase.During 2018, the Konza Prairie Biological Station experienced a record drought: flow ceased in the lower reaches of Kings Creek for the first time in over 40 years of observation, leaving a series of isolated pools. We studied how the drought affected aquatic insect emergence in 12 of these pools via elevated temperatures, decreased surface area, and concentration of predators (e.g. fishes and crayfish) over a four‐week period. We returned in 2020 and sampled emergence in the same pools over 2 weeks under non‐drought conditions to compare emergence between drought and non‐drought conditions.We found three overall patterns: (1) rates of areal emergence abundance and biomass (number or mg DM m⁻²d⁻¹) did not differ between drought and non‐drought conditions. In contrast, pool‐scale emergence abundance, but not biomass (number or mg DM pool⁻¹d⁻¹), was lower during drought conditions; (2) average midge body size was larger during the drought relative to the non‐drought conditions; (3) environmental variables (e.g. temperature, pool surface area, predator biomass) were not predictive of emergence during drought and non‐drought conditions.Fewer, but larger emergent midges (as seen under drought conditions) may represent a higher quality resource for terrestrial consumers than many smaller midges due to increased per‐capita energy yield. However, due to the overall decrease in water availability throughout the stream network, the overall emergence flux was concentrated in reaches with remaining water during the drought, making pools emergence subsidy hotspots. Overall, these contrasting responses underscore the complex nature of community responses to shifting climatic conditions. 

ABSTRACT Freshwater mussels (Bivalvia: Unionida) are among the most imperilled freshwater taxa. Yet, there is a lack of basic life history information for mussels, including data on their growth and longevity. These data help inform conservation efforts, as they can indicate whether species or populations may be vulnerable to decline and inform which species may be best adapted to certain habitats. We aimed to quantify growth and longevity in five mussel species from four river systems in the southeastern United States and test whether growth was related to stream flow. We also interpreted our findings in the context of life history theory.To model mussel growth and longevity, we cut radial thick sections from the shells of mussels and used high‐resolution photography to image the shells. We identified annual growth rings (annuli) and used von Bertalanffy growth models to estimate growth rate (K) and maximum age (A_max) across 13 mussel populations. We then used biochronological methods to remove age‐related variation in annual growth in each shell. We tested whether annual growth was correlated with stream flow using discharge‐based statistics.We found substantial variation inKandA_maxamong species and among populations of the same species.Kwas negatively related toA_max. We did not find consistent correlations between annual growth and stream flow.Our estimates ofKandA_maxalign with previous studies on closely related species and populations. They also match the eco‐evolutionary prediction that growth rate and longevity are negatively related. Life history theory predicts that short‐lived species with higher growth rates should be better adapted to environments with cyclical disturbance regimes, whereas longer‐lived species with low growth rates should be better adapted to stable environments. The lack of correlation between annual growth and stream flow suggests that mussel growth may be limited by other factors in our study system.While some species seem to have relatively narrow ranges for growth and longevity, other species show wide variation among populations. This highlights the need for species‐ and population‐specific conservation efforts. Fundamental life history information can be integrated with other species traits to predict how freshwater taxa may respond to ecological threats. 

Abstract Increases in species richness with habitat area (species–area relationship, or SAR) and increases in ecosystem function with species richness (biodiversity–ecosystem functioning, or BEF) are widely studied ecological patterns. Incorporating functional trait analysis into assemblage datasets may help clarify interpretations of SAR and BEF relationships in natural ecological systems. For example, life history theory can be used to make predictions about what species are most important in generating ecosystem function given a certain set of environmental conditions. We used quantitative assemblage data for freshwater mussels at nine sites in western Alabama, USA, to test for SAR and BEF relationships. At each site, we calculated species richness, mussel assemblage density, and two fundamental metrics of ecosystem function: biomass and secondary production. We also tested whether the proportional biomass and production contributions from species belonging to each of three life history strategies—opportunistic strategistsadapted to unstable or frequently disturbed habitats,periodic strategistsadapted to habitats subject to predictable large‐scale disturbances, andequilibrium strategistsadapted to stable habitats—varied longitudinally with stream drainage area, a proxy for habitat area. Species richness increased with stream size (SAR), and both biomass and production increased with species richness (BEF) and mussel density. There were few longitudinal changes in the proportional contributions of the different life history strategy classifications that we used, but the invasive clamCorbicula flumineacontributed proportionally more biomass and production at sites that had smaller drainage areas. This study provides further evidence for a clear longitudinal SAR in stream‐dwelling taxa. It also suggests BEF relationships for biomass and secondary production in natural assemblages but underscores the importance of assemblage density in BEF studies that use observational field data. Variation in proportional biomass and production contributions by different life history strategies was likely limited by the size of the stream size gradient in our study, as contributions were uniformly high for species with life history traits better adapted to stable and productive habitats such as mid‐sized rivers with low or predictable hydrologic disturbance frequencies. This highlights the need to understand how organisms' functional traits govern their relationships to the environment at different scales. 

Abstract The gut microbiome is influenced by host species and the environment, but how the environment influences the microbiome of animals introduced into a new ecosystem has rarely been investigated. Freshwater mussels are aquatic fauna, with some threatened or endangered species propagated in hatcheries and introduced into natural systems as part of conservation efforts. The effects of the environment on the freshwater mussel gut microbiome were assessed for two hatchery-propagated species (Lampsilis ovata, Lampsilis ornata) introduced into rivers within their natural range. Mussels were placed in rivers for 8 weeks, after which one subset was collected, another subset remained in that river, and a third subset was reciprocally transplanted to another river in the same river basin for a further 8 weeks. Gut microbiome composition and diversity were characterized for all mussels. After the initial 8 weeks, mussels showed increased gut bacterial species richness and distinct community composition compared to hatchery mussels, but gut microbiome diversity then decreased for mussels that remained in the same river for all 16 weeks. The gut bacterial community of mussels transplanted between rivers shifted to resemble that of mussels placed initially into the recipient river and that remained there for the whole study. All mussels showed high proportions of Firmicutes in their gut microbiome after 8 weeks, suggesting an essential role of this phylum in the gut of Lampsilis species. These findings show that the mussel gut microbiome shifts in response to new environments and provide insights into conservation strategies that involve species reintroductions. 

Abstract In freshwater ecosystems, consumers can play large roles in nutrient cycling by modifying nutrient availability for autotrophic and heterotrophic microbes. Nutrients released by consumers directly supportgreen food websbased on primary production andbrown food websbased on decomposition. While much research has focused on impacts of consumer driven nutrient dynamics on green food webs, less attention has been given to studying the effects of these dynamics on brown food webs.Freshwater mussels (Bivalvia: Unionidae) can dominate benthic biomass in aquatic systems as they often occur in dense aggregations that create biogeochemical hotspots that can control ecosystem structure and function through nutrient release. However, despite functional similarities as filter‐feeders, mussels exhibit variation in nutrient excretion and tissue stoichiometry due in part to their phylogenetic origin. Here, we conducted a mesocosm experiment to evaluate how communities of three phylogenetically distinct species of mussels individually and collectively influence components of green and brown food webs.We predicted that the presence of mussels would elicit a positive response in both brown and green food webs by providing nutrients and energy via excretion and biodeposition to autotrophic and heterotrophic microbes. We also predicted that bottom‐up provisioning of nutrients would vary among treatments as a result of stoichiometric differences of species combinations, and that increasing species richness would lead to greater ecosystem functioning through complementarity resulting from greater trait diversity.Our results show that mussels affect the functioning of green and brown food webs through altering nutrient availability for both autotrophic and heterotrophic microbes. These effects are likely to be driven by phylogenetic constraints on tissue nutrient stoichiometry and consequential excretion stoichiometry, which can have functional effects on ecosystem processes. Our study highlights the importance of measuring multiple functional responses across a gradient of diversity in ecologically similar consumers to gain a more holistic view of aquatic food webs. 

Abstract The loose‐equilibrium concept (LEC) predicts that ecological assemblages change transiently but return towards an earlier or average structure. The LEC framework can help determine whether assemblages vary within expected ranges or are permanently altered following environmental change.Long‐lived, slow‐growing animals typically respond slowly to environmental change, and their assemblage dynamics may respond over decades, which transcends most ecological studies. Unionid mussels are valuable for studying dynamics of long‐lived animals because they can live >50 years and occur in dense, species‐rich assemblages (mussel beds). Mussel beds can persist for decades, but disturbance can affect species differently, resulting in variable trajectories according to differences in species composition within and among rivers.We used long‐term data sets (10–40 years) from seven rivers in the eastern United States to evaluate the magnitude, pace and directionality of mussel assemblage change within the context of the LEC.Site trajectories varied within and among streams and showed patterns consistent with either the LEC or directional change. In streams that conformed to the LEC, rank abundance of dominant species remained stable over time, but directional change in other streams was driven by changes in the rank abundance and composition of dominant species.Characteristics of mussel assemblage change varied widely, ranging from those conforming to the LEC to those showing strong directional change. Conservation approaches that attempt to maintain or create a desired assemblage condition should acknowledge this wide range of possible assemblage trajectories and that the environmental factors that influence those changes remain poorly understood. 

Positive biodiversity–ecosystem functioning (BEF) relationships observed in experiments can be challenging to identify in natural communities. Freshwater animal communities are disproportionately harmed by global change that results in accelerated species loss. Understanding how animal-mediated ecosystems functions may change as a result of global change can help determine whether biodiversity or species-specific conservation will be effective at maintaining function. Unionid mussels represent half of imperiled species in freshwater ecosystems globally and perform important ecological functions such as water filtration and nutrient recycling. We explored BEF relationships for 22 naturally assembled mussel aggregations spanning three river basins. We used the Price equation to partition the contributions of species richness, composition, and context dependent interactions to two functions of interests: spatially-explicit standing-stock biomass (indirect proxy for function) and species-specific nitrogen (N) excretion rates (direct measure of N recycling). Random and non-random species loss each reduced biomass and N recycling. Many rare species with low contributions to biomass contributed to standing-stock biomass in all basins. Widespread species had variable function across sites, such that context dependent effects (CDEs) outweighed richness effects on standing-stock biomass in two basins, and were similar to richness effects in the third. Richness effects outweighed CDEs for N recycling. Thus, many species contributed a low proportion to overall N-recycling; a product we attribute to the high evenness and functional effect trait diversity associated with these communities. The loss of low-functioning species reduced the function of persisting species. This novel result using observational data adds evidence that positive species interactions, such as interspecific facilitation, may be a mechanism by which biodiversity enhances ecosystem functions. Our work stresses the importance of evaluating species-specific contributions to functions in diverse systems, such as nutrient cycling when maintaining specific animal-mediated functions is a management goal because indirect proxies may not completely characterize BEF relationships. 

Understanding patterns of diversity across macro (e.g. species-level) and micro (e.g. molecular-level) scales can shed light on community function and stability by elucidating the abiotic and biotic drivers of diversity within ecological communities. We examined the relationships among taxonomic and genetic metrics of diversity in freshwater mussels (Bivalvia: Unionidae), an ecologically important and species-rich group in the southeastern United States. Using quantitative community surveys and reduced-representation genome sequencing across 22 sites in seven rivers and two river basins, we surveyed 68 mussel species and sequenced 23 of these species to characterize intrapopulation genetic variation. We tested for the presence of species diversity–abundance correlations (i.e. the more-individuals hypothesis, MIH), species-genetic diversity correlations (SGDCs) and abundance-genetic diversity correlations (AGDCs) across all sites to evaluate relationships between different metrics of diversity. Sites with greater cumulative multispecies density (a standardized metric of abundance) had a greater number of species, consistent with the MIH hypothesis. Intrapopulation genetic diversity was strongly associated with the density of most species, indicating the presence of AGDCs. However, there was no consistent evidence for SGDCs. Although sites with greater overall densities of mussels had greater species richness, sites with higher genetic diversity did not always exhibit positive correlations with species richness, suggesting that there are spatial and evolutionary scales at which the processes influencing community-level diversity and intraspecific diversity differ. Our work reveals the importance of local abundance as indicator (and possibly a driver) of intrapopulation genetic diversity.