Coextirpation of host–affiliate relationships in rivers: the role of climate change,water withdrawal,and host‐specificity

The role of climate‐related disturbances on complex host–affiliate relationships remains understudied, largely because affiliate species vary in host use and are often differentially susceptible to disturbance relative to their hosts. Here we report the first set of host–affiliate species–discharge relationships (SDR) in freshwater and examine how anticipated shifts in water availability (flow) will impact coextirpations. We used SDR for freshwater mussels and fish across 11 regions (over 350 rivers) in the continental United States that we coupled to future water availability (2070) to model mussel and fish coextirpations. We also used river‐specific host–affiliate matrices (presence–absence) to evaluate how host‐specificity (mean number of hosts used by an affiliate) and host‐overlap (extent to which affiliates share hosts) relate to extirpation vulnerability. We found that the strength and predictability of SDR models vary geographically and that mussels were more susceptible to flow alterations than fish. These patterns of extirpations were strongest in the southeast where: (1) flow reductions are expected to be greatest; (2) more species are lost per unit flow; (3) and more mussels are expected to be lost per unit of fish. We also found that overall mussel losses associated with reduction in habitat (water availability) were greater than those associated with loss of fish hosts which we assumed to be a function of host redundancy. These findings highlight the utility of SDR as a tool for conservation efforts but they also demonstrate the potential severity of reductions in mussel and fish richness as consequence of climate change and water use. Mussels provide key ecosystem services but face multiple pronged attacks from reductions in flow, habitat, and fish hosts. These losses in biodiversity and ecosystem functions can translate into major effects on food webs and nutrient recycling.     

Ecosystem Processes Performed by Unionid Mussels in Stream Mesocosms: Species Roles and Effects of Abundance

Unionid mussels are a guild of freshwater, sedentary filter-feeders experiencing a global decline in both species richness and abundance. To predict how these losses may impact stream ecosystems we need to quantify the effects of both overall mussel biomass and individual species on ecosystem processes. In this study we begin addressing these fundamental questions by comparing rates of ecosystem processes for two common mussel species, Amblema plicata and Actinonaias ligamentina, across a range of abundance levels and at two trophic states (low and high productivity) in stream mesocosms. At both low and high productivity, community respiration, water column ammonia, nitrate, and phosphorus concentrations, and algal clearance rates were all linearly related to overall mussel biomass. After removing the effects of biomass with ANCOVA, we found few differences between species. In a separate series of experiments, nutrient excretion (phosphorus, ammonia, and molar N:P) and biodeposition rates were only marginally different between species. For the species studied here, functional effects of unionids in streams were similar between species and linearly related to biomass, indicating the potential for strong effects when overall mussel biomass is high and hydrologic residence times are long.     

Consumer Aggregations Drive Nutrient Dynamics and Ecosystem Metabolism in Nutrient-Limited Systems

Differences in animal distributions and metabolic demands can influence energy and nutrient flow in an ecosystem. Through taxa-specific nutrient consumption, storage, and remineralization, animals may influence energy and nutrient pathways in an ecosystem. Here we show these taxa-specific traits can drive biogeochemical cycles of nutrients and alter ecosystem primary production and metabolism, using riverine systems that support heterogeneous freshwater mussel aggregations. Freshwater unionid mussels occur as distinct, spatially heterogeneous, dense aggregations in rivers. They may influence rates of production and respiration because their activities are spatially concentrated within given stream reaches. Previous work indicates that mussels influence nutrient limitation patterns, algal species composition, and producer and primary consumer biomass. Here, we integrate measures of organismal rates, stoichiometry, community-scaled rates, and ecosystem rates, to determine the relative source–sink nutrient dynamics of mussel aggregations and their influence on net ecosystem processes. We studied areas with and without mussel aggregations in three nitrogen-limited rivers in southeastern Oklahoma, USA. We measured respiration and excretion rates of mussels and collected a subset of samples for tissue chemistry and for thin sectioning of the shell to determine growth rates at each site. This allowed us to assess nutrient remineralization and nutrient sequestration by mussels. These rates were scaled to the community. We also measured stream metabolism at three sites with and without mussels. We demonstrated that mussel species have distinct stoichiometric traits, vary in their respiration rates, and that mussel aggregations influence nutrient cycling and productivity. Across all mussel aggregations, we found that mussels excreted more nitrogen than they sequestered into tissue and excreted more phosphorus than they sequestered except at one site. Furthermore, gross primary productivity was significantly greater at reaches with mussels. Collectively, our results indicate that mussels have ecosystem-level impacts on nutrient availability and production in nutrient-limited rivers. Within these streams, mussels are affecting the movement of nutrients and altering nutrient spiralling.     

Invasive mussels modify the cycling,storage and distribution of nutrients and carbon in a large lake

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Freshwater mussel conservation: A global horizon scan of emerging threats and opportunities

We identified 14 emerging and poorly understood threats and opportunities for addressing the global conservation of freshwater mussels over the next decade. A panel of 17 researchers and stakeholders from six continents submitted a total of 56 topics that were ranked and prioritized using a consensus-building Delphi technique. Our 14 priority topics fell into five broad themes (autecology, population dynamics, global stressors, global diversity, and ecosystem services) and included understanding diets throughout mussel life history; identifying the drivers of population declines; defining metrics for quantifying mussel health; assessing the role of predators, parasites, and disease; informed guidance on the risks and opportunities for captive breeding and translocations; the loss of mussel–fish co-evolutionary relationships; assessing the effects of increasing surface water changes; understanding the effects of sand and aggregate mining; understanding the effects of drug pollution and other emerging contaminants such as nanomaterials; appreciating the threats and opportunities arising from river restoration; conserving understudied hotspots by building local capacity through the principles of decolonization; identifying appropriate taxonomic units for conservation; improved quantification of the ecosystem services provided by mussels; and understanding how many mussels are enough to provide these services. Solutions for addressing the topics ranged from ecological studies to technological advances and socio-political engagement. Prioritization of our topics can help to drive a proactive approach to the conservation of this declining group which provides a multitude of important ecosystem services.     

Soluble nitrogen and phosphorus excretion of exotic freshwater mussels (Dreissena spp.): potential impacts for nutrient remineralisation in western Lake Erie

1. Recent increases in phytoplankton biomass and the recurrence of cyanobacterial blooms in western Lake Erie, concomitant with a shift from a community dominated by zebra mussels (Dreissena polymorpha) to one dominated by quagga mussels (D. bugensis), led us to test for differences in ammonia‐nitrogen and phosphate‐phosphorus excretion rates of these two species of invasive molluscs. 2. We found significant differences in excretion rate both between size classes within a taxon and between taxa, with zebra mussels generally having greater nutrient excretion rates than quagga mussels. Combining measured excretion rates with measurements of mussel soft‐tissue dry weight and shell length, we developed nutrient excretion equations allowing estimation of nutrient excretion by dreissenids. 3. Comparing dreissenid ammonia and phosphate excretion with that of the crustacean zooplankton, we demonstrated that the mussels add to nitrogen and phosphorus remineralisation, shortening nitrogen and phosphorus turnover times, and, importantly, modify the nitrogen and phosphorus cycles in Lake Erie. The increased nutrient flux from dreissenids may facilitate phytoplankton growth and cyanobacterial blooms in well‐mixed and/or shallow areas of western Lake Erie.     

Long‐term decline of native freshwater mussel assemblages in a federally protected river

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Alteration of Ecosystem Function by Zebra Mussels in Oneida Lake: Impacts on Submerged Macrophytes

Dreissenid mussels (the zebra mussel Dreissena polymorpha and the quagga mussel D. bugensis) are ecosystem engineers that modify the physical environment by increasing light penetration. Such a change is likely to affect the distribution and diversity of submerged macrophytes. Filter-feeding by these mussels has been associated with increased water clarity in many North American and European lakes. In this study, we report the increase in water clarity of Oneida Lake, New York, USA, for 1975–2002 and argue that the increase was caused by zebra mussel invasion rather than declines in nutrients. Over the study period, although mean total phosphorus decreased significantly, the main increase in water clarity occurred after the zebra mussel invasion in 1991. The average depth receiving 1% surface light increased from 6.7 m to 7.8 m after the invasion of zebra mussels, representing a 23% areal expansion. The maximum depth of macrophyte colonization, as measured by diver and hydroacoustic surveys, increased from 3.0 m before the invasion of zebra mussels to 5.1 m after their establishment. In addition, macrophyte species richness increased, the frequency of occurrence increased for most species, and the composition of the macrophyte community changed from low-light–tolerant species to those tolerating a wide range of light conditions. Comparisons with observations reported in the literature indicate that increased light penetration alone could explain these changes in macrophyte distribution and diversity. Such changes will increase the importance of benthic primary production over pelagic production in the food web, thereby representing an overall alteration of ecosystem function, a process we refer to as “benthification”.     

Stoichiometry of regenerated nutrients differs between native and invasive freshwater mussels with implications for algal growth

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Long‐term mark‐and‐recapture study of a freshwater mussel reveals patterns of habitat use and an association between survival and river discharge

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The effect of Hurricane Katrina on the mussel assemblage of the Pearl River, Louisiana

We sampled the mussels of the Pearl River along the eastern border of Louisiana, USA, in 1997 and 2007. Hurricane Katrina followed the pathway of the river in August 2005, and we were interested in detecting any resulting decreases in mussel abundance or diversity. The mussel assemblage was relatively stable, despite the hurricane and a drought in the Pearl River Basin in 2006 and 2007. We detected no change in average species richness per site, and the total number of mussels collected per site (e.g., collection per unit effort) actually increased in 2007. Species importance curves comparing the two studies did, however, suggest a reduction from 29 to 23 species. Monte Carlo simulations and comparisons of assemblage similarity also suggested that Glebula rotundata increased at the expense of Potamilus purpuratus and Villosa lienosa, possibly because it was more tolerant of salt water intrusion. Eight of 13 mussels with adequate sample sizes had different adult size structures in the two surveys, either increasing or decreasing in mean shell length during the 10 years. Large individuals were rare for the two species that decreased in relative abundance in 2007. Semi-quantitative sampling appeared adequate here to compare relatively long-term changes in mussel diversity and adult size structure. However, more research on natural variation in assemblage structure and mussel size distributions is needed to understand the importance of disturbances like hurricanes or droughts.     

Phosphorus recycling by zebra mussels in relation to density and food resource availability

Using flow-through microcosms, we examined phosphorus (P) recycling by zebra mussels under conditions of nearly constant food resource supply and varying zebra mussel population densities (600–5200 ind./m²). At all density levels, zebra mussels filtered substantial algae, measured as chlorophyll biomass. Because chlorophyll biomass inputs were low throughout the study, zebra mussel biomass-specific rates of chlorophyll filtration declined with increasing density, suggesting food resource limitation at the higher densities. We observed net total P export and high zebra mussel biomass-specific rates of P recycling over time in microcosms at high zebra mussel densities. In systems with a low zebra mussel density, net total P export did not occur over time. Our results suggest the occurrence of P remineralization by zebra mussels and net loss associated with emaciation during periods of temporary starvation. These findings have implications for P dynamics since zebra mussels can be subjected to periods of starvation over seasonal and annual time scales.     

Water abstraction impacts stream ecosystem functioning via wetted‐channel contraction

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Annual winter water level drawdowns limit shallow‐water mussel densities in small lakes

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Increases in lake phytoplankton biomass caused by future climate‐driven changes to seasonal river flow

For the many lakes world‐wide with short residence times, changes to the rate of water throughput may have important effects on lake ecology. We studied relationships between current and predicted residence times and phytoplankton biomass using a eutrophic lake in the north‐west of England with an annual residence time averaging about 20 days, as a test case. Using 32 years of recent hydrological flow data for Bassenthwaite Lake, multiple sets of scaled flow for each year, and the process‐based phytoplankton response model, PROTECH, we modelled the effects of changing river flow on phytoplankton biomass in the lake. The impact on biomass was shown to depend on seasonal changes in flow rather than annual changes. Furthermore, there was a qualitative difference in impact depending on whether the nutrient loading to the lake came principally from flow‐independent sources, or from flow‐dependent ones. Predictions for changes in river flow under future climate scenarios in the north‐west of England have suggested that, despite little change in the annual flow magnitude, there will be a shift to greater flow in the winter and lesser flow in the summer. Applying these flow predictions to our modelling of Bassenthwaite Lake revealed that, with flow‐independent nutrient loading, and no overall increase in nutrient load, phytoplankton abundance in the summer could increase by up to 70%, including an increased proportion of Cyanobacteria. Conversely, were the loading completely dependent on the flow, the biomass would fall. In many parts of the world, river flow is expected to decrease in the summer even more than in England, suggesting these areas may expect substantial changes to seasonal phytoplankton biomass as a result of climate‐driven changes to seasonal river flow. Such changes would be in addition to any other changes owing to warming effects or eutrophication.     

Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Genome evolution and host‐microbiome shifts correspond with intraspecific niche divergence within harmful algal bloom‐forming Microcystis aeruginosa

Intraspecific niche divergence is an important driver of species range, population abundance and impacts on ecosystem functions. Genetic changes are the primary focus when studying intraspecific divergence; however, the role of ecological interactions, particularly host‐microbiome symbioses, is receiving increased attention. The relative importance of these evolutionary and ecological mechanisms has seen only limited evaluation. To address this question, we used Microcystis aeruginosa, the globally distributed cyanobacterium that dominates freshwater harmful algal blooms. These blooms have been increasing in occurrence and intensity worldwide, causing major economic and ecological damages. We evaluated 46 isolates of M. aeruginosa and their microbiomes, collected from 14 lakes in Michigan, USA, that vary over 20‐fold in phosphorus levels, the primary limiting nutrient in freshwater systems. Genomes of M. aeruginosa diverged along this phosphorus gradient in genomic architecture and protein functions. Fitness in low‐phosphorus lakes corresponded with additional shifts within M. aeruginosa including genome‐wide reductions in nitrogen use, an expansion of phosphorus assimilation genes and an alternative life history strategy of nonclonal colony formation. In addition to host shifts, despite culturing in common‐garden conditions, host‐microbiomes diverged along the gradient in taxonomy, but converged in function with evidence of metabolic interdependence between the host and its microbiome. Divergence corresponded with a physiological trade‐off between fitness in low‐phosphorus environments and growth rate in phosphorus‐rich conditions. Co‐occurrence of genotypes adapted to different nutrient environments in phosphorus‐rich lakes may have critical implications for understanding how M. aeruginosa blooms persist after initial nutrient depletion. Ultimately, we demonstrate that the intertwined effects of genome evolution, host life history strategy and ecological interactions between a host and its microbiome correspond with an intraspecific niche shift with important implications for whole ecosystem function.     

Spatio-temporal patterns of the decline of freshwater mussels in the Little South Fork Cumberland River,USA

The Little South Fork Cumberland River, Kentucky and Tennessee, USA, was a globally important conservation refugium for freshwater mussels (Mollusca: Unionidae) because it supported an intact example (26 species) of the unique Cumberland River mussel fauna including imperiled species. We used previous surveys and our 1997–1998 survey to reconstruct the historical fauna, to describe spatio-temporal patterns of density and number of species, and to evaluate the probable sequence and cause of observed mussel declines. We were specifically interested in better understanding how mussel assemblages respond to chronic disturbances, and how these changes manifest in persistence patterns. Density and numbers of species declined steadily from 1981 to 1998, but declines occurred first in the lower river (early 1980s), followed by declines in the upper river (late 1980s to early 1990s). Of the total species recorded from the Little South Fork, 17 (65%) are seemingly extirpated and five others appear near extirpation. Declines are associated with at least two, temporally distinct major insults. Lower river declines are associated with surface mining, whereas, oil extraction activities are implicated in upper river declines. Regardless of causal factors, species persistence was primarily a function of predecline population size with only the most numerous and widespread species surviving. At this time, the river appears lost as a conservation refugium for mussels despite its remoteness, predominantly forested watershed, and several layers of existing statutory and regulatory environmental safeguards. We suggest that the river could be restored and mussels reintroduced if an interagency task force is formed to identify and mitigate specific stressors now affecting most mussel species in the river.Nomenclature: Turgeon et al. (1998).     

Physical forces constrain the depth distribution of the abundant native mussel Elliptio complanata in lakes

1. Unionid mussels often account for a large portion of benthic biomass and contribute to nutrient cycling and sediment processes, but are thought to be limited to shallow areas (<2–3 m). 2. The depth distribution and body size of Elliptio complanata were compared in seven Canadian Shield lake basins of different sizes to test what factors determine the upper and lower limit of their depth range. Specifically, I tested whether (i) the upper range of their distribution is limited by exposure to winds and wave action and (ii) the lower range of their distribution is limited by the depth of the thermocline or by the boundary of mud deposition. 3. The average depth distribution of E. complanata shifted to greater depths in larger lake basins. When comparing individual transects, maximum mussel density was found deeper at more exposed sites. Mussel size decreased with increasing depth and was larger, on average, in larger lake basins. These results suggest that physical forces limit the upper range of mussel distribution in lakes. 4. The maximum depth at which mussels were found in different lakes was closely related to thermocline depth. However, mussels were commonly observed below the predicted depth of the mud deposition boundary. The thermocline limits the lower range of mussel distribution in lakes, probably by limiting food availability and by determining water temperature. Substratum type does not limit the lower distribution of mussels. 5. These results suggest that unionid mussels are present in the deeper parts of the littoral zone, especially in large lakes. Therefore, comparisons of mussel populations between sites and between lakes would be biased unless the full depth distribution of these mussels is considered. These results also suggest that long‐term changes in the thermal structure of lakes could affect the range of unionid mussel populations and their functional role in littoral ecosystems.     

Biomass loss and change in species dominance shift stream community excretion stoichiometry during severe drought

1. Animals contribute significantly to nutrient cycling through excretion, but most studies consider their effects under relatively benign abiotic conditions. Disturbances such as drought may alter animals’ nutrient contributions through shifts in species composition and biomass. Headwater streams are particularly vulnerable to extreme climate events and thus might show rapid changes in stream biota and their ecosystem effects.
2. We tested how biomass and subsequent ecosystem effects (nutrient cycling) of an intermittent prairie stream community changed during a drought. We quantified the biomass and contributions to nutrient cycling for assemblages comprising fishes, crayfish, and tadpoles in 12 isolated pools over 3 months encompassing the harshest drought on record for Kings Creek, KS, U.S.A. We predicted that macroconsumer biomass would decline with pool surface area and that differences in macroconsumer biomass and taxonomic composition would lead to different contributions of pool assemblages to nutrient cycling.
3. The biomass of pool assemblages declined with decreasing pool size, a pattern apparently driven by mortality, emigration, or metamorphosis. We also observed a change in assemblage structure of drying pools during drought relative to pool size, shifting dominance toward species with more drought-resistant traits. Accordingly, assemblage nitrogen (N) excretion rates declined as pool biomass was reduced, leading to a 58% reduction in N available to epilithic biofilms. Phosphorus (P) excretion rates declined from June to July, but increased in August, as species with high P excretion rates maintained similar proportional biomass and biomass of a non-native fish increased. Molar N:P of pool assemblage excretion declined significantly throughout the drought and coincided with loss of southern redbelly dace (Chrosomus erythrogaster: Cyprinidae).
4. Animal-mediated nutrient cycling was altered by the loss of biomass and stoichiometric traits of taxa that differed in their occurrences and ability to tolerate abiotic conditions during drought. Elevated availability of dissolved N in isolated pools may increase N uptake rates by biofilms during drought conditions, indicating the importance of N excreted by aggregated macroconsumers, especially those with unique stoichiometric traits. While the significance of shifts in the composition of freshwater communities to ecosystems is not entirely known, additional losses in ecosystem function and changes in community structure may follow episodes of severe drought.