Daniel C. Allen

Complex hydraulic and substrate variables limit freshwater mussel species richness and abundance

Abstract We examined how substrate and complex hydraulic variables limit the distribution of freshwater mussels. We sampled mussels and measured substrate and hydraulic variables (at low and high flows) at 6 sites in the Little River, Oklahoma. To test which variables were most limiting to mussel species richness and abundance, we evaluated univariate and multiple 95th-, 90th-, and 85th-quantile regression models using a model selection approach. Across all 3 quantiles analyzed, hydraulic variables related to substrate stability (relative shear stress ratio [RSS] and shear stress) at high flows most limited mussel species richness and abundance. High-flow substrate stability models performed the best, but models that used substrate variables (substrate size and heterogeneity) also performed relatively well. Models that used complex hydraulic variables estimated at low flows performed poorly compared to those using the same variables estimated at high flows, a result suggesting that hydraulic conditions at low flows do not limit mussel habitat in our system. Our results demonstrate that substrate stability at high flows is an important factor governing mussel distributions. Last, our quantile regression approach successfully quantified the limiting-factor relationships of substrate and hydraulic characteristics on mussel habitat, and this approach could be used in other studies investigating habitat requirements of aquatic organisms.

Burrowing behavior of freshwater mussels in experimentally manipulated communities

Abstract We experimentally manipulated mussel community structure and observed mussel burrowing behavior in mesocosms held in a greenhouse. Vertical positions, vertical movements, and horizontal movements of Actinonaias ligamentina, Amblema plicata, Fusconaia flava, and Obliquaria reflexa were recorded during five 11-d trials. Community structure was manipulated by constructing communities with 11 different diversity treatments crossed with 3 different density treatments. Vertical positions, vertical movements, and horizontal movements of mussels differed significantly among diversity treatments, and vertical movements differed among density treatments. Differences among diversity treatments were caused by differences in species composition because the burrowing activity of mussels in multispecies communities could be predicted additively from single-species communities. The species used in our study vary in body size, but differences among species were still significant after accounting for body length. We think that differences in species burrowing behavior might be a result of niche partitioning of vertical space, might be a result of differing effects of temperature between species, or might be related to mechanisms to avoid dislodgement during high flows. The burrowing behavior of freshwater mussels has implications for mussel sampling protocols, the sensitivity of mussels to zebra mussel attachment, and how mussels influence benthic ecosystems.

Density-dependent biodiversity effects on physical habitat modification by freshwater bivalves

Several decades of research have shown that biodiversity affects ecosystem processes associated with resource capture and the production of biomass within trophic levels. Although there are good reasons to expect that biodiversity influences non-trophic ecosystem processes, such as the physical creation or modification of habitat, studies investigating the role of biodiversity on physical processes are scarce. Here we report the results of a study using artificial streams to test the influence of freshwater mussel biodiversity on gravel erosion during high flows while manipulating mussel abundance. Mussel species vary in traits that should influence their effects on erosion, such as size, shell morphology, and burrowing behavior. We found that mussel species richness was associated with an increase in erosion at both low and high densities. Planned contrasts showed that the erosion observed in species mixtures was purely additive at low density, indicating that erosion in a species polyculture could routinely be predicted by the performance of monocultures. However, at high density certain combinations of species showed nonadditive effects on erosion, suggesting that organism abundance can fundamentally alter biodiversity effects. Although this may have been a result of altered species interactions at high density, our study design cannot confirm this.

Bottom‐up biodiversity effects increase resource subsidy flux between ecosystems

Although biodiversity can increase ecosystem productivity and adjacent ecosystems are often linked by resource flows between them, the relationship between biodiversity and resource subsidies is not well understood. Here we test the influence of biodiversity on resource subsidy flux by manipulating freshwater mussel species richness and documenting the effects on a trophic cascade from aquatic to terrestrial ecosystems. In a mesocosm experiment, mussel effects on algae were linked through stable isotope analyses to mussel-derived nitrogen subsidies, but mussel biodiversity effects on algal accumulation were not significant. In contrast, mussel biodiversity significantly increased aquatic insect emergence rates, because aquatic insects were responding to mussel-induced changes in algal community structure instead of algal accumulation. In turn, mussel biodiversity also significantly increased terrestrial spider abundance as spiders tracked increases in aquatic insect prey after a reproduction event. In a comparative field study, we found that sites with greater mussel species richness had higher aquatic insect emergence rates. These results show that, because food webs in adjacent ecosystems are often linked, biodiversity effects in one ecosystem can influence adjacent ecosystems as well.

Meta-analysis: abundance, behavior, and hydraulic energy shape biotic effects on sediment transport in streams

An increasing number of studies have emphasized the need to bridge the disciplines of ecology and geomorphology. A large number of case studies show that organisms can affect erosion, but a comprehensive understanding of biological impacts on sediment transport conditions is still lacking. We use meta-analysis to synthesize published data to quantify the effects of the abundance, body size, and behavior of organisms on erosion in streams. We also explore the influence of current velocity, discharge, and sediment grain size on the strength of biotic effects on erosion. We found that species that both increase erosion (destabilizers) and decrease erosion (stabilizers) can alter incipient sediment motion, sediment suspension, and sediment deposition above control conditions in which the organisms were not present. When abundance was directly manipulated, these biotic effects were consistently stronger in the higher abundance treatment, increasing effect sizes by 66%. Per capita effect size and per capita biomass were also consistently positively correlated. Fish and crustaceans were the most studied organisms, but aquatic insects increased the effect size by 550 x compared to other types of organisms after accounting for biomass. In streams with lower discharge and smaller grain sizes, we consistently found stronger biotic effects. Taken collectively, these findings provide synthetic evidence that biology can affect physical processes in streams, and these effects can be mediated by hydraulic energy. We suggest that future studies focus on understudied organisms, such as biofilms, conducting experiments under realistic field conditions, and developing hypotheses for the effect of biology on erosion and velocity currents in the context of restoration to better understand the forces that mediate physical disturbances in stream ecosystems.

Water as a trophic currency in dryland food webs

Water is essential for life on Earth, yet little is known about how water acts as a trophic currency, a unit of value in determining species interactions in terrestrial food webs. We tested the relative importance of groundwater and surface water in riparian food webs by manipulating their availability in dryland floodplains. Primary consumers (crickets) increased in abundance in response to added surface water and groundwater (contained in moist leaves), and predators (spiders and lizards) increased in abundance in response to added surface water, in spite of the presence of a river, an abundant water source. Moreover, the relative magnitude of organism responses to added water was greatest at the most arid site and lowest at the least arid site, mirroring cricket recruitment, which was greatest at the least arid site and lowest at the most arid site. These results suggest that water may be a key currency in terrestrial dryland food webs, which has important implications for predicting ecosystem response...

Early Life-history and Conservation Status of Venustaconcha Ellipsiformis (Bivalvia, Unionidae) in Minnesota

ABSTRACT The ellipse, Venustaconcha ellipsiformis (Bivalvia: Unionidae), was first recorded in Minnesota from the Straight River in 1987, but we knew little of its distribution in the state, brooding behavior, glochidial host fish relationships or status. To examine these questions, we followed standard procedures to conduct an extensive, qualitative statewide mussel survey, described mantle flapping behavior from field and laboratory observations and identified suitable glochidia hosts in the laboratory and from naturally infested fishes. We found extant populations in five Mississippi River tributaries in southeastern Minnesota: the Cannon, Cedar, Root, Upper Iowa and Zumbro rivers. This appears to be the extent of its historic range in Minnesota as no valid records were found elsewhere. Among these, the largest population occurred in the headwaters of the Root River drainage, which was also the only drainage where we observed recent recruitment. Brooding V. ellipsiformis quickly flap a small mantle extension, often in response to passing shadows or jarring of the substrate, or their mantle extensions may be slowly undulated. We identified 11 fish species as suitable hosts for V. ellipsiformis glochidia in the laboratory: brook stickleback (Culaea inconstans), mottled sculpin (Cottus bairdii), slimy sculpin (C. cognatus), logperch (Percina caprodes), mud darter (Etheostoma asprigene), rainbow darter (E. caeruleum), Iowa darter (E. exile), fantail darter (E. flabbelare), Johnny darter (E. nigrum), banded darter (E. zonale) and blackside darter (P. maculata). Rainbow darter, fantail darter and blackside darter were also found naturally infested with V. ellipsiformis glochidia. Venustaconcha ellipsiformis should remain classified as a “Threatened” species in Minnesota and management should include conserving populations within drainages due to its restricted range and likely low dispersal ability.

A Tale of Two Rivers: Implications of Water Management Practices for Mussel Biodiversity Outcomes During Droughts

Droughts often pose situations where stream water levels are lowest while human demand for water is highest. Here we present results of an observational study documenting changes in freshwater mussel communities in two southern US rivers during a multi-year drought. During a 13-year period water releases into the Kiamichi River from an impoundment were halted during droughts, while minimum releases from an impoundment were maintained in the Little River. The Kiamichi observed nearly twice as many low-flow events known to cause mussel mortality than the Little, and regression tree analyses suggest that this difference was influenced by reduced releases. During this period mussel communities in the Kiamichi declined in species richness and abundance, changes that were not observed in the Little. These results suggest that reduced releases during droughts likely led to mussel declines in one river, while maintaining reservoir releases may have sustained mussel populations in another.

Synthesis: comparing effects of resource and consumer fluxes into recipient food webs using meta-analysis.

Here we synthesize empirical research using meta-analysis to compare how consumer and resource fluxes affect recipient food webs. We tested the following hypotheses: (H1) The direct effects of resource fluxes (bottom-up) should be stronger than the direct effects of consumer fluxes (top-down), because resource fluxes are permanent (do not return to the food web in which they were produced) but consumer fluxes may not be (consumers can leave). (H2) Following H1, the indirect effects should attenuate (weaken) more quickly for consumer fluxes than for resource fluxes due to their direct effects being weaker, (H3) The effects of resource fluxes should be stronger when recipient food webs are in different ecosystems than donor food webs due to differences in elevation that accompany cross-ecosystem food web interfaces, often increasing flux quantity due to gravity, while the effects of consumer fluxes should be stronger when donor and recipient food webs are in the same ecosystem as they should more easily assimilate into the recipient food web. We found no differences in the magnitude of bottom-up and top-down direct effects for resource and consumer fluxes, but top-down direct effects were 122% stronger than top-down indirect effects. Indirect effects of prey and predator fluxes quickly attenuated while indirect effects of non-prey resource and herbivore fluxes did not, as the overall direct effects of prey and predator fluxes were 123% and 163% stronger than their indirect effects, respectively. This result suggests that the magnitude of indirect effects decrease as the trophic level of resource and consumer fluxes increases, and also contrasts with results from studies showing in situ top-down indirect effects are stronger than in situ bottom-up indirect effects. We found that resource and consumer flux effect sizes were similar when they occurred between ecosystems, but when they occurred within ecosystems predator flux effects were 107% stronger than nutrient flux effects. Finally, we found that observational studies had higher effect sizes than manipulative studies. Future research should focus on how resource and consumer fluxes might interact and generate feedbacks in empirical studies of natural food webs, and what ecological factors might affect their relative strength.

Plant biodiversity effects in reducing fluvial erosion are limited to low species richness.

It has been proposed that plant biodiversity may increase the erosion resistance of soils, yet direct evidence for any such relationship is lacking. We conducted a mesocosm experiment with eight species of riparian herbaceous plants, and found evidence that plant biodiversity significantly reduced fluvial erosion rates, with the eight-species polyculture decreasing erosion by 23% relative to monocultures. Species richness effects were largest at low levels of species richness, with little increase between four and eight species. Our results suggest that plant biodiversity reduced erosion rates indirectly through positive effects on root length and number of root tips, and that interactions between legumes and non-legumes were particularly important in producing biodiversity effects. Presumably, legumes increased root production of non-legumes by increasing soil nitrogen availability due to their ability to fix atmospheric nitrogen. Our data suggest that a restoration project using species from different functional groups might provide the best insurance to maintain long-term erosion resistance.