Ecosystem bioelement variability is associated with freshwater animal aggregations at the aquatic-terrestrial interface.

The impacts of animals on the biogeochemical cycles of major bioelements like C, N, and P are well-studied across ecosystem types. However, more than 20 elements are necessary for life. The feedbacks between animals and the biogeochemical cycles of the other bioelements are an emerging research priority. We explored how much freshwater mussels (Bivalvia: Unionoida) were related to variability in ecosystem pools of 10 bioelements (Ca, Cu, Fe, K, Mn, Na, Mg, P, S and Zn) in streams containing a natural mussel density gradient in the US Interior Highlands. We studied the concentrations of these bioelements across the aquatic-terrestrial interface-in the porewater of riverine gravel bars, and the emergent macrophyte Justicia americana. Higher mussel density was associated with increased calcium in gravel bars and macrophytes. Mussel density also correlated with variability in iron and other redox-sensitive trace elements in gravel bars and macrophytes, although this relationship was mediated by sediment grain size. We found that two explanations for the patterns we observed are worthy of further research: (1) increased calcium availability in gravel bars near denser mussel aggregations may be a product of the buildup and dissolution of shells in the gravel bar, and (2) mussels may alter redox conditions, and thus elemental availability in gravel bars with fine sediments, either behaviorally or through physical structure provided by shell material. A better understanding of the physical and biogeochemical impacts of animals on a wide range of elemental cycles is thus necessary to conserve the societal value of freshwater ecosystems.

White-tailed deer consumption of emergent macrophytes mediates aquatic-to-terrestrial nutrient flows.

Trophic interactions between mobile animals and their food sources often vector resource flows across ecosystem boundaries. However, the quality and quantity of such ecological subsidies may be altered by indirect interactions between seemingly unconnected taxa. We studied whether emergent macrophytes growing at the aquatic-terrestrial interface facilitate multi-step aquatic-to-terrestrial resource flows between streams and terrestrial herbivores. We also explored whether aquatic animal aggregations indirectly promote such resource flows by creating biogeochemical hotspots of nutrient cycling and availability.We tested whether white-tailed deer (Odocoileus virginianus) in eastern North America vector nutrient fluxes from streams to terrestrial ecosystems by consuming emergent macrophytes (Justicia americana) using isotope and nutrient analyses of fecal samples and motion-sensing cameras. We also tested whether mussel-generated biogeochemical hotspots might promote such fluxes by surveying the density and nutrient stoichiometry of J. americana beds growing in association with variable densities of freshwater mussels (Bivalvia: Unionoida).Fecal samples from riparian deer had 3% lower C:N and 20% lower C:P ratios than those in upland habitats. C and N isotopes suggested riparian deer ate both terrestrial and aquatic (J. americana) vegetation, whereas upland deer ate more terrestrial foods. Motion-sensing cameras showed deer eating J. americana more than twice as frequently at mussel-generated hotspots than non-mussel sites. However, mussels were not associated with variation in J. americana growth or N and P content-although N isotopes in J. americana leaves did suggest assimilation of animal-derived nutrients.Our findings suggest that white-tailed deer may conduct significant transfers of aquatic-derived nutrients into terrestrial habitats when they feed on macrophytes and defecate on land. Whether aquatic animal aggregations promote such resource flows by creating biogeochemical hotspots remains unresolved, but the nearly global distributions of the deer family (Cervidae) and of macrophytes suggest that cervid-driven aquatic-to-terrestrial nutrient flows may be widespread and ecologically important.

Animal aggregations promote emergent aquatic plant production at the aquatic-terrestrial interface.

The roles mobile animals and abiotic processes play as vectors for resource transfers between ecosystems ("subsidies") are well studied, but the idea that resources from animals with limited mobility may be transported across boundaries through intermediate taxa remains unexplored. Aquatic plants ("macrophytes") are globally distributed and may mediate transfers of aquatic-derived nutrients from aggregations of aquatic animals to terrestrial ecosystems when consumed by terrestrial herbivores. We used mesocosms (94 × 44 cm) to test whether aquatic animal-generated biogeochemical hotspots increase growth and nutrient content in macrophytes using the macrophyte Justicia americana and freshwater mussels. Justicia americana biomass production increased and belowground biomass allocation changed with increasing mussel density. At high mussel density, water-column phosphorus increased and carbon:phosphorus ratios in J. americana tissues decreased. We deployed motion-sensing cameras to explore herbivory on J. americana growing along the margins of the Kiamichi River, Oklahoma, and documented feeding by large mammals (Odocoileus virginianus, Sus scrofa, and Bos taurus). Thus, biogeochemical hotspots generated by aquatic animal aggregations can promote macrophyte production that subsequently is transferred to terrestrial animals. More broadly, this suggests that reductions in aquatic animal biomass may have bottom-up impacts that indirectly affect terrestrial ecosystems via plant-animal interactions bridging ecosystem boundaries.