Are large herbivores vectors of terrestrial subsidies for riverine food webs?

The tropical savannas of Africa have witnessed a dramatic reduction in native large mammalian herbivore populations. The consequences of these changes for terrestrial-aquatic food-web linkages are poorly documented. We used natural abundances of stable carbon and nitrogen isotopes (δ13C, δ15N) to determine spatial and temporal patterns in the importance of herbivore-mediated subsidies for consumers in the Mara River, Kenya. Potential primary producers (terrestrial C3 and C4 producers and periphyton) and consumers (invertebrates and fish) were collected during dry and wet seasons from different sites along the river, representing a gradient from forested highlands to natural savanna grasslands with high herbivore densities across mixed agricultural and livestock-dominated zones. Bayesian mixing models were used to estimate the relative contributions of terrestrial and algal sources of organic carbon supporting consumer trophic groups. Organic carbon sources differed for consumer groups and sites and with season. Overall, periphyton was the major energy source for most consumer groups during the dry season, but with wide 95% confidence intervals. During the wet season, the importance of terrestrial-derived carbon for consumers increased. The importance of C3 producers declined from 40 and 41% at the forested upper reaches to 20 and 8% at river reaches receiving hippo inputs during the dry and wet seasons, respectively. The reciprocal increase in the importance of C4 producers was higher than expected based on areal cover of riparian vegetation that was mainly C3. The importance of C4 producers notably increased from 18 and 10% at the forested upper reaches to 33 and 58% at river reaches receiving hippo inputs during the dry and wet seasons, respectively. This study highlights the importance of large herbivores to the functioning of riverine ecosystems and the potential implications of their loss from savanna landscapes that currently harbor remnant populations. Although the importance of C4 terrestrial carbon in most river systems has been reported to be negligible, this study shows that its importance can be mediated by large herbivores as vectors, which enhance energetic terrestrial-aquatic linkages in rivers in savanna landscapes