Large herbivorous wildlife and livestock differentially influence the relative importance of different sources of energy for riverine food webs

Abstract

In many regions of the world, large populations of native wildlife have declined or been replaced by livestock grazing areas and farmlands, with consequences for terrestrial-aquatic ecosystem connectivity and trophic resources supporting food webs in aquatic ecosystems. The river continuum concept (RCC) and the riverine productivity model (RPM) predict a shift of energy supplying aquatic food webs along rivers: from terrestrial inputs in low-order streams to autochthonous production in mid-sized rivers. In Afromontane-savanna landscapes, the shifting numbers of large mammalian wildlife present a physical continuum whose ecological implications for rivers is not clearly understood. Here, we studied the influence of replacing large wildlife (mainly hippos) with livestock on the fractional contribution of C3 vegetation, C4 grasses and periphyton on macroinvertebrates in the Mara River, which is an African montane-savanna river known to receive large subsidy fluxes of terrestrial organic matter and nutrients mediated by large mammalian herbivores (LMH), both wildlife and livestock, in its middle and lower reaches. Using stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes, we identified spatial patterns in the fractional contribution of allochthonous organic matter from C3 and C4 plants (woody vegetation and grasses, respectively) and autochthonous energy from periphyton for macroinvertebrates at various sites of the Mara River and its tributaries. Potential energy sources and invertebrates were sampled at 80 sites spanning stream orders 1 to 7, various catchment land uses (forest, agriculture and grasslands) and different loading rates of organic matter and nutrients by LMH (livestock and wildlife, i.e., hippopotamus). The fractional contribution of different sources of energy for macroinvertebrates along the river did not follow predictions of the RCC and RPM. First, the fractional contribution of C3 and C4 carbon was not related to river order or location along the fluvial continuum but to the loading of organic matter (dung) by both wildlife and livestock. Notably, C4 carbon was important for macroinvertebrates even in large river sections inhabited by hippos. Second, even in small 1st -3rd order forested streams, periphyton was a major source of energy for macroinvertebrates, and this was fostered by livestock inputs fuelling aquatic primary production throughout the river network. Importantly, our results show that replacing wildlife (hippos) with livestock shifts river systems towards greater reliance on autochthonous sources of energy through an algae-grazer pathway as opposed to reliance on allochthonous inputs of C4 carbon through a detrital pathway.

Keywords: Aquatic-terrestrial connectivity; C3 carbon; C4 carbon; MixSIAR models; Nutrient loading; Organic matter; Resource subsidies.