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

In many regions of the world, large populations of native wildlife have declined or been replaced by livestock grazing areas and farmlands, with consequences for terrestrial-aquatic ecosystem connectivity and trophic resources supporting food webs in aquatic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.

Livestock as vectors of organic matter and nutrient loading in aquatic ecosystems in African savannas.

Populations of large wildlife have declined in many landscapes around the world, and have been replaced or displaced by livestock. The consequences of these changes on the transfer of organic matter (OM) and nutrients from terrestrial to aquatic ecosystems are not well understood. We used behavioural data, excretion and egestion rates and C: N: P stoichiometry of dung and urine of zebu cattle, to develop a metabolism-based estimate of loading rates of OM (dung), C, N and P into the Mara River, Kenya. We also directly measured the deposition of OM and urine by cattle into the river during watering. Per head, zebu cattle excrete and/or egest 25.6 g dry matter (DM, 99.6 g wet mass; metabolism) - 27.7 g DM (direct input) of OM, 16.0-21.8 g C, 5.9-9.6 g N, and 0.3-0.5 g P per day into the river. To replace loading rates OM of an individual hippopotamus by cattle, around 100 individuals will be needed, but much less for different elements. In parts of the investigated sub-catchments loading rates by cattle were equivalent to or higher than that of the hippopotamus. The patterns of increased suspended materials and nutrients as a result of livestock activity fit into historical findings on nutrients concentrations, dissolved organic carbon and other variables in agricultural and livestock areas in the Mara River basin. Changing these patterns of carbon and nutrient transport and cycling are having significant effects on the structure and functioning of both terrestrial and aquatic ecosystems.

Hippopotamus are distinct from domestic livestock in their resource subsidies to and effects on aquatic ecosystems.

In many regions of the world, populations of large wildlife have been displaced by livestock, and this may change the functioning of aquatic ecosystems owing to significant differences in the quantity and quality of their dung. We developed a model for estimating loading rates of organic matter (dung) by cattle for comparison with estimated rates for hippopotamus in the Mara River, Kenya. We then conducted a replicated mesocosm experiment to measure ecosystem effects of nutrient and carbon inputs associated with dung from livestock (cattle) versus large wildlife (hippopotamus). Our loading model shows that per capita dung input by cattle is lower than for hippos, but total dung inputs by cattle constitute a significant portion of loading from large herbivores owing to the large numbers of cattle on the landscape. Cattle dung transfers higher amounts of limiting nutrients, major ions and dissolved organic carbon to aquatic ecosystems relative to hippo dung, and gross primary production and microbial biomass were higher in cattle dung treatments than in hippo dung treatments. Our results demonstrate that different forms of animal dung may influence aquatic ecosystems in fundamentally different ways when introduced into aquatic ecosystems as a terrestrially derived resource subsidy.