Spatial redistribution of nutrients by large herbivores and dung beetles in a savanna ecosystem

Abstract

Abstract1. Territorial or resting behaviour of large herbivores can cause strong local deposits of dung, in different places than where they graze. Additionally, dung beetles and other macrodetritivores can subsequently affect local nutrient budgets through post-depositional re-dispersion of dung and accompanying nutrients. Such horizontal displacement of nutrients by animals has strong implications for savanna ecosystem functioning, but remains poorly studied as it is notoriously difficult to accurately map these flows and incredibly time-consuming.2. In an African savanna, with alternating patches of lawn, bunch grasses and trees/ shrubs, we undertook such effort and studied nutrient aggregation and redistribution by different large herbivore functional groups and dung beetles for a full growing season. We used movable cages to quantify herbivore consumption rates and measured nutrient return through biweekly dung counts. Furthermore, we estimated the offtake of nitrogen (N) and phosphorus (P) by the dominant megagrazer (white rhinoceros) to middens (dung deposition hotspots). Last, we experimentally measured the removal amount and movement paths of telocoprid dung beetles to quantify their nutrient redistribution effects.3. Our estimates suggest white rhinoceros to cause a large export of nutrients from grazing areas towards middens resulting in negative nutrient budgets for both lawn and bunch grassland types. Mesograzers (50-600 kg) realized a net nitrogen input towards high forage quality lawn vegetation at the expense of lower quality bunch grasslands. Browsers caused a net flow from trees/shrubs towards grassland patches.4. Interestingly, while the magnitude of our estimated flows of N consumption and return by large herbivores were rather similar, the P returns were about half of what has been consumed. This is in agreement with ecological stoichiometry theory that predicts that large herbivores should recycle more N than P, due to their relatively high P demand. Furthermore, dung-rolling beetles had a small, but significant, directed movement from lawn to bunch grassland vegetation.5. Synthesis. We conclude that within-ecosystem nutrient redistributions by animals are important and approximately of the same order of magnitude as regional atmospheric nutrient in and outputs (e.g. fire emissions, atmospheric N deposition, biological N fixation), and hence are important for understanding savanna ecosystem functioning.

Synthesizing the effects of large, wild herbivore exclusion on ecosystem function

“…Leroux & Schmitz, 2015;Stark et al, 2015). Moose and hippopotamus, for example, move substantial quantities of nutrients between terrestrial and aquatic ecosystems, increasing nutrient availability and subsidizing consumers in recipient systems (Stears et al, 2018); similarly, rhinoceroses maintain nutrient (and secondarily, structural) heterogeneity via the lateral transfer of nutrients across a single savanna system (Veldhuis, Gommers, Olff, & Berg, 2017). Though exclosure experiments are generally inappropriate to study these landscape-scale effects of herbivores on nutrient cycling, recent synthesis nonetheless suggests that effects of such transfer likely vary across characteristics of both nutrient donor and recipient ecosystems and the herbivore species involved (Subalusky & Post, 2019).…”

Primary productivity and habitat protection predict elevational species richness and community biomass of large mammals on Mt. Kilimanjaro

“…Large mammals are of high ecological importance and are often used as flagship species for conservation (Williams, Burgess, & Rahbek, ). They play a crucial role in controlling ecosystem processes such as nutrient cycling and energy flow by turning over high amounts of biomass (Veldhuis, Gommers, Olff, & Berg, ). Meta‐analyses suggest that large mammals are particularly threatened by the loss of natural habitats and hunting (Hegerl et al, ).…”

Rolling pits of Hartmann’s mountain zebra (Zebra equus hartmannae) increase vegetation diversity and landscape heterogeneity in the Pre‐Namib

“…Deposits of feces and higher urine concentrations in the soil are a typical feature known from other resting places of larger animals such as Gemsbok (Dean & Milton, 1991a , 1991b ) or Kangaroo (Eldridge & Rath, 2002 ). In addition, such depressions frequently accumulate litter (Mallen‐Cooper et al., 2019 ), which is then together with the dung quickly converted into nutrients by the stronger mineralization due to a higher soil moisture (Eldridge & Rath, 2002 ; Veldhuis et al., 2018 ). As a result, these resting places and so the rolling pits of the Hartmann's mountain zebra form relatively moist and nutrient‐rich patches in the otherwise nutrient‐poor savanna (Augustine & McNaughton, 2006 ; McNaughton et al., 1988 ).…”