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  1. Free, publicly-accessible full text available February 1, 2024
  2. Austin, A (Ed.)
    Sympatric large mammalian herbivore species differ in diet composition, both by eating different parts of the same plant and by eating different plant species. Various theories proposed to explain these differences are not mutually exclusive, but are difficult to reconcile and confront with data. Moreover, whereas several of these ideas were originally developed with reference to within-plant partitioning (i.e., consumption of different tissues), they may analogously apply to partitioning of plant species; this possibility has received little attention. Plant functional traits provide a novel window into herbivore diets and a means of testing multiple hypotheses in a unified framework. We used DNA metabarcoding to characterize the diets of 14 sympatric large-herbivore species in an African savanna and analyzed diet composition in light of 27 functional traits that we measured locally for 204 plant species. Plant traits associated with the deep phylogenetic split between grasses and eudicots formed the primary axis of resource partitioning, affirming the generality and importance of the grazer-browser spectrum. A secondary axis comprised plant traits relevant to herbivore body size. Plant taxa in the diets of large-bodied species were lower on average in digestible energy and protein, taller on average (especially among grazers), and tended to be higher in tensile strength, zinc, stem-specific density, and potassium (and lower in sodium, stem dry matter content, and copper). These results are consistent with longstanding hypotheses linking body size with forage quality and height, yet they also suggest the existence of undiscovered links between herbivore body size and a set of rarely considered food-plant traits. We also tested the novel hypothesis that the leaf economic spectrum (LES), a major focus in plant ecology, is an axis of resource partitioning in large-herbivore assemblages; we found that the LES was a minor axis of individual variation within a few species, but had little effect on interspecific dietary differentiation. Synthesis. These results identify key plant traits that underpin the partitioning of food-plant species in large-herbivore communities and suggest that accounting for multiple plant traits (and tradeoffs among them) will enable a deeper understanding of herbivore-plant interaction networks. 
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  3. Abstract

    Many populations of consumers consist of relatively specialized individuals that eat only a subset of the foods consumed by the population at large. Although the ecological significance of individual‐level diet variation is recognized, such variation is difficult to document, and its underlying mechanisms are poorly understood. Optimal foraging theory provides a useful framework for predicting how individuals might select different diets, positing that animals balance the “opportunity cost” of stopping to eat an available food item against the cost of searching for something more nutritious; diet composition should be contingent on the distribution of food, and individual foragers should be more selective when they have greater energy reserves to invest in searching for high‐quality foods. We tested these predicted mechanisms of individual niche differentiation by quantifying environmental (resource heterogeneity) and organismal (nutritional condition) determinants of diet in a widespread browsing antelope (bushbuck,Tragelaphus sylvaticus) in an African floodplain‐savanna ecosystem. We quantified individuals' realized dietary niches (taxonomic richness and composition) using DNA metabarcoding of fecal samples collected repeatedly from 15 GPS‐collared animals (range 6–14 samples per individual, median 12). Bushbuck diets were structured by spatial heterogeneity and constrained by individual condition. We observed significant individual‐level partitioning of food plants by bushbuck both within and between two adjacent habitat types (floodplain and woodland). Individuals with home ranges that were closer together and/or had similar vegetation structure (measured using LiDAR) ate more similar diets, supporting the prediction that heterogeneous resource distribution promotes individual differentiation. Individuals in good nutritional condition had significantly narrower diets (fewer plant taxa), searched their home ranges more intensively (intensity‐of‐use index), and had higher‐quality diets (percent digestible protein) than those in poor condition, supporting the prediction that animals with greater endogenous reserves have narrower realized niches because they can invest more time in searching for nutritious foods. Our results support predictions from optimal foraging theory about the energetic basis of individual‐level dietary variation and provide a potentially generalizable framework for understanding how individuals' realized niche width is governed by animal behavior and physiology in heterogeneous landscapes.

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  4. Ecological niche differences are necessary for stable species coexistence but are often difficult to discern. Models of dietary niche differentiation in large mammalian herbivores invoke the quality, quantity, and spatiotemporal distribution of plant tissues and growth forms but are agnostic toward food plant species identity. Empirical support for these models is variable, suggesting that additional mechanisms of resource partitioning may be important in sustaining large-herbivore diversity in African savannas. We used DNA metabarcoding to conduct a taxonomically explicit analysis of large-herbivore diets across southeastern Africa, analyzing ∼4,000 fecal samples of 30 species from 10 sites in seven countries over 6 y. We detected 893 food plant taxa from 124 families, but just two families—grasses and legumes—accounted for the majority of herbivore diets. Nonetheless, herbivore species almost invariably partitioned food plant taxa; diet composition differed significantly in 97% of pairwise comparisons between sympatric species, and dissimilarity was pronounced even between the strictest grazers (grass eaters), strictest browsers (nongrass eaters), and closest relatives at each site. Niche differentiation was weakest in an ecosystem recovering from catastrophic defaunation, indicating that food plant partitioning is driven by species interactions, and was stronger at low rainfall, as expected if interspecific competition is a predominant driver. Diets differed more between browsers than grazers, which predictably shaped community organization: Grazer-dominated trophic networks had higher nestedness and lower modularity. That dietary differentiation is structured along taxonomic lines complements prior work on how herbivores partition plant parts and patches and suggests that common mechanisms govern herbivore coexistence and community assembly in savannas. 
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  5. null (Ed.)
  6. The world’s largest carnivores are declining and now occupy mere fractions of their historical ranges. Theory predicts that when apex predators disappear, large herbivores should become less fearful, occupy new habitats, and modify those habitats by eating new food plants. Yet experimental support for this prediction has been difficult to obtain in large-mammal systems. Following the extirpation of leopards and African wild dogs from Mozambique’s Gorongosa National Park, forest-dwelling antelopes (bushbuck, Tragelaphus sylvaticus ) expanded into treeless floodplains, where they consumed novel diets and suppressed a common food plant (waterwort, Bergia mossambicensis ). By experimentally simulating predation risk, we demonstrate that this behavior was reversible. Thus, whereas anthropogenic predator extinction disrupted a trophic cascade by enabling rapid differentiation of prey behavior, carnivore restoration may just as rapidly reestablish that cascade. 
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  7. Abstract

    Crop raiding by wildlife poses major threats to both wildlife conservation and human well‐being in agroecosystems worldwide. These threats are particularly acute in many parts of Africa, where crop raiders include globally threatened megafauna such as elephants, and where smallholder agriculture is a primary source of human livelihood. One framework for understanding herbivore feeding behaviour, the forage‐maturation hypothesis, predicts that herbivores should align their movements with intermediate forage biomass (i.e., peak green‐up); this phenomenon is known as “surfing the green wave.” Crop‐raiding elephants, however, often consume not just foliage, but also fruits and tubers (e.g., maize and potatoes), which generally mature after seasonal peaks in photosynthetic activity. Thus, although elephants have been reported to surf the green wave in natural habitats, they may utilize a different strategy in cultivated landscapes by selecting crops that are “browning down.”

    We sought to understand the factors that underpin movement of elephants into agricultural landscapes.

    In Mozambique's Gorongosa National Park, we used movement data from GPS‐collared elephants, together with precipitation records, remotely sensed estimates of landscape greenness (NDVI), DNA‐based diet analysis, measurements of plant nutritional quality and survey‐based metrics of crop availability to understand spatiotemporal variation in elephant crop‐raiding behaviour.

    Elephants tracked peak NDVI while foraging inside the Park. During the dry season, however, when NDVI within the Park declined and availability of mature crops was high, crop raiding increased dramatically, and elephants consistently selected crop plants that were browning down while foraging in cultivated landscapes. Crops contained significantly higher digestible energy than wild food plants, but comparable (and sometimes lower) levels of digestible protein, suggesting that this foraging strategy maximized energy rather than protein intake.

    Our study is the first to combine GPS tracking data with high‐resolution diet analysis and community‐based reporting of crop availability to reveal fine‐scale plasticity in foraging behaviour of elephants at the human–wildlife interface. Our results extend the forage‐maturation hypothesis by showing that elephants surf waves of plant brown‐down in cultivated landscapes. These findings can aid efforts to reduce human–elephant conflict by enabling wildlife managers to prioritize mitigation actions in time and space with limited resources.

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  8. Abstract

    Major disturbances can temporarily remove factors that otherwise constrain population abundance and distribution. During such windows of relaxed top‐down and/or bottom‐up control, ungulate populations can grow rapidly, eventually leading to resource depletion and density‐dependent expansion into less‐preferred habitats. Although many studies have explored the demographic outcomes and ecological impacts of these processes, fewer have examined the individual‐level mechanisms by which they occur. We investigated these mechanisms in Gorongosa National Park, where the Mozambican Civil War devastated large‐mammal populations between 1977 and 1992. Gorongosa’s recovery has been marked by proliferation of waterbuck (Kobus ellipsiprymnus), an historically marginal 200‐kg antelope species, which is now roughly 20‐fold more abundant than before the war. We show that after years of unrestricted population growth, waterbuck have depleted food availability in their historically preferred floodplain habitat and have increasingly expanded into historically avoided savanna habitat. This expansion was demographically skewed: mixed‐sex groups of prime‐age individuals remained more common in the floodplain, while bachelors, loners, and subadults populated the savanna. By coupling DNA metabarcoding and forage analysis, we show that waterbuck in these two habitats ate radically different diets, which were more digestible and protein‐rich in the floodplain than in savanna; thus, although individuals in both habitats achieved positive net energy balance, energetic performance was higher in the floodplain. Analysis of daily activity patterns from high‐resolution GPS‐telemetry, accelerometry, and animal‐borne video revealed that savanna waterbuck spent less time eating, perhaps to accommodate their tougher, lower‐quality diets. Waterbuck in savanna also had more ectoparasites than those in the floodplain. Thus, plasticity in foraging behavior and diet selection enabled savanna waterbuck to tolerate the costs of density‐dependent spillover, at least in the short term; however, the already poorer energetic performance of these individuals implies that savanna occupancy may become prohibitively costly as heterospecific competitors and predators continue to recover in Gorongosa. Our results suggest that behavior can provide a leading indicator of the onset of density‐dependent limitation and the likelihood of subsequent population decline, but that reliable inference hinges on understanding the mechanistic basis of observed behavioral shifts.

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  9. Abstract

    Megafauna assemblages have declined or disappeared throughout much of the world, and many efforts are underway to restore them. Understanding the trophic ecology of such reassembling systems is necessary for predicting recovery dynamics, guiding management, and testing general theory. Yet, there are few studies of recovering large‐mammal communities, and fewer still that have characterized food‐web structure with high taxonomic resolution.

    In Gorongosa National Park, large herbivores have rebounded from near‐extirpation following the Mozambican Civil War (1977–1992). However, contemporary community structure differs radically from the prewar baseline: medium‐sized ungulates now outnumber larger bodied species, and several apex carnivores remain locally extinct.

    We used DNA metabarcoding to quantify diet composition of Gorongosa’s 14 most abundant large‐mammal populations. We tested five hypotheses: (i) the most abundant populations exhibit greatest individual‐level dietary variability; (ii) these populations also have the greatest total niche width (dietary diversity); (iii) interspecific niche overlap is high, with the diets of less‐abundant species nested within those of more‐abundant species; (iv) partitioning of forage species is stronger in more structurally heterogeneous habitats; and (v) selectivity for plant taxa converges within guilds and digestive types, but diverges across them.

    Abundant (and narrow‐mouthed) populations exhibited higher among‐individual dietary variation, but not necessarily the greatest dietary diversity. Interspecific dietary overlap was high, especially among grazers and in structurally homogenous habitat, whereas niche separation was more pronounced among browsers and in heterogeneous habitat. Patterns of selectivity were similar for ruminants—grazers and browsers alike—but differed between ruminants and non‐ruminants.

    Synthesis. The structure of this recovering food web was consistent with several hypotheses predicated on competition, habitat complexity, and herbivore traits, but it differed from patterns observed in more intact assemblages. We propose that intraspecific competition in the fastest‐recovering populations has promoted individual variation and a more nested food web, wherein rare species use subsets of foods eaten by abundant species, and that this scenario is reinforced by weak predation pressure. Future work should test these conjectures and analyse how the taxonomic dietary niche axis studied here interacts with other mechanisms of diet partitioning to affect community reassembly following wildlife declines.

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