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 largemammal 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.
Size‐structured differences in resource use stabilize species coexistence in animal communities, but what behavioral mechanisms underpin these niche differences? Behavior is constrained by morphological and physiological traits that scale allometrically with body size, yet the degree to which behaviors exhibit allometric scaling remains unclear; empirical datasets often encompass broad variation in environmental context and phylogenetic history, which complicates the detection and interpretation of scaling relationships between size and behavior. We studied the movement and foraging behaviors of three sympatric, congeneric spiral‐horned antelope species (Tragelaphus spp.) that differ in body mass—bushbuck (26–40 kg), nyala (57–83 kg), and kudu (80–142 kg)—in an African savanna ecosystem where (i) food was patchily distributed due to ecosystem engineering by fungus‐farming termites and (ii) predation risk was low due to the extirpation of several large carnivores. Because foraging behavior is directly linked to traits that scale allometrically with size (e.g., metabolic rate, locomotion), we hypothesized that habitat use and diet selection would likewise exhibit nonlinear scaling relationships. All three antelope species selected habitat near termitaria, which are hotspots of abundant, high‐quality forage. Experimental removal of forage from termite mounds sharply reduced use of those mounds by bushbuck, confirming that habitat selection was resource driven. Strength of selection for termite mounds scaled negatively and nonlinearly with body mass, as did recursion (frequency with which individuals revisited locations), whereas home‐range area and mean step length scaled positively and nonlinearly with body mass. All species disproportionately ate mound‐associated plant taxa; nonetheless, forage selectivity and dietary composition, richness, and quality all differed among species, reflecting the partitioning of shared food resources. Dietary protein exhibited the theoretically predicted negative allometric relationship with body mass, whereas digestible‐energy content scaled positively. Our results demonstrate cryptic size‐based separation along spatial and dietary niche axes—despite superficial similarities among species—consistent with the idea that body‐size differentiation is driven by selection for divergent resource‐acquisition strategies, which in turn underpin coexistence. Foraging and space‐use behaviors were nonlinearly related to body mass, supporting the hypothesis that behavior scales allometrically with size. However, explaining the variable functional forms of these relationships is a challenge for future research.
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 Ryan A. Long and Robert M. Pringle contributed equally.
Elucidating factors that contribute to citation rates of scientific articles can help scientists write manuscripts that have a stronger influence on their scientific field and are accessible to a broad audience. Using a cohort of 778 articles published in The Journal of Wildlife Management from 2011–2015, we examined how visibility strategies, article structure, and focal system (all factors authors can predominantly control) influenced the accumulation of citations over various time frames within the first 5 years after publication, and the number of days until an article received its first citation. Visibility strategies (e.g., open access, increasing the Altmetric Attention Score, and self‐citations) all influenced the number of citations accrued following publication. Citations were more stochastic 1 year following publication compared to 5 years following publication, with only 20.1% of papers receiving a citation after 1 year compared to 92.5% of papers receiving a citation after 5 years. Our model explained much more of the variation after 5 years compared to after only 1 year (R2 = 0.57 and 0.12, respectively). The number of factors significantly associated with citation rates increased as the timeframe of our analysis increased. After 5 years, factors associated with article structure (e.g., number of references), focal system (e.g., methods papers), and visibility all increased citation counts of papers. Our work suggests citation rates within wildlife ecology are influenced by a number of controllable factors, and that authors pursuing a variety of visibility strategies can increase the influence of an article on science and management.
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