The inappropriate use of time-to-independence biases estimates of activity patterns of free-ranging mammals derived from camera traps.

Measuring and comparing activity patterns provide key insights into the behavioral trade-offs that result in animal activity and their extrinsic and intrinsic drivers. Camera traps are a recently emerged source of data for sampling animal activity used to estimate activity patterns. However, nearly 70% of studies using such data to estimate activity patterns apply a time-to-independence data filter to discard appreciable periods of sampling effort. This treatment of activity as a discrete event emerged from the use of camera trap data to estimate animal abundances, but does not reflect the continuous nature of behavior, and may bias resulting estimates of activity patterns. We used a large, freely available camera trap dataset to test the effects of time to independence on the estimated activity of eight medium- to large-sized African mammals. We show that discarding data through the use of time-to-independence filters causes substantial losses in sample sizes and differences in the estimated activity of species. Activity patterns estimated for herbivore species were more affected by the application of time-to-independence data filters than carnivores, this extending to estimates of potential interactions (activity overlap) between herbivore species. We hypothesize that this pattern could reflect the typically more abundant, social, and patch-specific foraging patterns of herbivores and suggest that this effect may bias estimates of predator-prey interactions. Activity estimates of rare species, with less data available, may be particularly vulnerable to loss of data through the application of time-to-independence data filters. We conclude that the application of time-to-independence data filters in camera trap-based estimates of activity patterns is not valid and should not be used.

The generality of cryptic dietary niche differences in diverse large-herbivore assemblages.

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.

Trophic rewilding as a climate change mitigation strategy?

The loss of megafauna at the terminal Pleistocene has been linked to a wide range of Earth-system-level changes, such as altered greenhouse gas budgets, fire regimes and biome-level vegetation changes. Given these influences and feedbacks, might part of the solution for mitigating anthropogenic climate change lie in the restoration of extant megafauna to ecosystems? Here, we explore the potential role of trophic rewilding on Earth's climate system. We first provide a novel synthesis of the various ways that megafauna interact with the major drivers of anthropogenic climate change, including greenhouse gas storage and emission, aerosols and albedo. We then explore the role of rewilding as a mitigation tool at two scales: (i) current and near-future opportunities for national or regional climate change mitigation portfolios, and (ii) more radical opportunities at the global scale. Finally, we identify major knowledge gaps that complicate the complete characterization of rewilding as a climate change mitigation strategy. Our perspective is urgent since we are losing the Earth's last remaining megafauna, and with it a potential option to address climate change.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Diet shifts by adult flightless dung beetles Circellium bacchus, revealed using DNA metabarcoding, reflect complex life histories.

Life history changes may change resource use. Such shifts are not well understood in the dung beetles, despite recognised differences in larval and adult feeding ability. We use the flightless dung beetle Circellium bacchus to explore such shifts, identifying dung sources of adults using DNA metabarcoding, and comparing these with published accounts of larval dung sources. C. bacchus is traditionally considered to specialise on the dung of large herbivores for both larval and adult feeding. We successfully extracted mammal DNA from 151 adult C. bacchus fecal samples, representing 16 mammal species (ranging from elephants to small rodents), many of which are hitherto undescribed in the diet. Adult C. bacchus showed clear dung source preferences, especially for large herbivores inhabiting dense-cover vegetation. Our approach also confirmed the presence of cryptic taxa in the study area, and we propose that this may be used for biodiversity survey and monitoring purposes. Murid rodent feces were the most commonly fed-upon dung source (77.5%) for adult C. bacchus, differing markedly from the large and megaherbivore dung sources used for larval rearing. These findings support the hypothesis of life history-specific shifts in resource use in dung beetles, and reveal a hitherto unsuspected, but ecologically important, role of these dung beetles in consuming rodent feces. The differences in feeding abilities of the larval and adult life history stages have profound consequences for their resource use and foraging strategies, and hence the ecological role of dung beetles. This principle and its ecological consequences should be explored in other scarabaeids.

Expert-derived monitoring thresholds for impacts of megaherbivores on vegetation cover in a protected area.

Monitoring is meant to inform conservation authorities, yet managers often don't know when to respond to monitoring results. One of the reasons is that management often lacks consensus on monitoring thresholds for intervention. This results in aimless monitoring without a clear directive on when monitoring indicates a trajectory towards an unacceptable state or impending change, which possibly necessitates intervention. Although experts rarely provide simple, measureable and quantifiable monitoring thresholds as required by management, they are often more comfortable expressing opinions on whether a specific area is desirable or not. This allows thresholds to be reverse engineered: by getting experts to identify sites as desirable and undesirable, field variables can subsequently be measured to derive the boundary between subjectively identified desirable and undesirable states. Such a boundary provides a defendable point for management to assess and consider intervention. Here we describe the identification of monitoring thresholds by defining the limits of desirable canopy cover, derived from expert stakeholder preferences, in the Sundays Spekboom Thicket vegetation of the Addo Elephant National Park, South Africa. The park has experienced variable utilization intensity by large herbivores, especially elephant. For years managers have grappled with the question of what percentage shrub canopy cover is desirable as a management target, but science has failed to provide this. Using experts to assess pre-selected sites as desirable or undesirable across a range of canopy covers, we showed that a canopy cover of ∼65% (±15%) would be desirable for expert stakeholders. We then used satellite imagery to map canopy cover, providing managers for the first time with a large-scale map of canopy cover, indicating desirability status. This approach was useful for facilitating joint-decision making between conservation agencies and stakeholders on tangible indicators of achieving goals, and may be useful in fostering relationships, trust, mutual understanding and transparency, characteristics critical for managing complex socio-ecological systems.

Long­term monitoring reveals differing impacts of elephants on elements of a canopy shrub community.

The conservation management of southern Africa's elephants focuses on identifying and mitigating the extent and intensity of impacts on biological diversity. However, variation in the intensity of elephant effects between elements of biodiversity is seldom explored, which limits our ability to interpret the scale of the impacts. Our study quantifies >50 years of impacts in the succulent thickets of the Addo Elephant National Park, South Africa, contrasting hypotheses for the resilience of the canopy shrubs (a key functional guild) to elephants with those that argue the opposite. We also assess the impacts between elements of the community, ranging from community composition and structure to the structure of individual canopy species. We show the vulnerability of the canopy shrubs to transformation as the accumulated influences of elephants alter community composition and structure. The pattern of transformation is similar to that caused by domestic herbivores, which leads us to predict that elephants will eventually bring about landscape-level degradation and a significant loss of biodiversity. While we expected the canopy species to show similar declining trends in structure, providing insight into the response of the community as a whole, we demonstrate an uneven distribution of impacts between constituent elements; most of the canopy dominants exhibited little change, resisting removal. This implies that these canopy dominants might not be useful indicators of community change in thickets, a pattern that is likely repeated among the canopy trees of savanna systems. Our findings suggest that predicting elephant impacts, and finding solutions to the so-called "elephant problem," require a broader and more integrated understanding of the mechanisms driving the changes between elements of biodiversity at various spatial and temporal scales.

Shift in black rhinoceros diet in the presence of elephant: evidence for competition?

In African large herbivore assemblages, megaherbivores dominate the biomass and utilise the greatest share of available resources. Consequently, they are considered a separate trophic guild that structures the food niches of coexisting large herbivores. However, there exists little empirical evidence on how food resources are shared within this guild, and none for direct competition for food between megaherbivores. Using the histological analysis of faeces, we explore this phenomenon for African elephant Loxodonta africana and black rhinoceros Diceros bicornis in the Addo Elephant National Park, South Africa, where the accumulated impacts of elephant have reduced browse availability. Despite being unable to generalise beyond our study sites, our observations support the predictions of competition theory (as opposed to optimality theory) by showing (1) a clear seasonal separation in resource use between these megaherbivores that increased as resource availability declined, and (2) rhinoceros changed their selectivity in the absence of elephant (using an adjacent site) by expanding and shifting their diet along the grass-browse continuum, and in relation to availability. Although black rhinoceros are generally considered strict browsers, the most significant shift in diet occurred as rhinoceros increased their preferences for grasses in the presence of elephant. We speculate that the lack of specialised grazing adaptations may increase foraging costs in rhinoceros, through reduced harvest- and handling-efficiencies of grasses. In the short-term, this may be off-set by an enhanced tolerance for low quality food and by seasonally mobilising fat reserves; however, the long-term fitness consequences require further study. Our data suggest that managing elephant at high densities may compromise the foraging opportunities of coexisting browsers. This may be particularly important in small, fenced areas and overlapping preferred habitats where impacts intensify.

Understanding long-term variations in an elephant piosphere effect to manage impacts.

Surface water availability is a key driver of elephant impacts on biological diversity. Thus, understanding the spatio-temporal variations of these impacts in relation to water is critical to their management. However, elephant piosphere effects (i.e. the radial pattern of attenuating impact) are poorly described, with few long-term quantitative studies. Our understanding is further confounded by the complexity of systems with elephant (i.e. fenced, multiple water points, seasonal water availability, varying population densities) that likely limit the use of conceptual models to predict these impacts. Using 31 years of data on shrub structure in the succulent thickets of the Addo Elephant National Park, South Africa, we tested elephant effects at a single water point. Shrub structure showed a clear sigmoid response with distance from water, declining at both the upper and lower limits of sampling. Adjacent to water, this decline caused a roughly 300-m radial expansion of the grass-dominated habitats that replace shrub communities. Despite the clear relationship between shrub structure and ecological functioning in thicket, the extent of elephant effects varied between these features with distance from water. Moreover, these patterns co-varied with other confounding variables (e.g. the location of neighboring water points), which limits our ability to predict such effects in the absence of long-term data. We predict that elephant have the ability to cause severe transformation in succulent thicket habitats with abundant water supply and elevated elephant numbers. However, these piosphere effects are complex, suggesting that a more integrated understanding of elephant impacts on ecological heterogeneity may be required before water availability is used as a tool to manage impacts. We caution against the establishment of water points in novel succulent thicket habitats, and advocate a significant reduction in water provisioning at our study site, albeit with greater impacts at each water point.