Proactive cursorial and ambush predation risk avoidance in four African herbivore species.

Most herbivores must balance demands to meet nutritional requirements, maintain stable thermoregulation and avoid predation. Species-specific predator and prey characteristics determine the ability of prey to avoid predation and the ability of predators to maximize hunting success. Using GPS collar data from African wild dogs, lions, impala, tsessebes, wildebeest and zebra in the Okavango Delta, Botswana, we studied proactive predation risk avoidance by herbivores. We considered predator activity level in relation to prey movement, predator and prey habitat selection, and preferential use of areas by prey. We compared herbivore behaviour to lion and wild dog activity patterns and determined the effect of seasonal resource availability and prey body mass on anti-predator behaviour. Herbivore movement patterns were more strongly correlated with lion than wild dog activity. Habitat selection by predators was not activity level dependent and, while prey and predators differed to some extent in their habitat selection, there were also overlaps, probably caused by predators seeking habitats with high prey abundance. Areas favoured by lions were used by herbivores more when lions were less active, whereas wild dog activity level was not correlated with prey use. Prey body mass was not a strong predictor of the strength of proactive predation avoidance behaviour. Herbivores showed stronger anti-predator behaviours during the rainy season when resources were abundant. Reducing movement when top predators are most active and avoiding areas with a high likelihood of predator use during the same periods appear to be common strategies to minimize predation risk. Such valuable insights into predator-prey dynamics are only possible when using similar data from multiple sympatric species of predator and prey, an approach that should become more prevalent given the ongoing integration of technological methods into ecological studies.

Dispersal and connectivity in increasingly extreme climatic conditions.

While climate change has been shown to impact several life-history traits of wild-living animal populations, little is known about its effects on dispersal and connectivity. Here, we capitalize on the highly variable flooding regime of the Okavango Delta to investigate the impacts of changing environmental conditions on the dispersal and connectivity of the endangered African wild dog (Lycaon pictus). Based on remote sensed flood extents observed over 20 years, we derive two extreme flood scenarios: a minimum and a maximum flood extent, representative of very dry and very wet environmental periods. These conditions are akin to those anticipated under increased climatic variability, as it is expected under climate change. Using a movement model parameterized with GPS data from dispersing individuals, we simulate 12,000 individual dispersal trajectories across the ecosystem under both scenarios and investigate patterns of connectivity. Across the entire ecosystem, surface water coverage during maximum flood extent reduces dispersal success (i.e., the propensity of individuals to disperse between adjacent subpopulations) by 12% and increases dispersal durations by 17%. Locally, however, dispersal success diminishes by as much as 78%. Depending on the flood extent, alternative dispersal corridors emerge, some of which in the immediate vicinity of human-dominated landscapes. Notably, under maximum flood extent, the number of dispersing trajectories moving into human-dominated landscapes decreases by 41% at the Okavango Delta's inflow, but increases by 126% at the Delta's distal end. This may drive the amplification of human-wildlife conflict. While predicting the impacts of climate change on environmental conditions on the ground remains challenging, our results highlight that environmental change may have significant consequences for dispersal patterns and connectivity, and ultimately, population viability. Acknowledging and anticipating such impacts will be key to effective conservation strategies and to preserve vital dispersal corridors in light of climate change and other human-related landscape alterations.

Higher Mortality Is Not a Universal Cost of Dispersal: A Case Study in African Wild Dogs.

AbstractMortality is considered one of the main costs of dispersal. A reliable evaluation of mortality, however, is often hindered by a lack of information about the fate of individuals that disappear under unexplained circumstances (i.e., missing individuals). Here, we addressed this uncertainty by applying a Bayesian mortality analysis that inferred the fate of missing individuals according to information from individuals with known fate. Specifically, we tested the hypothesis that mortality during dispersal is higher than mortality among nondispersers using 32 years of mark-resighting data from a free-ranging population of the endangered African wild dog (Lycaon pictus) in northern Botswana. Contrary to expectations, we found that mortality during dispersal was lower than mortality among nondispersers, indicating that higher mortality is not a universal cost of dispersal. Our findings suggest that group living can incur costs for certain age classes, such as limited access to resources as group density increases, that exceed the mortality costs associated with dispersal. By challenging the accepted expectation of higher mortality during dispersal, we urge for further investigations of this key life history trait and propose a robust statistical approach to reduce bias in mortality estimates.

Increasing ambient temperatures trigger shifts in activity patterns and temporal partitioning in a large carnivore guild.

Shifts in species' interactions are implicated as an important proximate cause underpinning climate-change-related extinction. However, there is little empirical evidence on the pathways through which climate conditions, such as ambient temperature, impact community dynamics. The timing of activities is a widespread behavioural adaptation to environmental variability, and temporal partitioning is a key mechanism that facilitates coexistence, especially within large carnivore communities. We investigated temperature impacts on community dynamics through its influence on the diel activity of, and temporal partitioning amongst, four sympatric species of African large carnivores: lions (Panthera leo), leopards (Panthera pardus), cheetahs (Acinonyx jubatus) and African wild dogs (Lycaon pictus). Activity of all species was shaped by a combination of light availability and temperature, with most species becoming more nocturnal and decreasing activity levels with increasing temperatures. A nocturnal shift was most pronounced in cheetahs, the most diurnal species during median temperatures. This shift increased temporal overlap between cheetahs and other carnivore species by up to 15.92%, highlighting the importance of considering the responses of interacting sympatric species when inferring climate impacts on ecosystems. Our study provides evidence that temperature can significantly affect temporal partitioning within a carnivore guild by generating asymmetrical behavioural responses amongst functionally similar species.

A three-step approach for assessing landscape connectivity via simulated dispersal: African wild dog case study.

Context: Dispersal of individuals contributes to long-term population persistence, yet requires a sufficient degree of landscape connectivity. To date, connectivity has mainly been investigated using least-cost analysis and circuit theory, two methods that make assumptions that are hardly applicable to dispersal. While these assumptions can be relaxed by explicitly simulating dispersal trajectories across the landscape, a unified approach for such simulations is lacking. Objectives: Here, we propose and apply a simple three-step approach to simulate dispersal and to assess connectivity using empirical GPS movement data and a set of habitat covariates. Methods: In step one of the proposed approach, we use integrated step-selection functions to fit a mechanistic movement model describing habitat and movement preferences of dispersing individuals. In step two, we apply the parameterized model to simulate dispersal across the study area. In step three, we derive three complementary connectivity maps; a heatmap highlighting frequently traversed areas, a betweenness map pinpointing dispersal corridors, and a map of inter-patch connectivity indicating the presence and intensity of functional links between habitat patches. We demonstrate the applicability of the proposed three-step approach in a case study in which we use GPS data collected on dispersing African wild dogs (Lycaon pictus) inhabiting northern Botswana. Results: Using step-selection functions we successfully parametrized a detailed dispersal model that described dispersing individuals' habitat and movement preferences, as well as potential interactions among the two. The model substantially outperformed a model that omitted such interactions and enabled us to simulate 80,000 dispersal trajectories across the study area. Conclusion: By explicitly simulating dispersal trajectories, our approach not only requires fewer unrealistic assumptions about dispersal, but also permits the calculation of multiple connectivity metrics that together provide a comprehensive view of landscape connectivity. In our case study, the three derived connectivity maps revealed several wild dog dispersal hotspots and corridors across the extent of our study area. Each map highlighted a different aspect of landscape connectivity, thus emphasizing their complementary nature. Overall, our case study demonstrates that a simulation-based approach offers a simple yet powerful alternative to traditional connectivity modeling techniques. It is therefore useful for a variety of applications in ecological, evolutionary, and conservation research. Supplementary Information: The online version contains supplementary material available at 10.1007/s10980-023-01602-4.

When to stay and when to leave? Proximate causes of dispersal in an endangered social carnivore.

Reliable estimates of birth, death, emigration and immigration rates are fundamental to understanding and predicting the dynamics of wild populations and, consequently, inform appropriate management actions. However, when individuals disappear from a focal population, inference on their fate is often challenging. Here we used 30 years of individual-based mark-recapture data from a population of free-ranging African wild dogs Lycaon pictus in Botswana and a suite of individual, social and environmental predictors to investigate factors affecting the decision to emigrate from the pack. We subsequently used this information to assign an emigration probability to those individuals that were no longer sighted within their pack (i.e. missing individuals). Natal dispersal (i.e. emigration from the natal pack) showed seasonal patterns with female dispersal peaking prior to the mating season and male dispersal peaking at the beginning of the wet season. For both sexes, natal dispersal rate increased in the absence of unrelated individuals of the opposite sex in the pack. Male natal dispersal decreased with increasing number of pups in the pack and increased in larger packs. Female natal dispersal decreased with increasing number of pups in larger packs, but increased with increasing number of pups in smaller packs. Individuals of both sexes were less likely to exhibit secondary dispersal (i.e. emigration from a pack other than the natal pack) if they were dominant and if many pups were present in the pack. Our models predicted that 18% and 25% of missing females and males, respectively, had likely dispersed from the natal pack, rather than having died. A misclassification of this order of magnitude between dispersal and mortality can have far-reaching consequences in the evaluation and prediction of population dynamics and persistence, and potentially mislead conservation actions. Our study showed that the decision to disperse is context-dependent and that the effect of individual, social and environmental predictors differs between males and females and between natal and secondary dispersal related to different direct and indirect fitness consequences. Furthermore, we demonstrated how a thorough understanding of the proximate causes of dispersal can be used to assign a dispersal probability to missing individuals. Knowledge of causes of dispersal can then be used within an integrated framework to more reliably estimate mortality rates.

Tourist photographs as a scalable framework for wildlife monitoring in protected areas.

Protected areas are critical to conservation efforts in the face of rapid biodiversity declines [1]. Yet the resources for conservation are often limited and shared amongst many competing priorities [2]. As a consequence, even basic monitoring surveys are absent within most protected areas [3]. Although a range of wildlife monitoring methods exist, considerable focused survey effort is often required to yield accurate and precise estimates [4]. This makes monitoring difficult to sustain or replicate, limiting access to the data required for evidence-based conservation decisions. Citizen-scientists have been proposed as an important complement to the finite resources available for basic monitoring within protected areas [5]; however, the full potential of this approach has yet to be realised. Wildlife tourists and guides are especially focussed on encountering and photographing fauna and flora, yet the data collected in these efforts is rarely harnessed for conservation monitoring within protected areas. A detailed understanding of photographic tourism's potential role in wildlife monitoring has been lacking, but is essential for the development of new tools to harness the data being collected through tourism. Here, we demonstrate that tourist-contributed data can aid wildlife monitoring in protected areas by providing population estimates of large carnivores comparable to those from traditional survey methods. Our approach could capitalize upon the immense number of wildlife photographs being taken daily as part of the global > 30-billion USD, wildlife-based tourism industry.

Risk avoidance in sympatric large carnivores: reactive or predictive?

1. Risks of predation or interference competition are major factors shaping the distribution of species. An animal's response to risk can either be reactive, to an immediate risk, or predictive, based on preceding risk or past experiences. The manner in which animals respond to risk is key in understanding avoidance, and hence coexistence, between interacting species. 2. We investigated whether cheetahs (Acinonyx jubatus), known to be affected by predation and competition by lions (Panthera leo) and spotted hyaenas (Crocuta crocuta), respond reactively or predictively to the risks posed by these larger carnivores. 3. We used simultaneous spatial data from Global Positioning System (GPS) radiocollars deployed on all known social groups of cheetahs, lions and spotted hyaenas within a 2700 km(2) study area on the periphery of the Okavango Delta in northern Botswana. The response to risk of encountering lions and spotted hyaenas was explored on three levels: short-term or immediate risk, calculated as the distance to the nearest (contemporaneous) lion or spotted hyaena, long-term risk, calculated as the likelihood of encountering lions and spotted hyaenas based on their cumulative distributions over a 6-month period and habitat-associated risk, quantified by the habitat used by each of the three species. 4. We showed that space and habitat use by cheetahs was similar to that of lions and, to a lesser extent, spotted hyaenas. However, cheetahs avoided immediate risks by positioning themselves further from lions and spotted hyaenas than predicted by a random distribution. 5. Our results suggest that cheetah spatial distribution is a hierarchical process, first driven by resource acquisition and thereafter fine-tuned by predator avoidance; thus suggesting a reactive, rather than a predictive, response to risk.

Fear of the dark or dinner by moonlight? Reduced temporal partitioning among Africa's large carnivores.

Africa is home to the last intact guild of large carnivores and thus provides the only opportunity to investigate mechanisms of coexistence among large predator species. Strong asymmetric dominance hierarchies typically characterize guilds of large carnivores; but despite this asymmetry, subdominant species may persist alongside their stronger counterparts through temporal partitioning of habitat and resources. In the African guild, the subdominant African wild dogs and cheetahs are routinely described as diurnal and crepuscular. These activity patterns have been interpreted to result from the need to avoid encounters with the stronger, nocturnal spotted hyenas and lions. However, the idea that diel activity patterns of carnivore species are strongly shaped by competition and predation has recently been challenged by new observations. In a three-year study in the Okavango Delta, we investigated daily activity patterns and temporal partitioning for wild dogs, cheetahs, spotted hyenas and lions by fitting radio collars that continuously recorded activity bursts, to a total of 25 individuals. Analysis of activity patterns throughout the 24-h cycle revealed an unexpectedly high degree of temporal overlap among the four species. This was mainly due to the extensive and previously undescribed nocturnal activity of wild dogs and cheetahs. Their nocturnal activity fluctuated with the lunar cycle, represented up to 40% of the diel activity budget and was primarily constrained by moonlight availability. In contrast, the nocturnal activity patterns of lions and hyenas were unaffected by moonlight and remained constant over the lunar cycle. Our results suggest that other ecological factors such as optimal hunting conditions have shaped the diel activity patterns of subdominant, large predators. We suggest that they are "starvation driven" and must exploit every opportunity to obtain a meal. The benefits of activity on moonlit nights therefore offset the risks of encountering night-active predators and competitors.