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.

Population density and ranging behaviour of a generalist carnivore varies with human population.

Canid species are highly adaptable, including to urban and peri-urban areas, where they can come into close contact with people. Understanding the mechanisms of wild canid population persistence in these areas is key to managing any negative impacts. The resource dispersion hypothesis predicts that animal density increases and home range size decreases as resource concentration increases, and may help to explain how canids are distributed in environments with an urban-natural gradient. In Australia, dingoes have adapted to human presence, sometimes living in close proximity to towns. Using a targeted camera trap survey and spatial capture-recapture models, we estimated spatial variation in the population density and detection rates of dingoes on Worimi Country in the Great Lakes region of the NSW coast. We tested whether dingo home range and population densities varied across a gradient of human population density, in a mixed-use landscape including, urban, peri-urban, and National Park environs. We found human population density to be a strong driver of dingo density (ranging from 0.025 to 0.433 dingoes/km2 across the natural-urban gradient), and to have a negative effect on dingo home range size. The spatial scale parameter changed depending on survey period, being smaller in the peak tourism period, when human population increases in the area, than in adjacent survey periods, potentially indicating reduced home range size when additional resources are available. Our study highlights the potential value of managing anthropogenic resource availability to manage carnivore densities and potential risk of human-carnivore interactions.

How territoriality and sociality influence the habitat selection and movements of a large carnivore.

While territoriality is one of the key mechanisms influencing carnivore space use, most studies quantify resource selection and movement in the absence of conspecific influence or territorial structure. Our analysis incorporated social information in a resource selection framework to investigate mechanisms of territoriality and intra-specific competition on the habitat selection of a large, social carnivore. We fit integrated step selection functions to 3-h GPS data from 12 collared African wild dog packs in the Okavango Delta and estimated selection coefficients using a conditional Poisson likelihood with random effects. Packs selected for their neighbors' 30-day boundary (defined as their 95% kernel density estimate) and for their own 90-day core (defined as their 50% kernel density estimate). Neighbors' 30-day boundary had a greater influence on resource selection than any habitat feature. Habitat selection differed when they were within versus beyond their neighbors' 30-day boundary. Pack size, pack tenure, pup presence, and seasonality all mediated how packs responded to neighbors' space use, and seasonal dynamics altered the strength of residency. While newly-formed packs and packs with pups avoided their neighbors' boundary, older packs and those without pups selected for it. Packs also selected for the boundary of larger neighboring packs more strongly than that of smaller ones. Social structure within packs has implications for how they interact with conspecifics, and therefore how they are distributed across the landscape. Future research should continue to investigate how territorial processes are mediated by social dynamics and, in turn, how territorial structure mediates resource selection and movement. These results could inform the development of a human-wildlife conflict (HWC) mitigation tool by co-opting the mechanisms of conspecific interactions to manage space use of endangered carnivores.

Size matters: Natural experiments suggest the dear enemy effect is moderated by pack size in African wild dogs.

Remote monitoring of communal marking sites, or latrines, provides a unique opportunity to observe undisturbed scent marking behaviour of African wild dogs (Lycaon pictus). We used remote camera trap observations in a natural experiment to test behavioural scent mark responses to rivals (either familiar neighbours or unfamiliar strangers), to determine whether wild dogs exhibit the "dear enemy" or "nasty neighbour" response. Given that larger groups of wild dogs represent a greater threat to smaller groups, including for established residents, we predicted that the overarching categories "dear enemy" vs. "nasty neighbour" may be confounded by varying social statuses that exists between individual dyads interacting. Using the number of overmarks as a metric, results revealed an interaction between sender and receiver group size irrespective of familiarity consistent with this prediction: in general, individuals from large resident packs overmarked large groups more than they overmarked smaller groups, whereas individuals from smaller packs avoided overmarking larger groups, possibly to avoid detection. Monitoring a natural system highlights variables such as pack size that may be either overlooked or controlled during scent presentation experiments, influencing our ability to gain insights into the factors determining territorial responses to rivals.

Male dingo urinary scents code for age class and wild dingoes respond to this information.

Chemical information in canid urine has been implicated in territoriality and influences the spacing of individuals. We identified the key volatile organic compound (VOC) components in dingo (Canis lupus dingo) urine and investigated the potential role of scents in territorial spacing. VOC analysis, using headspace gas chromatography-mass spectrometry (GC-MS), demonstrated that the information in fresh urine from adult male dingoes was sufficient to allow statistical classification into age categories. Discriminant function analyses demonstrated that the relative amounts or combinations of key VOCs from pre-prime (3-4 years), prime (5-9 years), and post-prime (≥10 years) males varied between these age categories, and that scents exposed to the environment for 4 (but not 33) days could still be classified to age categories. Further, a field experiment showed that dingoes spent less time in the vicinity of prime male dingo scents than other scents. Collectively, these results indicate that age-related scent differences may be discriminable by dingoes. Previous authors have suggested the potential to use scent as a management tool for wild canids by creating an artificial territorial boundary/barrier. Our results suggest that identifying the specific signals in prime-age male scents could facilitate the development of scent-based tools for non-lethal management.

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.

Long-term, climate-driven phenological shift in a tropical large carnivore.

Understanding the degree to which animals are shifting their phenology to track optimal conditions as the climate changes is essential to predicting ecological responses to global change. Species at low latitudes or high trophic levels are theoretically expected to exhibit weaker phenological responses than other species, but limited research on tropical systems or on top predators impedes insight into the contexts in which these predictions are upheld. Moreover, a lack of phenological studies on top predators limits understanding of how climate change impacts propagate through entire ecosystems. Using a 30-y dataset on endangered African wild dogs (Lycaon pictus), we examined changes in reproductive phenology and temperatures during birthing and denning over time, as well as potential fitness consequences of these changes. We hypothesized that their phenology would shift to track a stable thermal range over time. Data from 60 packs and 141 unique pack-years revealed that wild dogs have delayed parturition by 7 days per decade on average in response to long-term warming. This shift has led to temperatures on birthing dates remaining relatively stable but, contrary to expectation, has led to increased temperatures during denning periods. Increased denning temperatures were associated with reduced reproductive success, suggesting that a continued phenological shift in the species may become maladaptive. Such results indicate that climate-driven shifts could be more widespread in upper trophic levels than previously appreciated, and they extend theoretical understanding of the species traits and environmental contexts in which large phenological shifts can be expected to occur as the climate changes.

Real-time drone derived thermal imagery outperforms traditional survey methods for an arboreal forest mammal.

Koalas (Phascolarctos cinereus) are cryptic and currently face regional extinction. The direct detection (physical sighting) of individuals is required to improve conservation management strategies. We provide a comparative assessment of three survey methods for the direct detection of koalas: systematic spotlighting (Spotlight), remotely piloted aircraft system thermal imaging (RPAS), and the refined diurnal radial search component of the spot assessment technique (SAT). Each survey method was repeated on the same morning with independent observers (03:00-12:00 hrs) for a total of 10 survey occasions at sites with fixed boundaries (28-76 ha) in Port Stephens (n = 6) and Gilead (n = 1) in New South Wales between May and July 2019. Koalas were directly detected on 22 occasions during 7 of 10 comparative surveys (Spotlight: n = 7; RPAS: n = 14; and SAT: n = 1), for a total of 12 unique individuals (Spotlight: n = 4; RPAS: n = 11; SAT: n = 1). In 3 of 10 comparative surveys no koalas were detected. Detection probability was 38.9 ± 20.03% for Spotlight, 83.3 ± 11.39% for RPAS and 4.2 ± 4.17% for SAT. Effective detectability per site was 1 ± 0.44 koalas per 6.75 ± 1.03 hrs for Spotlight (1 koala per 6.75 hrs), 2 ± 0.38 koalas per 4.35 ± 0.28 hrs for RPAS (1 koala per 2.18 hrs) and 0.14 ± 0.14 per 6.20 ± 0.93 hrs for SAT (1 koala per 43.39 hrs). RPAS thermal imaging technology appears to offer an efficient method to directly survey koalas comparative to Spotlight and SAT and has potential as a valuable conservation tool to inform on-ground management of declining koala populations.

Addressing inequality and intolerance in human-wildlife coexistence.

Millennia of human conflict with wildlife have built a culture of intolerance toward wildlife among some stakeholders. We explored 2 key obstacles to improved human-wildlife coexistence: coexistence inequality (how the costs and benefits of coexisting with wildlife are unequally shared) and intolerance. The costs of coexisting with wildlife are often disproportionately borne by the so-called global south and rural communities, and the benefits often flow to the global north and urban dwellers. Attitudes and behaviors toward wildlife (tolerance versus intolerance) vary with social and cultural norms. We suggest more empathetic advocacy is needed that, for example, promotes conservation while appropriately considering those who bear the costs of conflict with wildlife. To achieve more equitable cost-sharing, we suggest limiting the costs incurred by those most affected or by sharing those costs more widely. For example, we advocate for the development of improved wildlife compensation schemes, increasing the scale of rewilding efforts, and preventing wildlife-derived revenue leaching out of the local communities bearing the costs of coexistence.

Soluciones para la Desigualdad y la Intolerancia en la Coexistencia Humano - Fauna Resumen Los milenios de conflicto entre los humanos y la fauna han construido una cultura de intolerancia hacia la fauna entre algunos actores. Exploramos dos obstáculos importantes para la mejora de la coexistencia humano - fauna: la desigualdad de coexistencia (cómo los costos y los beneficios de la coexistencia con la fauna están compartidos de una manera desigual) y la intolerancia. Los costos de coexistir con la fauna generalmente están asumidos de manera desproporcional por las llamadas comunidades del sur global o rurales, y los beneficios de convivir con la fauna generalmente fluyen hacia el norte mundial y hacia los habitantes de zonas urbanas. Las actitudes y comportamientos hacia la fauna (tolerancia versus intolerancia) varían con las normas culturales y sociales. Sugerimos la necesidad de una defensa más empática que, por ejemplo, promueva la conservación a la vez que considera de manera apropiada a aquellos que asumen los costos del conflicto con la fauna. Para lograr costos compartidos más equitativos sugerimos limitar los costos incurridos por aquellos más afectados o compartir los costos de manera más amplia. Por ejemplo, abogamos por el desarrollo de esquemas mejorados de compensación de fauna, el incremento de la escala de los esfuerzos por el retorno a la vida silvestre y la prevención del secuestro de ingresos derivados de la fauna fuera de las comunidades locales que asumen los costos de la coexistencia.

Parsimonious test of dynamic interaction.

In recent years, there have been significant advances in the technology used to collect data on the movement and activity patterns of humans and animals. GPS units, which form the primary source of location data, have become cheaper, more accurate, lighter and less power-hungry, and their accuracy has been further improved with the addition of inertial measurement units. The consequence is a glut of geospatial time series data, recorded at rates that range from one position fix every several hours (to maximize system lifetime) to ten fixes per second (in high dynamic situations). Since data of this quality and volume have only recently become available, the analytical methods to extract behavioral information from raw position data are at an early stage of development. An instance of this lies in the analysis of animal movement patterns. When investigating solitary animals, the timing and location of instances of avoidance and association are important behavioral markers. In this paper, a novel analytical method to detect avoidance and association between individuals is proposed; unlike existing methods, assumptions about the shape of the territories or the nature of individual movement are not needed. Simulations demonstrate that false positives (type I error) are rare (1%-3%), which means that the test rarely suggests that there is an association if there is none.

Sneeze to leave: African wild dogs (Lycaon pictus) use variable quorum thresholds facilitated by sneezes in collective decisions.

In despotically driven animal societies, one or a few individuals tend to have a disproportionate influence on group decision-making and actions. However, global communication allows each group member to assess the relative strength of preferences for different options among their group-mates. Here, we investigate collective decisions by free-ranging African wild dog packs in Botswana. African wild dogs exhibit dominant-directed group living and take part in stereotyped social rallies: high energy greeting ceremonies that occur before collective movements. Not all rallies result in collective movements, for reasons that are not well understood. We show that the probability of rally success (i.e. group departure) is predicted by a minimum number of audible rapid nasal exhalations (sneezes), within the rally. Moreover, the number of sneezes needed for the group to depart (i.e. the quorum) was reduced whenever dominant individuals initiated rallies, suggesting that dominant participation increases the likelihood of a rally's success, but is not a prerequisite. As such, the 'will of the group' may override dominant preferences when the consensus of subordinates is sufficiently great. Our findings illustrate how specific behavioural mechanisms (here, sneezing) allow for negotiation (in effect, voting) that shapes decision-making in a wild, socially complex animal society.

The effect of relatedness and pack size on territory overlap in African wild dogs.

BACKGROUND: Spacing patterns mediate competitive interactions between conspecifics, ultimately increasing fitness. The degree of territorial overlap between neighbouring African wild dog (Lycaon pictus) packs varies greatly, yet the role of factors potentially affecting the degree of overlap, such as relatedness and pack size, remain unclear. We used movement data from 21 wild dog packs to calculate the extent of territory overlap (20 dyads). RESULTS: On average, unrelated neighbouring packs had low levels of overlap restricted to the peripheral regions of their 95% utilisation kernels. Related neighbours had significantly greater levels of peripheral overlap. Only one unrelated dyad included overlap between 75%-75% kernels, but no 50%-50% kernels overlapped. However, eight of 12 related dyads overlapped between their respective 75% kernels and six between the frequented 50% kernels. Overlap between these more frequented kernels confers a heightened likelihood of encounter, as the mean utilisation intensity per unit area within the 50% kernels was 4.93 times greater than in the 95% kernels, and 2.34 times greater than in the 75% kernels. Related packs spent significantly more time in their 95% kernel overlap zones than did unrelated packs. Pack size appeared to have little effect on overlap between related dyads, yet among unrelated neighbours larger packs tended to overlap more onto smaller packs' territories. However, the true effect is unclear given that the model's confidence intervals overlapped zero. CONCLUSIONS: Evidence suggests that costly intraspecific aggression is greatly reduced between related packs. Consequently, the tendency for dispersing individuals to establish territories alongside relatives, where intensively utilised portions of ranges regularly overlap, may extend kin selection and inclusive fitness benefits from the intra-pack to inter-pack level. This natural spacing system can affect survival parameters and the carrying capacity of protected areas, having important management implications for intensively managed populations of this endangered species.

An exploratory clustering approach for extracting stride parameters from tracking collars on free-ranging wild animals.

Changes in stride frequency and length with speed are key parameters in animal locomotion research. They are commonly measured in a laboratory on a treadmill or by filming trained captive animals. Here, we show that a clustering approach can be used to extract these variables from data collected by a tracking collar containing a GPS module and tri-axis accelerometers and gyroscopes. The method enables stride parameters to be measured during free-ranging locomotion in natural habitats. As it does not require labelled data, it is particularly suitable for use with difficult to observe animals. The method was tested on large data sets collected from collars on free-ranging lions and African wild dogs and validated using a domestic dog.

Additive opportunistic capture explains group hunting benefits in African wild dogs.

African wild dogs (Lycaon pictus) are described as highly collaborative endurance pursuit hunters based on observations derived primarily from the grass plains of East Africa. However, the remaining population of this endangered species mainly occupies mixed woodland savannah where hunting strategies appear to differ from those previously described. We used high-resolution GPS and inertial technology to record fine-scale movement of all members of a single pack of six adult African wild dogs in northern Botswana. The dogs used multiple short-distance hunting attempts with a low individual kill rate (15.5%), but high group feeding rate due to the sharing of prey. Use of high-level cooperative chase strategies (coordination and collaboration) was not recorded. In the mixed woodland habitats typical of their current range, simultaneous, opportunistic, short-distance chasing by dogs pursuing multiple prey (rather than long collaborative pursuits of single prey by multiple individuals) could be the key to their relative success in these habitats.

Energy cost and return for hunting in African wild dogs and cheetahs.

African wild dogs (Lycaon pictus) are reported to hunt with energetically costly long chase distances. We used high-resolution GPS and inertial technology to record 1,119 high-speed chases of all members of a pack of six adult African wild dogs in northern Botswana. Dogs performed multiple short, high-speed, mostly unsuccessful chases to capture prey, while cheetahs (Acinonyx jubatus) undertook even shorter, higher-speed hunts. We used an energy balance model to show that the energy return from group hunting and feeding substantially outweighs the cost of multiple short chases, which indicates that African wild dogs are more energetically robust than previously believed. Comparison with cheetah illustrates the trade-off between sheer athleticism and high individual kill rate characteristic of cheetahs, and the energetic robustness of frequent opportunistic group hunting and feeding by African wild dogs.

Fine-scale spatiotemporal patterns of genetic variation reflect budding dispersal coupled with strong natal philopatry in a cooperatively breeding mammal.

The relatedness structure of animal populations is thought to be a critically important factor underlying the evolution of mating systems and social behaviours. While previous work has shown that population structure is shaped by many biological processes, few studies have investigated how these factors vary over time. Consequently, we explored the fine-scale spatiotemporal genetic structure of an intensively studied population of cooperatively breeding banded mongooses (Mungos mungo) over a 10-year period. Overall population structure was strong (average F(ST)  = 0.129) but groups with spatially overlapping territories were not more genetically similar to one another than noncontiguous groups. Instead, genetic differentiation was associated with historical group-fission (budding) events, with new groups diverging from their parent groups over time. Within groups, relatedness was high within but not between the sexes, although the latter increased over time since group formation due to group founders being replaced by philopatric young. This trend was not mirrored by a decrease in average offspring heterozygosity over time, suggesting that close inbreeding may often be avoided, even when immigration into established groups is virtually absent and opportunities for extra-group matings are rare. Fine-scale spatiotemporal population structure could have important implications in social species, where relatedness between interacting individuals is a vital component in the evolution of patterns of inbreeding avoidance, reproductive skew and kin-selected helping and harming.

Elevated prolactin levels immediately precede decisions to babysit by male meerkat helpers.

Recent studies suggest that decisions to care for the offspring of others in societies of cooperative vertebrates may have a hormonal basis. The crucial question of whether changes in hormone levels immediately precede or merely follow bouts of offspring care, however, remains largely unanswered. Here, we show that in wild groups of cooperatively breeding meerkats, male helpers that decided to babysit for the day had significantly higher levels of prolactin, coupled with lower levels of cortisol, before initiating a babysitting session compared with similarly aged individuals that decided to forage. In addition, these hormonal differences disappeared over the course of the day, suggesting that hormone levels changed in a fundamentally different way in meerkats that babysat versus those that foraged. In contrast, long-term contributions to babysitting were not significantly associated with plasma levels of prolactin, cortisol, or testosterone in individual male helpers. Our results show, for the first time, that elevated levels of prolactin may immediately precede bouts of helping behavior but differ from recent findings on the same study population in which plasma levels of cortisol, but not prolactin, were significantly and positively associated with rates of pup feeding by male helpers. Together, these results lend significant weight to the idea that decisions to help in cooperative vertebrates have a hormonal basis, although different hormones appear to be associated with different types of care.

Cortisol levels are positively associated with pup-feeding rates in male meerkats.

In societies of cooperative vertebrates, individual differences in contributions to offspring care are commonly substantial. Recent attempts to explain the causes of this variation have focused on correlations between contributions to care and the protein hormone prolactin, or the steroid hormone testosterone. However, such studies have seldom considered the importance of other hormones or controlled for non-hormonal factors that are correlative with both individual hormone levels and contributions to care. Using multivariate statistics, we show that hormone levels explain significant variation in contributions to pup-feeding by male meerkats, even after controlling for non-hormonal effects. However, long-term contributions to pup provisioning were significantly and positively correlated with plasma levels of cortisol rather than prolactin, while plasma levels of testosterone were not related to individual patterns of pup-feeding. Furthermore, a playback experiment that used pup begging calls to increase the feeding rates of male helpers gave rise to parallel increases in plasma cortisol levels, whilst prolactin and testosterone levels remained unchanged. Our findings confirm that hormones can explain significant amounts of variation in contributions to offspring feeding, and that cortisol, not prolactin, is the hormone most strongly associated with pup-feeding in cooperative male meerkats.

Adaptive size modification by dominant female meerkats.

In species of cooperative insects that live in large groups, selection for increased fecundity has led to the evolution of an increased body size among female reproductives, but whether this is also true of cooperative vertebrates is unknown. Among vertebrates, morphological modification of female breeders has only been documented in a single species; in naked mole rats (Heterocephalus glaber), acquisition of alpha status is associated with a significant increase in body size through an elongation of the lumbar vertebrae. Here we provide evidence of morphological modification among breeding females of a cooperative carnivore, the meerkat (Suricata suricatta), and demonstrate that this modification is likely to be adaptive. The same female meerkats were significantly larger when they were dominant than when they were subordinate. This increased body size was not explained by differences in age, foraging efficiency, or investment in offspring care, but may have arisen, in part, through increased levels of hormone that govern bone growth. Increases in body size are likely to result in fitness benefits, for large females delivered larger litters and had heavier offspring, both of which are known to correlate positively with measures of breeding success in meerkats. Our results suggest that the acquisition of alpha status in female meerkats is associated with an adaptive increase in body size and hence that morphological modification of female vertebrates may be more widespread than has been previously supposed.