Achilles' heel of sociality revealed by energetic poverty trap in cursorial hunters

This study empirically tests two foundation ecological theories: (1) pack hunting is a driver for the evolution of sociality; and (2) species have a finite energy potential, whereby increased maintenance costs result in decreased reproductive effort. Using activity and prey data from 22 packs of African wild dogs (Lycaon pictus), we parameterized a model detailing the energetic cost/benefit of cooperative hunting. Larger pack size increased foraging time, prey size, and capture probability while reducing chase distance, resulting in a rapidly increasing net rate of energy intake up to a pack size of five, which peaked at 10 individuals and then declined. With a streamlined body plan necessary for hypercursoriality limiting stomach capacity in smaller packs, it was demonstrated that the group hunting benefit will rather accrue to widely foraging predators than to "sit-and-wait" ones. Reproductive effort, measured by the number of pups born, revealed smaller litters with decreasing pack size, validated finite energy theory, and highlighted a "poverty trap" where smaller groups have lower foraging gains, smaller litters, and increased vulnerability to extirpation. Consequently, these results demonstrated a mechanistic example of pervasive selection for maximal body size (Cope's rule), leading to a macroevolutionary ratchet, where sociality linked to hypercursoriality is betrayed by an Achilles' heel.     

Does captivity influence territorial and hunting behaviour? Assessment for an ex situ reintroduction program of African lions Panthera leo

Maintaining a territory and being able to hunt are imperative for the success of African lion Panthera leo prides. We aimed to determine whether captive‐origin prides display similar territorial and hunting behaviour to wild lions. Behaviours and locations of two captive‐origin prides and one wild pride were collected through direct observation. All prides established territories, and core areas corresponded to resource requirements. There was no evidence that pride origin affected territorial or hunting behaviour. Captive‐origin prides exhibited behaviours that lead us to be optimistic about each pride's ability to establish and defend a territory successfully, and to hunt, following reintroduction.     

Territory quality determines social group composition in Ethiopian wolves Canis simensis

1. We contrast the value of four different models to predict variation in territory size as follows: resource density (the ideal free distribution), population density, group size and intruder pressure (relative resource-holding potential). In the framework of the resource dispersion hypothesis, we test the effect of resource abundance and spatial variation in resource distribution on the age/sex composition of social groups. 2. We explore these drivers of territory size and group size/composition in Ethiopian wolves Canis simensis in the Bale Mountains, Ethiopia, using fine-scale distribution maps of their major prey species based on satellite-derived vegetation maps. 3. The number of adult males is correlated with territory size, while prey density, wolf population density and intruder pressure are not associated with territory size. On average, each additional adult male increases territory size by 1.18 km(2). 4. Prey abundance increases with territory size (average biomass accumulation of 6.5 kg km(-2)), and larger territories provide greater per capita access to prime foraging habitat and prey. 5. The age/sex composition of wolf packs is more closely related to territory quality than territory size. Subordinate adult females are more likely to be present in territories with greater proportions of prime giant molerat Tachyoryctes macrocephalus habitat (i.e. >80% of Web Valley territories and >20% in Sanetti/Morebawa), and more yearlings (aged 12-23 months) occur in territories with greater overall prey biomass. 6. Wolf packs with restricted access to good foraging habitat tend to defend more exclusive territories, having a lower degree of overlap with neighbouring packs. 7. The greater per capita access to prey in large groups suggests a strong evolutionary advantage of collaborative territorial defence in this species, although the relative costs of territorial expansion vs. exclusion depend upon the spatial distribution of resources. We propose a model whereby territory size is determined by the number of adult males, with the presence of subordinate females and yearlings dependent on the quality of habitat, and the abundance and distribution of prey, incorporated within territory boundaries.     

Fence management and time since pack formation influence African wild dog escapes from protected areas in South Africa

In human-dominated and highly fragmented landscapes, keeping wildlife within reserve boundaries is vital for conservation success. In South Africa, fences are a widely employed conservation management tool for protected areas and are successful in mitigating human-wildlife conflict. However, fences are permeable, and predators are able to cross through reserve fences. African wild dogs (Lycaon pictus) often leave fenced boundaries, resulting in high capture and translocation costs. Moreover, when wild dog packs (up to 30 individuals) leave fenced reserves they enter human-dominated landscapes where they face strong persecution and livestock predation incurs high costs. The factors driving packs to leave managed reserves are poorly understood, thus, to effectively manage wild dogs in fenced systems, it is important to understand why they leave reserve boundaries. There are several hypotheses as to why wild dogs cross through reserve fences, including inter- and intra-specific competition, social behaviour, management, prey density and environmental variability. Using a long-term dataset comprising 32 resident packs across five reserves, we investigated the relative strength of these hypotheses on the probability of wild dogs exiting a fenced reserve. During the 14-year study period, we recorded 154 exit events. We found that the interaction of fence integrity and time since pack formation were the primary factors affecting the probability of a pack leaving a reserve. When fence integrity was poor, escape probability decreased with pack age likely due to the exploratory behaviour of new packs. When fence integrity was average, escape probability increased with pack age likely due to the fitness benefits of holding larger and more exclusive territories as packs age. When fence integrity was good, the probability of a pack escaping was very low (only 1% occurrence). The implications of this research suggest that the primary management consideration for reducing wild dog escapes from fenced reserves should be maintaining adequate reserve-wide fence integrity, rather than focusing on social structure or drivers of inter- and intra-specific competition.     

Conservation implications of prey responses to wild dogs Lycaon pictus during the denning season on wildlife ranches

The spread of game ranching in southern Africa provides opportunities for the reestablishment of populations of endangered wild dogs extirpated by livestock ranchers. However, this potential has not been realized, partly because of negative rancher perceptions. Some ranchers believe that wild dogs impart costs by killing wildlife that could be utilized consumptively. Others complain that wild dogs make ungulates 'skittish' and cause local reductions in prey densities while denning. We compared the skittishness and density of prey species inside and outside the denning home ranges of nine wild dog packs in Zimbabwe. Wild dogs had no impact on prey skittishness, but prey species did occur at lower densities inside denning home ranges. In some scenarios, and particularly on fenced game ranches, wild dogs could cause prey population declines during denning. On small game ranches, the use of fences as a tool by wild dogs during hunting can increase the proportion of large prey species in their diet by up to 11 times, and thus increase the minimum area required to support that diet. In addition, game fencing is typically permeable to wild dogs but not their prey, preventing the recovery of prey populations through the natural influx of prey animals into the denning area following departure of the dogs. Wild dogs could thus impose significant financial costs to game ranchers hosting denning packs. Our findings emphasize the importance of promoting the formation of conservancies, where neighbouring landowners remove boundary fences to create larger contiguous wildlife areas.     

The effects of anthrax on endangered African wild dogs (Lycaon pictus)

African wild dogs ( Lycaon pictus ) are large, social canids formerly found throughout sub-Saharan Africa. Wild dogs are now endangered; only six nations hold populations larger than 100 individuals. Considerable evidence suggests that disease plays a central role in the regulation of some wild dog populations. In particular, it has been suggested that epizootics of rabies, canine distemper and anthrax can have strong local effects on wild dog numbers. Resolution of the regulatory importance of these diseases has been hampered by lack of data from wild populations. Here we report on an outbreak of anthrax among wild dogs in the Selous Game Reserve in southern Tanzania, describing clinical signs and diagnosis, measuring mortality, and testing for effects on hunting success and movements. Only one of five packs under study was affected. Immediate mortality was mild, with none of 18 adults (0%) and four of 24 pups (17%) dying. Mortality was significantly higher among puppies (<1 year old) than among adults. Mortality of individuals that showed signs of disease but did not die immediately was not significantly elevated over the following six months. The hunting success and movement patterns of the pack were not affected by the outbreak. Collectively, these data suggest that African wild dogs possess a degree of resistance to anthrax.     

Implications of African wild dog (Lycaon pictus) denning on the density and distribution of a key prey species: addressing myths and misperceptions

African wild dogs (Lycaon pictus) are often the least popular large carnivore among game ranchers because of their perceived impact on prey populations. Landowner perceptions include that wild dogs greatly deplete prey during their three-month denning period, take prey that could otherwise be sold for hunting and cause prey to move away from the vicinity of their den sites. Landowners’ tolerance towards African wild dogs could thus be improved with a more rigorous understanding of the actual impact of wild dogs on prey populations during the denning period. Using impala density data and wild dog denning records from Sango Ranch in the Savé Valley Conservancy, Zimbabwe, we compared impala densities between pre-denning, denning and post-denning season and between inside and outside the denning home range. Our results indicate that wild dog denning does not cause a significant local reduction in prey around the den and does not cause prey to move away from denning areas. However prey species did occur in lower density inside the denning home ranges than outside, in all seasons. This result indicates that wild dogs select dens in areas of lower prey density, perhaps as an avoidance mechanism for lions. Accordingly, contrary to what landowners believe, wild dogs do not have a significant impact on prey populations during their denning season.     

TErritorial defence and mating success in males of the small heath butterfly, coenonympha pamphilus L. (Lepidoptera: Satyridae)

It has long been recognized that territorial defence in male butterflies must be some sort of mating strategy and that the territories are used as mating stations. However, so far, no systematic study has established the adaptive significance of territorial behaviour. This study is an attempt to fill this gap. By comparing the distribution of territories with the distribution of locations where wild and released virgin females mated this study shows that, in the small heath butterfly (Coenonympha pamphilus), males in territories have higher mating success than males outside territories. This supported the hypotheses that the function of territorial behaviour is to secure more matings and that the territories are mating stations. Wing length measurements suggest the same. Large males residing in territories tended to mate more often than small males, which were usually found outside territories. Since resident males were larger than non-resident males, this size difference was used to see how territorial occupancy influenced longevity. Mark-recapture of measured males revealed no significant correlation between wing length and the further life expectancy of males, strongly suggesting that the mating success of males in territories is also higher when measured over their whole lifetime.     

Habitat selection and polygyny in breeding Corn Buntings Miliaria calandra

Unlike many other polygynous passerine species, female Corn Buntings Miliaria calandra apparently do not suffer costs by pairing polygynously, yet it is unclear whether this is because polygynous males hold the highest quality territories or because pairing with polygynous males is unimportant in determining female reproductive success. Male Corn Buntings on North Uist, Scotland, consistently defended territories which contained nesting habitat, and females often foraged outside male territories when provisioning nestlings. Females showed strong preferences for nesting in uncultivated land, and 80% of nests were under Hogweed Heracleum sphondylium, possibly because this provided cover against predation and the weather. When provisioning nestlings, females showed strong preferences for foraging in cereal crops, probably because this habitat provided better food resources and/or better cover from predators. Males were unpaired or paired with one to three females per breeding season, but variation in territory size or vegetation composition did not explain differences in the number of females paired with individual males. We suggest that when females neither gain benefits nor suffer costs by breeding polygynously, and males do not differ greatly in the areas of habitat selected, polygyny can arise through random female settlement within the nesting habitat.     

Territorial defense against various food competitors in the Tanganyikan benthophagous cichlid <Emphasis Type="Italic">Neolamprologus tetracanthus</Emphasis>

Territorial defense of nonbreeding female Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid, was investigated. Females defended territories (=home ranges, ca. 1m across) against a variety of intruding fishes. Conspecific females were usually attacked outside the territories, heterospecific benthivores (shrimp eaters) and omnivores near the border of the territories, and piscivores, algae and detritus feeders, and herbivores inside the territories. Females used some parts of the sandy substrate in the territories for foraging (foraging areas). Territorial defense prevented most of the conspecific females and benthivores from intruding into the foraging areas. In omnivores, piscivores, and algae and detritus feeders, about half the intruders were repelled from the foraging areas, although herbivores were infrequently repelled in the areas. Soon after removal of the resident females, many food competitors invaded the foraging areas and eagerly devoured prey, suggesting that the territories are maintained for food resource protection from these competitors. Females are likely to discriminate intruding fishes and change their territorial defense primarily on the basis of the degree of dietary overlap, resulting in monofunctional serial territories.     

Spatial and temporal patterns of neutral and adaptive genetic variation in the endangered African wild dog (Lycaon pictus)

Deciphering patterns of genetic variation within a species is essential for understanding population structure, local adaptation and differences in diversity between populations. Whilst neutrally evolving genetic markers can be used to elucidate demographic processes and genetic structure, they are not subject to selection and therefore are not informative about patterns of adaptive variation. As such, assessments of pertinent adaptive loci, such as the immunity genes of the major histocompatibility complex (MHC), are increasingly being incorporated into genetic studies. In this study, we combined neutral (microsatellite, mtDNA) and adaptive (MHC class II DLA‐DRB1 locus) markers to elucidate the factors influencing patterns of genetic variation in the African wild dog (Lycaon pictus); an endangered canid that has suffered extensive declines in distribution and abundance. Our genetic analyses found all extant wild dog populations to be relatively small (N_e < 30). Furthermore, through coalescent modelling, we detected a genetic signature of a recent and substantial demographic decline, which correlates with human expansion, but contrasts with findings in some other African mammals. We found strong structuring of wild dog populations, indicating the negative influence of extensive habitat fragmentation and loss of gene flow between habitat patches. Across populations, we found that the spatial and temporal structure of microsatellite diversity and MHC diversity were correlated and strongly influenced by demographic stability and population size, indicating the effects of genetic drift in these small populations. Despite this correlation, we detected signatures of selection at the MHC, implying that selection has not been completely overwhelmed by genetic drift.     

Small pack size imposes a trade-off between hunting and pup-guarding in the painted hunting dog Lycaon pictus

The painted hunting dog or African wild dog, Lycaon pictus,is one of the most endangered large carnivores in Africa, withextinction predicted within a few decades if their dramaticdecline is not stopped. It has recently been hypothesized thatbecause of their constraining need for helpers, group sizewas of major importance in obligate cooperative breeding species,and that the resulting likely existence of a threshold numberof adults could create an Allee effect, increasing the groupextinction risk. One example where the importance for a criticalnumber of adults may have major repercussions for painted huntingdogs concerns baby-sitting, or pup-guarding, a behavior typicalof obligate cooperative breeders. We propose that, as forgoingthis behavior is costly because pup guards have the potentialto decrease pup mortality, its use is costly too, especiallyin small packs, because helpers are strongly needed for theircooperative foraging (hunting, protecting the kill and bringingback food to the pups). We present a simple model showing howpup-guarding imposes a cost because it implies that less foodper hunt is brought back to more individuals at the den. Wecomplete these analyses with empirical tests of the effectof pack size on the probability of pup-guarding, from fielddata from the Hwange population in Zimbabwe. Our model, aswell as our 5 years of empirical data, both suggest a criticalthreshold at a size of about five individuals.     

Effects of habitat use on food acquisition and food caching during a nonbreeding season in a winter-breeding,food-storing passerine

ABSTRACT

Capsule: Wintering male Bull-headed Shrikes Lanius buchepalus preferred vegetable fields with perch sites to search for and detect terrestrial prey, and males occupying territories with large areas of vegetable fields acquired more prey and cached more food.

Aims: To better understand effects of habitat use on food acquisition and food caching of the Bull-headed Shrike, we investigated relationships between habitat quality (measured through foraging-site selection and foraging success) and food caching during the non-breeding season.

Methods: We monitored 66 territorial male shrikes during the non-breeding season from 2014 to 2016, and collected data on foraging-site selection, foraging success, and food caching.

Results: Our field observations showed that male shrikes preferred to forage over vegetable fields and that males occupying territories incorporating large areas of that habitat were able to acquire more food items and store more food caches in their territories during the nonbreeding season.

Conclusion: We suggest that for male Bull-headed Shrikes, a winter-breeding food-storing passerine, the quality of habitat in the nonbreeding season has the potential to affect their subsequent fitness.     

Defence, intrusion and the evolutionary stability of territoriality

Territorial behaviour can only be adaptive if its costs are outweighed by its benefits. Territorial individuals incur costs by defending their territories against intruders. Usually these intruders are assumed to be non-territorial floaters attempting to take over the whole territory or neighbours trying to extend the borders of their own territory. We instead investigate how costs and benefits of territorial behaviour are affected by neighbours which invade to steal resources on a territory.We show analytically that in the absence of defence intrusion into neighbouring territories always pays and that even if territories are defended intrusion levels can still be high. Using a more detailed simulation model we find that territory defence usually disappears from the population even if owners have a strong advantage over intruders in terms of fighting costs or foraging efficiency. Defence and thus territoriality can only be evolutionarily stable if fighting costs for the intruder relative to the productivity of the territory are very high or if crossing the borders between territories carries additional costs.Our results show that stealing of resources by neighbours can have a considerable effect on the evolutionary stability of territory defence and thus territoriality itself. A more mechanistic model of territorial behaviour is needed to incorporate these kinds of mechanisms into a general theory on the evolution of territoriality.     

Territorial behavior of Western Sandpipers on their nonbreeding grounds: effect of sex and foraging interference

ABSTRACT Nonbreeding shorebirds often alternate social structure between anonymous flocks and territorial behavior in response to different environmental factors. To evaluate specific drivers for one species, we studied the spacing behavior of wintering Western Sandpipers (Calidris mauri) at Bahía Santa María, northwestern Mexico, using behavioral observations. The density, population structure, and territorial behavior of Western Sandpipers differed among three habitat types. Cattail marshes supported 110 birds per ha, 49% males, and no territorial birds. Mangrove flats supported 288 birds per ha, 58% males, and 5% territorial birds. Brackish flats supported 365 birds per ha, 76% males, and 7% territorial birds. Territories consisted of rectangular strips (5–9 m long, N= 77). Territory length was not related to either bird density or number of territorial birds by plot, but was positively related to nearest bird distance. Aggression rate was inversely related to territory length, suggesting that territory length is set by the costs of defense. Foraging rate was independent of territory length, and prey densities in territories did not differ from those in areas used by nonterritorial birds. Males were more likely to be territorial and had a higher aggression rate than females, suggesting that males, which forage more on surface prey, were more affected by foraging interference. Our results suggest that the territorial behavior of Western Sandpipers in our study was an opportunistic strategy to reduce foraging interference. The variation in spacing behavior we documented provides evidence that interference competition affects the social structure of Western Sandpipers during the nonbreeding season.     

Genetic diversity and spatial genetic structure of African wild dogs (<Emphasis Type="Italic">Lycaon pictus</Emphasis>) in the Greater Limpopo transfrontier conservation area

The Greater Limpopo Transfrontier Conservation Area (GLTFCA) is one of the last refuges for the endangered African wild dog and hosts roughly one-tenth of the global population. Wild dogs in this area are currently threatened by human encroachment, habitat fragmentation and scarcity of suitable connecting habitat between protected areas. We derived genetic data from mitochondrial and nuclear markers to test the following hypotheses: (i) demographic declines in wild dogs have caused a loss of genetic variation, and (ii) Zimbabwean and South African populations in the GLTFCA have diverged due to the effects of isolation and genetic drift. Genetic patterns among five populations, taken with comparisons to known information, illustrate that allelic richness and heterozygosity have been lost over time, presumably due to effects of inbreeding and genetic drift. Genetic structuring has occurred due to low dispersal rates, which was most apparent between Kruger National Park and the Zimbabwean Lowveld. Immediate strategies to improve gene flow should focus on increasing the quality of habitat corridors between reserves in the GLTFCA and securing higher wild dog survival rates in unprotected areas, with human-mediated translocation only undertaken as a last resort.     

Hunting success of lions affected by the moon's phase in a wooded habitat

Variation in moonlight affects foraging, hunting and vigilance behaviours in many nocturnal species. Here, we explore the effect of moonlight on the movement and hunting behaviour of African lions. Previous studies found bright moonlight is associated with reduced hunting success; however, those studies were largely undertaken in open habitats where predators are easily seen by prey species on moonlit nights. In this study, we explored whether moonlight affected hunting behaviour and success in a largely wooded environment. Measures of short‐term lion movements (distance moved, displacement and path tortuosity) derived from GPS telemetry data were used as indicators of movement behaviour. Field observations of belly distension were used to assess recent food intake. Lions had greater belly distension (indicating feeding success) on dark nights. However, this change in feeding success was not reflected by lion movement patterns—there was no evidence that these changed across moon phases. There was no evidence that lions used more covered habitats on brighter nights to facilitate concealment.     

The Effect of Simulated African Wild Dog Presence on Anti‐predator Behaviour of Kudu and Impala

In this study, we examined the behavioural, temporal and spatial effects of simulated African wild dog (Lycaon pictus) presence on its two main prey species: kudu (Tragelaphus strepsiceros) and impala (Aepyceros melampus). We spread African wild dog faeces around waterholes and played African wild dog sounds at different intervals to mimic immediate and non‐immediate predation pressure. We looked at anti‐predator behaviour at both a herd and individual level and distinguished between high‐quality (detracts from all other activities), high‐cost vigilance and low‐quality (used to monitor the surrounding in spare time), low‐cost vigilance to determine costs involved. We found that simulated African wild dog presence had little effect on anti‐predator behaviour of their free‐ranging prey. Only when immediate predation risk was mimicked did kudu invest in (additional) high‐quality vigilance, whereas impala showed no response. Regardless of direct cues of African wild dog presence, behavioural adjustments to reduce predation risk were primarily based on environmental factors such as time of the day and broad‐scale habitat structure. Predators have been shown to utilize waterholes to hunt, and prey species are therefore likely to maximize anti‐predator behaviour in this high‐risk environment based on environmental variables affecting predation risk, the main predator within the system, and water requirements, leaving little flexibility to respond to (simulated) African wild dog presence.     

Diet composition and foraging success in generalist predators: Are specialist individuals better foragers?

Factors affecting individual diet specialization in generalist populations and the relationship between diet and foraging success remain poorly studied, particularly in terrestrial wide-ranging predators. We studied whether individual variations in diet in Montagu's harrier males (determined through a combination of direct foraging observations and pellet analysis) were associated with patterns of foraging habitat selection and foraging success of 12 radiotracked males during the breeding period. We found important differences in diet composition and breadth between individuals. Diet diversity was negatively related to hunting success: the most efficient individuals in terms of hunting success had the most specialized diet. This study also suggests an important role of individual foraging habitat selection in explaining individual diet, as the proportion of different prey types in the diet was associated with habitat composition within the home range, with higher proportion of those habitats that held higher abundances of their more frequent prey. This study thus provides evidence of individual diet specialization having a knock-on effect on foraging efficiency in a wide-ranging raptor and highlights the role of individual behaviour as a driving force of intra-population niche variation.