Hunting income patterns among the Hadza: big game, common goods, foraging goals and the evolution of the human diet.

The assumption that large mammal hunting and scavenging are economically advantageous to hominid foragers is examined in the light of data collected among the Hadza of northern Tanzania. Hadza hunters disregard small prey in favour of larger forms (mean adult mass greater than or equal to 40 kg). Here we report experimental data showing that hunters would reduce their mean rates if they included small animals in the array they target. Still, daily variance in large animal hunting returns is high, and the risk of failure correspondingly great, significantly greater than that associated with small game hunting and trapping. Sharing large kills reduces the risk of meatless days for big game hunters, and obviates the problem of storing large amounts of meat. It may be unavoidable if large carcasses cannot be defended economically against the demands of other consumers. If so, then large prey are common goods. A hunter may gain no consumption advantage from his own big game acquisition efforts. We use Hadza data to model this 'collective action' problem, and find that an exclusive focus on large game with extensive sharing is not the optimal strategy for hunters concerned with maximizing their own chances of eating meat. Other explanations for the emergence and persistence of this practice must be considered.     

Predator size and prey size–gut capacity ratios determine kill frequency and carcass production in terrestrial carnivorous mammals

Carnivore kill frequency is a fundamental part of predator–prey interactions, which are important shapers of ecosystems. Current field kill frequency data are rare and existing models are insufficiently adapted to carnivore functional groups. We developed a kill frequency model accounting for carnivore mass, prey mass, pack size, partial consumption of prey and carnivore gut capacity. Two main carnivore functional groups, small prey‐feeders versus large prey‐feeders, were established based on the relationship between stomach capacity (C) and pack corrected prey mass (iMprey). Although the majority of small prey‐feeders is below, and of large prey‐feeders above a body mass of 10–20 kg, both occur across the whole body size spectrum, indicating that the dichotomy is rather linked to body size‐related ecology than physiology. The model predicts a negative relationship between predator size and kill frequency for large prey‐feeders. However, for small prey‐feeders, this negative relationship was absent. When comparing carnivore prey requirements to estimated stomach capacity, small carnivores may have to eat to their full capacity repeatedly per day, requiring fast digestion and gut clearance. Large carnivores do not necessarily have to eat to full gastric capacity per day, or do not need to eat every day, which in turn reduces kill frequencies or drives other ecological processes such as scavenging, kleptoparasitism, and partial carcass consumption. Where ecological conditions allow, large prey‐feeding appears attractive for carnivores, which can thus reduce activities related to hunting. This is particularly so for large carnivores, who can achieve distinct reductions in hunting activity due to their relatively large gut capacity.     

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.     

Aposematism: what should our starting point be?

The evolution of aposematism is considered to be a major evolutionary problem because if new aposematic forms emerged in defended cryptic populations, they would face the dual problems of rarity and conspicuousness. We argue that this commonly assumed starting point might not have wide validity. We describe a novel evolutionary computer model in which prey evolve secondary defences and become conspicuous by moving widely over a visually heterogeneous habitat. Unless crypsis imposes high opportunity costs (for instance, preventing prey from efficient foraging, thermoregulation and communication), costly secondary defences are not predicted to evolve at all. However, when crypsis imposes opportunity costs, prey evolve secondary defences that facilitate raised behavioural conspicuousness as prey exploit opportunities within their environment. Optimal levels of secondary defence and of behavioural conspicuousness increase with population sizes and the costs imposed by crypsis. When prey are already conspicuous by virtue of their behaviours, the evolution of aposematic appearances (bright coloration, etc.) is much easier to explain because aposematic traits add little further costs of conspicuousness, but can bring large benefits.     

Prey preferences of the leopard (Panthera pardus)

Leopards Panthera pardus have a catholic diet and are generally thought to prey on medium-sized ungulates; however, knowledge on which species are actually preferred and avoided is lacking, along with an understanding of why such preferences arise. Twenty-nine published and four unpublished studies of leopard diet that had relative prey abundance estimates associated with them were analysed from 13 countries in 41 different spatial locations or temporal periods throughout the distribution of the leopard. A Jacobs' index value was calculated for each prey species in each study and the mean of these was then tested against a mean of 0 using t or sign tests for preference or avoidance. Leopards preferentially prey upon species within a weight range of 10–40 kg. Regression plots suggest that the most preferred mass of leopard prey is 25 kg, whereas the mean body mass of significantly preferred prey is 23 kg. Leopards prefer prey within this body mass range, which occur in small herds, in dense habitat and afford the hunter minimal risk of injury during capture. Consequently, impala, bushbuck and common duiker are significantly preferred, with chital likely to also be preferred with a larger sample size from Asian sites. Species outside the preferred weight range are generally avoided, as are species that are restricted to open vegetation or that have sufficient anti-predator strategies. The ratio of mean leopard body mass with that of their preferred prey is less than 1 and may be a reflection of their solitary hunting strategy. This model will allow us to predict the diet of leopards in areas where dietary information is lacking, also providing information to assist wildlife managers and conservation bodies on predator carrying capacity and predator–prey interactions.     

The cost of parental care: prey hunting in a digger wasp

Trivers's concept of parental investment is an integral partof modern evolutionary biology. "Parental investment" is definedas any parental expenditure that benefits a current progenyat the expense of a parent's ability to reproduce in the future.Because future costs are hard to quantify, other currencieswere used that were thought to be related to the actual costs.However, the validity of these alternative measures has rarely been established, at least in insects. Specifically, these measureswere not shown to represent costs at all. We investigated provisioningbehavior in a sphecid wasp, the European beewolf, Philanthustriangulum F., and tested whether prey hunting entails futurecosts to the female wasp and thus represents parental investment.We increased as well as decreased the females' hunting effortexperimentally and determined their hunting success on the following day. Furthermore, we analyzed the correlation betweenhunting rate of unrestricted females and their life span andassessed the effect of an experimentally decreased huntingeffort on life span. The future rate of bee hunting decreasedwhen hunting expenditure was increased (in the field) and viceversa (both in the field and in the laboratory). In contrast, there was no trade-off between hunting rate and life span, andlife span was not affected by an experimentally decreased huntingeffort (in the laboratory). Because prey hunting entails costsin terms of a reduced rate of prey hunting in the future, itmeets Trivers' definition of parental investment.     

Potential impact of size-selective planktivory on phosphorus release by zooplankton

A laboratory model of phosphorus release by Daphnia rosea was implemented for published zooplankton data from lakes characterized by different abundances of size-selective planktivores. Size-selective feeding reduces average prey size and increases P release per unit biomass. At the system level, decreased prey standing crop associated with higher planktivore abundance could balance the size dependent increase in P release rate. However, estimates of both net reduction and net increase in rate of P release from zooplankton resulted from model application. Size-selective feeding might be important not only in energetic or evolutionary relationships between predator and prey but also in determination of the relative importance of different pathways of phosphorus flow through pelagic systems.     

Body size,energetic and foraging mode of raptors in central Chile

Summary An inferential analysis of the foraging mode (opportunist or mediate by prey selection) of a taxonomic assemblage of raptors in central Chile was conducted. The analysis of energetic aspects such as daily requirements of predators and energy supplied by the preys, with estimations of prey relative abundances and their incidence in the diet of raptors, led us to conclude that contrarly to the opportunistic hunting mode suggested by others authors, these predators apparently present prey selection (on a biomass basis). This phenomenon is particularly evident in raptors of small body size.     

Hunted Woolly Monkeys (Lagothrix poeppigii) Show Threat-Sensitive Responses to Human Presence

Responding only to individuals of a predator species which display threatening behaviour allows prey species to minimise energy expenditure and other costs of predator avoidance, such as disruption of feeding. The threat sensitivity hypothesis predicts such behaviour in prey species. If hunted animals are unable to distinguish dangerous humans from non-dangerous humans, human hunting is likely to have a greater effect on prey populations as all human encounters should lead to predator avoidance, increasing stress and creating opportunity costs for exploited populations. We test the threat sensitivity hypothesis in wild Poeppigi''s woolly monkeys (Lagothrix poeppigii) in Yasuní National Park, Ecuador, by presenting human models engaging in one of three behaviours “hunting”, “gathering” or “researching”. These experiments were conducted at two sites with differing hunting pressures. Visibility, movement and vocalisations were recorded and results from two sites showed that groups changed their behaviours after being exposed to humans, and did so in different ways depending on the behaviour of the human model. Results at the site with higher hunting pressure were consistent with predictions based on the threat sensitivity hypothesis. Although results at the site with lower hunting pressure were not consistent with the results at the site with higher hunting pressure, groups at this site also showed differential responses to different human behaviours. These results provide evidence of threat-sensitive predator avoidance in hunted primates, which may allow them to conserve both time and energy when encountering humans which pose no threat.     

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.     

Conspicuous visual signals do not coevolve with increased body size in marine sea slugs

Many taxa use conspicuous colouration to attract mates, signal chemical defences (aposematism) or for thermoregulation. Conspicuousness is a key feature of aposematic signals, and experimental evidence suggests that predators avoid conspicuous prey more readily when they exhibit larger body size and/or pattern elements. Aposematic prey species may therefore evolve a larger body size due to predatory selection pressures, or alternatively, larger prey species may be more likely to evolve aposematic colouration. Therefore, a positive correlation between conspicuousness and body size should exist. Here, we investigated whether there was a phylogenetic correlation between the conspicuousness of animal patterns and body size using an intriguing, understudied model system to examine questions on the evolution of animal signals, namely nudibranchs (opisthobranch molluscs). We also used new ways to compare animal patterns quantitatively with their background habitat in terms of intensity variance and spatial frequency power spectra. In studies of aposematism, conspicuousness is usually quantified using the spectral contrast of animal colour patches against its background; however, other components of visual signals, such as pattern, luminance and spectral sensitivities of potential observers, are largely ignored. Contrary to our prediction, we found that the conspicuousness of body patterns in over 70 nudibranch species decreased as body size increased, indicating that crypsis was not limited to a smaller body size. Therefore, alternative selective pressures on body size and development of colour patterns, other than those inflicted by visual hunting predators, may act more strongly on the evolution of aposematism in nudibranch molluscs.     

What Enables Size-Selective Trophy Hunting of Wildlife?

Although rarely considered predators, wildlife hunters can function as important ecological and evolutionary agents. In part, their influence relates to targeting of large reproductive adults within prey populations. Despite known impacts of size-selective harvests, however, we know little about what enables hunters to kill these older, rarer, and presumably more wary individuals. In other mammalian predators, predatory performance varies with knowledge and physical condition, which accumulates and declines, respectively, with age. Moreover, some species evolved camouflage as a physical trait to aid in predatory performance. In this work, we tested whether knowledge-based faculty (use of a hunting guide with accumulated experience in specific areas), physical traits (relative body mass [RBM] and camouflage clothing), and age can predict predatory performance. We measured performance as do many hunters: size of killed cervid prey, using the number of antler tines as a proxy. Examining ∼4300 online photographs of hunters posing with carcasses, we found that only the presence of guides increased the odds of killing larger prey. Accounting for this effect, modest evidence suggested that unguided hunters presumably handicapped with the highest RBM actually had greater odds of killing large prey. There was no association with hunter age, perhaps because of our coarse measure (presence of grey hair) and the performance trade-offs between knowledge accumulation and physical deterioration with age. Despite its prevalence among sampled hunters (80%), camouflage had no influence on size of killed prey. Should these patterns be representative of other areas and prey, and our interpretations correct, evolutionarily-enlightened harvest management might benefit from regulatory scrutiny on guided hunting. More broadly, we suggest that by being nutritionally and demographically de-coupled from prey and aided by efficient killing technology and road access, wildlife hunters in the developed world might have overcome many of the physical, but not knowledge-based, challenges of hunting.     

Evidence of prey depletion around lesser kestrel Falco naumanni colonies and its short term negative consequences

Predation may reduce prey numbers in such extent that prey may be depleted, which has negative effects on predator populations. Prey depletion would be more likely when the number of predators increase and/or concentrate their activity in a certain area, as is the case of colonial birds. As a matter of fact, food depletion due to intraspecific competition is considered a major cost of coloniality, and several studies have shown indirect evidence of this. However, no direct measures of food depletion have been provided along with its consequences for the fitness of the colony inhabitants. We carried out a field study with the lesser kestrel Falco naumanni, a raptor that breeds in colonies ranging from two to dozens of pairs. During the nestling period we sampled the main prey of the kestrels around different sized colonies at increasing distances. At the same time, we recorded hunting distances and prey delivery rates to the nest. In addition, we monitored the reproductive success in colonies of different sizes. Lesser kestrels feed their nestlings mainly with grasshoppers and these prey became depleted through the season in the surroundings of the large colonies. Prey depletion made kestrels fly longer distances to forage and prey delivery rates to the nest decreased. Lower feeding rates were not compensated by bringing larger prey, hence, the net amount of energy provided to the chicks decreased with the date in large colonies. By contrast, none of this occurred around small colonies, where both prey abundance and hunting distance remained constant throughout the season. As a consequence, the seasonal decline in the reproductive success (number of fledglings and fledgling body condition) was greater the larger the colony. Thus, these results evidence that food depletion and its fitness costs are related to colony size, as they are suffered by the kestrels breeding in large colonies but not by those settled at small ones. Finally, the consequences of prey depletion on the demographic dynamics and the regulation of colony size are discussed.     

Predation on mammals by the chimpanzee (Pan troglodytes)

With respect to prey selectivity and predation frequency, chimpanzees (Pan troglodytes) show local differences as well as diachronic variability within the same population. When data on predation from three long-term studies at Mahale, Gombe, and Tai are compared, some differences and similarities emerge; Mahale is more like Gombe than Tai in regard of prey selection but features of hunting at Tai with respect to predation frequency are not conspicuous. The most responsible factor for diversity in prey selectivity is a distinct “prey image” maintained by chimpanzees of different populations, although it is necessary to clarify in future studies why and how such tradition develops. Relative body size of chimpanzees to prey species and/or the degree of cooperation among members of a hunting party may explain the variability in prey size selected at each site, the latter influencing the frequency of successful hunts at the same time. Although various degrees of habituation and different sampling methods including artificial feeding might have obscured the real differences, recent data from the three populations do not seem to be biased greatly by such factors. Nevertheless, it is still difficult to make strict comparisons due to the lack of sufficient standardized data across the three populations on the frequency of hunting and predation. It is suggested that the size or demographic trend of a chimpanzee unit-group, especially the number of adult males included, necessarily influences its hunting frequency as well as its prey profile. It is also suggested that factors which bring these males together into a party (e.g. fruit abundance, swollen females, conflict between unit-groups etc.) strongly affect theactual hunting and kill rates. Other possible factors responsible for the local differences are forest structure (e.g. tree height), skilful “hero” chimpanzees, and competition with sympatric carnivorous animals. A total of at least 32 species have been recorded as prey mammals of chimpanzees from 12 study sites and the most common prey mammals are primates (18 species), of which 13 species are forest monkeys. Forest monkeys, colobine species in particular, are often the most common victims of the predation by chimpanzees at each site. We may point out a tendency toward selective hunting for the forest monkeys in terms of the selectivity of prey fauna among all three subspecies of chimpanzees, including populations living in drier environment. The mode of chimpanzee hunting seems to correspond to the highest available biomass of gregarious, arboreal monkeys in the forest, colobine species in particular. In contrast, bonobos (P. paniscus) are less carnivorous than chimpanzees, only rarely preying on a few species of small mammals. The sharp contrast of the two allied species in their predatory tendencies appears to have something to do with the differences in the structure of primary production between their habitats.     

Predator-prey size relationships in an African large-mammal food web

1. Size relationships are central in structuring trophic linkages within food webs, leading to suggestions that the dietary niche of smaller carnivores is nested within that of larger species. However, past analyses have not taken into account the differing selection shown by carnivores for specific size ranges of prey, nor the extent to which the greater carcass mass of larger prey outweighs the greater numerical representation of smaller prey species in the predator diet. Furthermore, the top-down impact that predation has on prey abundance cannot be assessed simply in terms of the number of predator species involved. 2. Records of found carcasses and cause of death assembled over 46 years in the Kruger National Park, South Africa, corrected for under-recording of smaller species, enabled a definitive assessment of size relationships between large mammalian carnivores and their ungulate prey. Five carnivore species were considered, including lion (Panthera leo), leopard (Panthera pardus), cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus) and spotted hyena (Crocuta crocuta), and 22 herbivore prey species larger than 10 kg in adult body mass. 3. These carnivores selectively favoured prey species approximately half to twice their mass, within a total prey size range from an order of magnitude below to an order of magnitude above the body mass of the predator. The three smallest carnivores, i.e. leopard, cheetah and wild dog, showed high similarity in prey species favoured. Despite overlap in prey size range, each carnivore showed a distinct dietary preference. 4. Almost all mortality was through the agency of a predator for ungulate species up to the size of a giraffe (800-1200 kg). Ungulates larger than twice the mass of the predator contributed substantially to the dietary intake of lions, despite the low proportional mortality inflicted by predation on these species. Only for megaherbivores substantially exceeding 1000 kg in adult body mass did predation become a negligible cause of mortality. 5. Hence, the relative size of predators and prey had a pervasive structuring influence on biomass fluxes within this large-mammal food web. Nevertheless, the large carnivore assemblage was dominated overwhelmingly by the largest predator, which contributed the major share of animals killed across a wide size range.     

Body size in the Eurasian lynx in Sweden: dependence on prey availability

The Eurasian lynx (Lynx lynx) is a common predator of both roe deer (Capreolus capreolus) and reindeer (Rangifer tarandus) in Sweden. We investigated the influence of prey availability, latitude, sex, and age on body size and body mass variation of the Eurasian lynx in Sweden, using data from 243 specimens whose locality of capture, year of capture, sex, and age were known. We found that both body size and body mass of the lynx in Sweden are mainly affected by the lynx sex and age but also by the availability of prey during the first year of life. Body size and body mass of lynx as well as the density of roe deer increased from Central Sweden to South. Furthermore, body size and body mass of lynx increased from Central Sweden to North (i.e. within the reindeer husbandry area). Lynx body size was slightly smaller within the reindeer husbandry area (approximately north of latitudes 62°–63°N) compared to outside, probably because reindeer are more difficult prey to hunt, as well as being migratory and thus an unpredictable prey for the Eurasian lynx compared to the non-migratory roe deer. Our results support a growing body of evidence showing that food availability at growth has a major effect on body size of animals.     

To kill a kangaroo: understanding the decision to pursue high-risk/high-gain resources

In this paper, we attempt to understand hunter–gatherer foraging decisions about prey that vary in both the mean and variance of energy return using an expected utility framework. We show that for skewed distributions of energetic returns, the standard linear variance discounting (LVD) model for risk-sensitive foraging can produce quite misleading results. In addition to creating difficulties for the LVD model, the skewed distributions characteristic of hunting returns create challenges for estimating probability distribution functions required for expected utility. We present a solution using a two-component finite mixture model for foraging returns. We then use detailed foraging returns data based on focal follows of individual hunters in Western Australia hunting for high-risk/high-gain (hill kangaroo) and relatively low-risk/low-gain (sand monitor) prey. Using probability densities for the two resources estimated from the mixture models, combined with theoretically sensible utility curves characterized by diminishing marginal utility for the highest returns, we find that the expected utility of the sand monitors greatly exceeds that of kangaroos despite the fact that the mean energy return for kangaroos is nearly twice as large as that for sand monitors. We conclude that the decision to hunt hill kangaroos does not arise simply as part of an energetic utility-maximization strategy and that additional social, political or symbolic benefits must accrue to hunters of this highly variable prey.     

Home range size variation in a recovering wolf population: evaluating the effect of environmental, demographic, and social factors

Home range size in mammals is a key ecological trait and an important parameter in conservation planning, and has been shown to be influenced by ecological, demographic and social factors in animal populations. Information on space requirements is especially important for carnivore species which range over very large areas and often come into direct conflict with human interest. We used long-term telemetry-location data from a recovering wolf population in Scandinavia to investigate variation in home range size in relation to environmental and social characteristics of the different packs. Wolves showed considerable variation in home range size, which ranged from 259 to 1,676 km². Although wolf density increased fourfold during the study period, we found no evidence that intraspecific competition influenced range size. Local variation in moose density, which was the main prey for most packs, did not influence wolf home range size. Home ranges increased with latitude and elevation and decreased with increased roe deer density. Although prey biomass alone did not influence range size, our data suggest that there is a correlation between habitat characteristics, choice of prey species and possible hunting success, which currently combine to shape home range size in Scandinavian wolves.     

Diet shift induced rapid evolution of size and function in a predatory bird

A predator’s body size correlates with its prey size. Change in the diet may call for changes in the hunting mode and traits determining hunting success. We explored long-term trends in sternum size and shape in the northern goshawk by applying geometric morphometrics. Tetraonids, the primary prey of the goshawk, have decreased and been replaced by smaller birds in the diet. We expected that the size of the goshawk has decreased accordingly more in males than females based on earlier observations of outer morphology. We also expected changes in sternum shape as a function of changes in hunting mode. Size of both sexes has decreased during the preceding decades (1962?2008), seemingly reflecting a shift in prey size and hunting mode. Female goshawks hunting also mammalian prey tend to have a pronouncedly “Buteo-type” sternum compared to males preying upon birds. Interestingly, the shrinkage of body size resulted in an increasingly “Buteo-type” sternum in both sexes. In addition, the sternum shape in birds that died accidentally (i.e., fit individuals) was more Buteo-type than in starved ones, hinting that selection was towards a Buteo-type sternum shape. We conclude that these observed patterns are likely due to directional selection driven by changes in the diet towards smaller and more agile prey. On the other hand, global warming is predicted to also cause a decrease in size, thus these two scenarios are inseparable. Because of difficulties in studying fitness-related phenotypic changes of large raptors in the field, time series of museum exemplars collected over a wide geographical area may give answers to this conundrum.