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.     

Dangerous prey and daring predators: a review

How foragers balance risks during foraging is a central focus of optimal foraging studies. While diverse theoretical and empirical work has revealed how foragers should and do manage food and safety from predators, little attention has been given to the risks posed by dangerous prey. This is a potentially important oversight because risk of injury can give rise to foraging costs similar to those arising from the risk of predation, and with similar consequences. Here, we synthesize the literature on how foragers manage risks associated with dangerous prey and adapt previous theory to make the first steps towards a framework for future studies. Though rarely documented, it appears that in some systems predators are frequently injured while hunting and risk of injury can be an important foraging cost. Fitness costs of foraging injuries, which can be fatal, likely vary widely but have rarely been studied and should be the subject of future research. Like other types of risk‐taking behaviour, it appears that there is individual variation in the willingness to take risks, which can be driven by social factors, experience and foraging abilities, or differences in body condition. Because of ongoing modifications to natural communities, including changes in prey availability and relative abundance as well as the introduction of potentially dangerous prey to numerous ecosystems, understanding the prevalence and consequences of hunting dangerous prey should be a priority for behavioural ecologists.     

Complex interactions between components of individual prey specialization affect mechanisms of niche variation in a grasshopper-hunting wasp

1.?Individual foraging behaviour defines the use of resources by a given population and its variation in different ways such as, for example, unpredictable interactions between taxon-biased and size-biased selection. Here we investigated how the environmental availability of prey and individual specialization, for both prey taxa and prey size, shape niche variation across generations in the grasshopper-hunting digger wasp Stizus continuus. 2.?The population of S.?continuus expressed selective predation, females mainly hunting species encountered on large bushes; diet changed across generations, due more to size increase in potential prey than to changes in the orthopteran community. 3.?Individual females of both generations weakly overlapped the size and taxa of prey, and the niche width of the second generation increased for both prey size and taxa. 4.?The greater variance in prey size in the environment accounted for the enlarged prey size niche of the second generation, but the load-lifting constraints of the wasps maintained individual prey size specialization constant. In contrast, the enlarged prey taxon niche paralleled a smaller overlap of diets between wasps in the second generation. 5.?Increased niche width in the S.?continuus population was thus achieved in two ways. Regarding prey size, all individuals shifted towards the use of the full set of available resources (parallel release). For prey taxa, according to the classical niche variation hypothesis, individuals diverged to minimize resource use overlap and perhaps intraspecific competition. These two mechanisms were observed for the first time simultaneously in a single predator population.     

Specialisation in prey capture drives coexistence among sympatric spider‐hunting wasps

1. Sister taxa that coexist in the same space and time often face competition due to the use of similar resources. However, some closely related species can adopt fine‐grained specialisation in resource use to coexist. This study investigated niche overlap between three sympatric spider‐hunting wasp species of the genus Trypoxylon (Hymenoptera: Crabronidae) known to nest in three of the habitats found in the study area. 2. First, the co‐occurrence of these wasp species in the three habitats was estimated, as a proxy for potential competition. Then, the following hypotheses were tested: (i) niche partitioning is seen more often between species that co‐occur in a habitat, whereas there is niche overlap between species nesting in distinct habitats (prey specialisation hypothesis); and (ii) wasp species capture prey according to their size (physical constraint hypothesis). 3. Two pairs of wasp species were found consistently nesting in the same habitat. Niche partitioning based on prey taxa occurred regardless of the habitat preference. It was also found that differences in the size of wasps reflected distinctions in the size of their prey. 4. These findings were consistent over the years, showing that the significance of specialisation in foraging activities and physical constraints during prey capture can play key roles in the coexistence of sympatric species. The distinctions in the foraging strategies of these wasps are discussed, as well as potential mechanisms driving the evolution in prey specialisation, with insights for future studies.     

Benefits of Group Foraging Depend on Prey Type in a Small Marine Predator,the Little Penguin

Group foraging provides predators with advantages in over-powering prey larger than themselves or in aggregating small prey for efficient exploitation. For group-living predatory species, cooperative hunting strategies provide inclusive fitness benefits. However, for colonial-breeding predators, the benefit pay-offs of group foraging are less clear due to the potential for intra-specific competition. We used animal-borne cameras to determine the prey types, hunting strategies, and success of little penguins (Eudyptula minor), a small, colonial breeding air-breathing marine predator that has recently been shown to display extensive at-sea foraging associations with conspecifics. Regardless of prey type, little penguins had a higher probability of associating with conspecifics when hunting prey that were aggregated than when prey were solitary. In addition, success was greater when individuals hunted schooling rather than solitary prey. Surprisingly, however, success on schooling prey was similar or greater when individuals hunted on their own than when with conspecifics. These findings suggest individuals may be trading-off the energetic gains of solitary hunting for an increased probability of detecting prey within a spatially and temporally variable prey field by associating with conspecifics.     

Intragroup competition predicts individual foraging specialisation in a group‐living mammal

Individual foraging specialisation has important ecological implications, but its causes in group‐living species are unclear. One of the major consequences of group living is increased intragroup competition for resources. Foraging theory predicts that with increased competition, individuals should add new prey items to their diet, widening their foraging niche (‘optimal foraging hypothesis’). However, classic competition theory suggests the opposite: that increased competition leads to niche partitioning and greater individual foraging specialisation (‘niche partitioning hypothesis’). We tested these opposing predictions in wild, group‐living banded mongooses (Mungos mungo), using stable isotope analysis of banded mongoose whiskers to quantify individual and group foraging niche. Individual foraging niche size declined with increasing group size, despite all groups having a similar overall niche size. Our findings support the prediction that competition promotes niche partitioning within social groups and suggest that individual foraging specialisation may play an important role in the formation of stable social groupings.     

Social learning of hunting skills in juvenile marsh harriers <Emphasis Type="Italic">Circus aeruginosus</Emphasis>

The impact of social factors on the improvement of hunting skills of juvenile marsh harriers during their first autumnal migration were studied in SE Poland. While foraging with adult birds, juveniles performed more dives on prey both in terms of number of trials and rates. Hunting sessions of juveniles were more efficient in the presence of adults than in the absence of adults. Juveniles hunting with adults and other juveniles could select adequate habitat patches in which access to prey is easier. The role of vertical and horizontal transmission of information in the development of hunting skills in juvenile marsh harrier were confirmed because faster development of hunting ability was achieved in the social hunting after the end of their postfledging dependency period.     

The ecological conditions that favor tool use and innovation in wild bottlenose dolphins (Tursiops sp.)

Dolphins are well known for their exquisite echolocation abilities, which enable them to detect and discriminate prey species and even locate buried prey. While these skills are widely used during foraging, some dolphins use tools to locate and extract prey. In the only known case of tool use in free-ranging cetaceans, a subset of bottlenose dolphins (Tursiops sp.) in Shark Bay, Western Australia habitually employs marine basket sponge tools to locate and ferret prey from the seafloor. While it is clear that sponges protect dolphins' rostra while searching for prey, it is still not known why dolphins probe the substrate at all instead of merely echolocating for buried prey as documented at other sites. By 'sponge foraging' ourselves, we show that these dolphins target prey that both lack swimbladders and burrow in a rubble-littered substrate. Delphinid echolocation and vision are critical for hunting but less effective on such prey. Consequently, if dolphins are to access this burrowing, swimbladderless prey, they must probe the seafloor and in turn benefit from using protective sponges. We suggest that these tools have allowed sponge foraging dolphins to exploit an empty niche inaccessible to their non-tool-using counterparts. Our study identifies the underlying ecological basis of dolphin tool use and strengthens our understanding of the conditions that favor tool use and innovation in the wild.     

Seasonal Trophic Niche Shift and Cascading Effect of a Generalist Predator Fish

Few studies have examined how foraging niche shift of a predator over time cascade down to local prey communities. Here we examine patterns of temporal foraging niche shifts of a generalist predator (yellow catfish, Pelteobagrus fulvidraco) and the abundance of prey communities in a subtropical lake. We predicted that the nature of these interactions would have implications for patterns in diet shifts and growth of the predator. Our results show significant decreases in planktivory and benthivory from late spring to summer and autumn, whereas piscivory increased significantly from mid-summer until late autumn and also increased steadily with predator body length. The temporal dynamics in predator/prey ratios indicate that the predation pressure on zooplankton and zoobenthos decreased when the predation pressure on the prey fish and shrimps was high. Yellow catfish adjusted their foraging strategies to temporal changes in food availability, which is in agreement with optimal foraging theory. Meanwhile the decrease in planktivory and benthivory of yellow catfish enabled primary consumers, such as zooplankton and benthic invertebrates, to develop under low grazing pressure via trophic cascading effects in the local food web. Thus, yellow catfish shifts its foraging niche to intermediate consumers in the food web to benefit the energetic demand on growth and reproduction during summer, which in turn indirectly facilitate the primary consumers. In complex food webs, trophic interactions are usually expected to reduce the strength and penetrance of trophic cascades. However, our study demonstrates strong associations between foraging niche of piscivorous fish and abundance of prey. This relationship appeared to be an important factor in producing top-down effects on both benthic and planktonic food webs.     

Dichromatism in relation to the trophic biology of predatory cichlid fishes in Lake Tanganyika, East Africa

A population of the piscivorous Tanganyikan cichlid, Lepidiolamprologus profundicola contains individually distinct pale and dark colour morphs: a dichromatism not related to age or sex. The 'dark form' frequently targets prey from the shaded space under rocks, while the 'plae form' targets prey in well-iluminated open water. This hunting site specialization is not ascribable to the differences of microhabitat that the predators of each form encounter. The main function of the dichromatism is apparently cryptic, as a camouflage for hunting. Interspecific comparisons among 19 species of carnivorous Tanganyikan cichlid fishes reveal that dichromatic taxa generally chase an active prey: fishes, scales of fishes or highly mobile free-swimming crustacea. In contrast, cichlid species foraging on a sessile or sluggish prey are monochromatic. This finding strongly supports the hypothesis that a pale-dark dichromatism serves to optimize foraging efficiency in predatory cichlids hunting an active prey.     

Foraging Trade‐offs between Prey Size,Delivery Rate and Prey Type: How Does Niche Breadth and Early Learning of the Foraging Niche Affect Food Delivery?

Optimal foraging theory suggests that avian parents should prefer the most energetically efficient (largest) prey items when delivering food to offspring at a central place. However, during periods of high demand, selectivity of prey may decline, leading to the delivery of smaller and/or less nutritious items. We compared foraging trade‐offs between great tits (Parus major) which had a wider feeding niche than blue tits (Cyanistes caeruleus). We also compared the foraging efficiency of cross‐fostered young, which had learned the spatial foraging niche and prey size of the foreign species, to that of control conspecifics. Mean delivery rates did not differ between control and cross‐fostered parents of either species but as delivery rates increased, prey size declined for both species and both treatment groups. However, across the range of increasing delivery rates, parents were able to increase the total biomass of prey delivered. Cross‐fostering did not alter the proportion of different prey taxa in the diet, but cross‐fostered birds shifted the size of the prey taken to that of their foster species. Consistent with their broader feeding niche, great tits, but not blue tits, incorporated more unpalatable items (flies) as delivery rates increased. Although great tits foraged less efficiently in the blue tit niche, paradoxically, blue tits seem to deliver more prey biomass when foraging in the great tit niche.     

Polychromatism of the scale-eaterPerissodus microlepis (Cichlidae,Teleostei) in relation to foraging behavior

Variation in foraging behavior in a population of the scale-eaterPerissodus microlepis was studied on the northwest coast of Lake Tanganyika. Differences in body coloration were found among adult and subadult individuals, which were classified into 4 color morphs designated as Beige, Dark, Grey and Stripe. These color morphs were not strictly related to either sex or size. Each morph spent much time in specific microhabitats and had a major hunting technique that differed from other morphs. Beige morph. which predominated in number, ambushed prey at open surfaces of the substrate, whereas Dark morph used the shade of rock as an ambush site. Grey morph mixed in schools of fisheds hovering in midwater to attack school members, and Stripe morph cruised in the water column and stooped mainly at bottom-fishes. Prey preference differed among the morphs corresponding to their hunting techniques but successful attack rates were similar among them. Observations of marked individuals demonstrated adherence to particular hunting techniques and, in some cases, to particular hunting sites. Intraspecific foraging specialization is discussed in relation to the function of body color and diversity of life styles of prey fishes.     

Dietary niche breadth in a local community of passerine birds: an analysis using phylogenetic contrasts

The analysis of a local community of forest passerines (13 species) using phylogenetic contrasts shows a correlation between body size of bird species and mean prey size, minimum prey size, maximum prey size and the size range of dietary items. This suggests that larger birds drop small prey taxa from their prey list, because of the difficulty of capturing very small prey, for energetic reasons or because of microhabitat usage. We find some support for the third hypothesis. Dietary niche breadth calculated across prey taxa is not related to body size. Dietary niche breadth, however, is correlated with size-corrected measurements of the bill and locomotor apparatus. Long and slender bills increase the dietary niche breadth. Thus subtle differences constrain foraging and the techniques of extracting certain prey taxa form crevices. Dietary niche breadth and foraging diversity are positively correlated with population density: at least locally dietary generalists occur at higher breeding densities than specialists.     

Evolution of reversed sexual size dimorphism and role partitioning among predatory birds, with a size scaling of flight performance

To explain the adaptive significance of sex role partitioning and reversed sexual size dimorphism among raptors, owls and skuas, where females are usually larger than males, we combine several previous hypotheses with some new ideas. Owing to their structural and behavioural adaptations for prey capture, predatory birds have better prospects than other birds of defending their offspring against nest predators. This makes sex role partitioning advantageous; one parent guards the offspring while the other forages for the family. Further, among predators hunting alert prey such as vertebrates, two mates because of interference may not procur much more food than would one mate hunting alone. By contrast, two mates feeding on less alert prey may together obtain almost twice as much food as one mate hunting alone. For these reasons, partitioning of breeding labours might be adaptive only in predatory birds. An initial imbalance favours female nest guarding and male foraging: the developing eggs might be damaged if the female attacks prey; their mass might reduce her flight performance; she must visit the nest to lay; and the male feeds her before she lays (‘courtship feeding’). Increased female body size should enhance egg production, incubation, ability to tear apart prey for the young, and, in particular, offspring protection in predatory birds. Efficient foraging during the breeding period then becomes most important for the male. This imposes great demands on aerial agility in males, particularly among predators of agile prey. Flight performance decreases with increasing size in five of six aspects explored. The male must therefore not be too large in relation to the most important prey. For these reasons, he should be smaller than the female. Among predatory birds, size dimorphism increases with the proportion of birds in the diet, which may be explained as follows. Adult birds have mainly one type of predators: other predatory birds. Because almost only these specialists exploit adult birds, they carry out most of the cropping of this prey. A predator of easier prey competes with many other kinds of predators, which considerably reduce prey abundance in its territory. This is not so for predators of adult birds. Further, because birds are extremely agile, the specialized predator can hunt efficiently only within a limited size range of birds, whose flight skill it can match. Increased size dimorphism among these predators therefore should be particularly important for enlarging the combined food base of the pair. A bird specialist may consume much of the available prey in the suitable size range during the breeding period. When the predator's young are large enough to defend themselves, the female aids better by hunting than by guarding the chicks. It is advantageous among bird specialists if she hunts prey of other sizes than does the male, who has by then reduced prey abundance in his prey size class. But among predatory birds hunting easier prey the female gains little by hunting outside the male's prey spectrum, because other kinds of predators will have reduced the prey abundance outside as well as inside the male's preferred size range. Intra-pair food separation through large sexual size dimorphism therefore should be particularly advantageous among predators of birds. This may be the main reason why the degree of size dimorphism increases with the dietary proportion of birds.     

Assumption validity in human optimal foraging: The Barí hunters of Venezuela as a test case

Optimal foraging theory is being used increasingly as a means of understanding human foraging behavior. One of the central assumptions of optimal foraging theory is that prey items or patches are encountered sequentially and as a Poisson process. Using empirical data gathered on the Barí hunters of Venezuela, we assess the validity of this central assumption. We compare our observed distribution of encounter frequencies with an expected Poisson distribution, utilizing chisquare tests and graphic representations. The results are strikingly consonant with the expected Poisson distribution and lend support to the applicability of optimal foraging models to human hunting behavior.     

Influence of Group Size on the Success of Wolves Hunting Bison

An intriguing aspect of social foraging behaviour is that large groups are often no better at capturing prey than are small groups, a pattern that has been attributed to diminished cooperation (i.e., free riding) in large groups. Although this suggests the formation of large groups is unrelated to prey capture, little is known about cooperation in large groups that hunt hard-to-catch prey. Here, we used direct observations of Yellowstone wolves (Canis lupus) hunting their most formidable prey, bison (Bison bison), to test the hypothesis that large groups are more cooperative when hunting difficult prey. We quantified the relationship between capture success and wolf group size, and compared it to previously reported results for Yellowstone wolves hunting elk (Cervus elaphus), a prey that was, on average, 3 times easier to capture than bison. Whereas improvement in elk capture success levelled off at 2–6 wolves, bison capture success levelled off at 9–13 wolves with evidence that it continued to increase beyond 13 wolves. These results are consistent with the hypothesis that hunters in large groups are more cooperative when hunting more formidable prey. Improved ability to capture formidable prey could therefore promote the formation and maintenance of large predator groups, particularly among predators that specialize on such prey.     

Foraging mode affects the evolution of egg size in generalist predators embedded in complex food webs

Ecological networks incorporate myriad biotic interactions that determine the selection pressures experienced by the embedded populations. We argue that within food webs, the negative scaling of abundance with body mass and foraging theory predict that the selective advantages of larger egg size should be smaller for sit‐and‐wait than active‐hunting generalist predators, leading to the evolution of a difference in egg size between them. Because body mass usually scales negatively with predator abundance and constrains predation rate, slightly increasing egg mass should simultaneously allow offspring to feed on more prey and escape from more predators. However, the benefits of larger offspring would be relatively smaller for sit‐and‐wait predators because (i) due to their lower mobility, encounters with other predators are less common, and (ii) they usually employ a set of alternative hunting strategies that help to subdue relatively larger prey. On the other hand, for active predators, which need to confront prey as they find them, body‐size differences may be more important in subduing prey. This difference in benefits should lead to the evolution of larger egg sizes in active‐hunting relative to sit‐and‐wait predators. This prediction was confirmed by a phylogenetically controlled analysis of 268 spider species, supporting the view that the structure of ecological networks may serve to predict relevant selective pressures acting on key life history traits.     

Experimental study of cognitive aspects of ambivalent foraging as exemplified by the great tit

A hypothesis of ambivalent foraging is proposed based on ideas about dual treating of the prey by a consumer: the food value attracts while the danger repulses. The foraging strategy of the great tit was investigated experimentally with the use of artificial “food patches” with variable amounts of dangerous prey (live red wood ants) and non-dangerous prey (fly larvae). With non-dangerous prey, the behavior of the birds corresponded to the known marginal value theorem: they proceeded with foraging until the resources were exhausted. We found the threshold amount of dangerous prey that prevents tits from hunting.     

Snow tracking reveals different foraging patterns of red foxes and pine martens

Red fox (Vulpes vulpes) shares similar prey preferences and co-occurs with several other carnivores, and is together with pine marten (Martes martes), the most common mesocarnivore in the northern boreal forest. Voles are important prey for both species, but it is unclear to what extent they compete for the same food resources in winter. Here, we use 2139 km and 533 km of meticulous snow tracking of red foxes and pine martens to evaluate their food niches. We measured hunting and digging behaviour, whether successful or not, and the effect of snow depth and temperature. Pine martens were restricted to forested habitats, whereas red foxes used a wide range of habitats. Red foxes were found to dig more often than pine martens, 0.67 vs. 0.39 digging events per kilometre. Hunting was less common and similar in both species, about 0.1 hunting event per kilometre. Pine martens were more efficient in hunting and finding food remains compared to red foxes. Increasing snow depth reduced hunting success and also reduced dig success of red foxes. Food niche overlap was small. Red foxes used mostly voles and carrion remains of ungulates, whereas pine martens used cached eggs and small birds. We suggest that caching eggs is an important strategy for pine martens to survive winter in northern latitudes. Snow depth was important for capturing voles, and thick snow cover appeared to mask the effect of vole peaks. Intensified land use, as clear-cutting and leaving slaughter remains from harvest, will benefit red foxes on the expense of pine martens. The ongoing climate change with warmer winters and less snow will likely further benefit the red fox.