Short‐term prey field lability constrains individual specialisation in resource selection and foraging site fidelity in a marine predator

Spatio‐temporally stable prey distributions coupled with individual foraging site fidelity are predicted to favour individual resource specialisation. Conversely, predators coping with dynamic prey distributions should diversify their individual diet and/or shift foraging areas to increase net intake. We studied individual specialisation in Scopoli's shearwaters (Calonectris diomedea) from the highly dynamic Western Mediterranean, using daily prey distributions together with resource selection, site fidelity and trophic‐level analyses. As hypothesised, we found dietary diversification, low foraging site fidelity and almost no individual specialisation in resource selection. Crucially, shearwaters switched daily foraging tactics, selecting areas with contrasting prey of varying trophic levels. Overall, information use and plastic resource selection of individuals with reduced short‐term foraging site fidelity allow predators to overcome prey field lability. Our study is an essential step towards a better understanding of individual responses to enhanced environmental stochasticity driven by global changes, and of pathways favouring population persistence.     

Within-colony feeding selectivity by a corallivorous reef fish: foraging to maximize reward?

Foraging theory predicts that individuals should choose a prey that maximizes energy rewards relative to the energy expended to access, capture, and consume the prey. However, the relative roles of differences in the nutritive value of foods and costs associated with differences in prey accessibility are not always clear. Coral‐feeding fishes are known to be highly selective feeders on particular coral genera or species and even different parts of individual coral colonies. The absence of strong correlations between the nutritional value of corals and prey preferences suggests other factors such as polyp accessibility may be important. Here, we investigated within‐colony feeding selectivity by the corallivorous filefish, Oxymonacanthus longirostris, and if prey accessibility determines foraging patterns. After confirming that this fish primarily feeds on coral polyps, we examined whether fish show a preference for different parts of a common branching coral, Acropora nobilis, both in the field and in the laboratory experiments with simulated corals. We then experimentally tested whether nonuniform patterns of feeding on preferred coral species reflect structural differences between polyps. We found that O. longirostris exhibits nonuniform patterns of foraging in the field, selectively feeding midway along branches. On simulated corals, fish replicated this pattern when food accessibility was equal along the branch. However, when food access varied, fish consistently modified their foraging behavior, preferring to feed where food was most accessible. When foraging patterns were compared with coral morphology, fish preferred larger polyps and less skeletal protection. Our results highlight that patterns of interspecific and intraspecific selectivity can reflect coral morphology, with fish preferring corals or parts of coral colonies with structural characteristics that increase prey accessibility.     

Invasive house mice facing a changing environment on the Sub‐Antarctic Guillou Island (Kerguelen Archipelago)

Adaptation to new environments is a key feature in evolution promoting divergence in morphological structures under selection. The house mouse (Mus musculus domesticus) introduced on the Sub‐Antarctic Guillou Island (Kerguelen Archipelago) had and still has to face environmental conditions that likely shaped the pattern and pace of its insular evolution. Since mouse arrival on the island, probably not more than two centuries ago, ecological conditions dramatically differed from those available to their Western European commensal source populations. In addition, over the last two decades, the plant and animal communities of Guillou Island were considerably modified by the eradication of rabbits, the effects of climate change and the spread of invasive species detrimental to native communities. Under such a changing habitat, the mouse response was investigated using a morphometric quantification of mandible and molar tooth, two morphological structures related to food processing. A marked differentiation of the insular mice compared with their relatives from Western Europe was documented for both mandibles and molar shapes. Moreover, these shapes changed through the 16 years of the record, in agreement with expectations of drift for the molar, but more than expected by chance for the mandible. These results suggest that mice responded to the recent changes in food resources, possibly with a part of plastic variation for the mandible prone to bone remodelling. This pattern exemplifies the intricate interplay of evolution, ecology and plasticity that is a probable key of the success of such an invasive rodent facing pronounced shifts in food resources exploitation under a changing environment.     

Divergent in shape and convergent in function: Adaptive evolution of the mandible in Sub‐Antarctic mice

Convergent evolution in similar environments constitutes strong evidence of adaptive evolution. Transported with people around the world, house mice colonized even remote areas, such as Sub‐Antarctic islands. There, they returned to a feral way of life, shifting towards a diet enriched in terrestrial macroinvertebrates. Here, we test the hypothesis that this triggered convergent evolution of the mandible, a morphological character involved in food consumption. Mandible shape from four Sub‐Antarctic islands was compared to phylogeny, tracing the history of colonization, and climatic conditions. Mandible shape was primarily influenced by phylogenetic history, thus discarding the hypothesis of convergent evolution. The biomechanical properties of the jaw were then investigated. Incisor in‐lever and temporalis out‐lever suggested an increase in the velocity of incisor biting, in agreement with observations on various carnivorous and insectivorous rodents. The mechanical advantage related to incisor biting also revealed an increased functional performance in Sub‐Antarctic populations, and appears to be an adaptation to catch prey more efficiently. The amount of change involved was larger than expected for a plastic response, suggesting microevolutionary processes were evolved. This study thus denotes some degree of adaptive convergent evolution related to changes in habitat‐related changes in dietary items in Sub‐Antarctic mice, but only regarding simple, functionally relevant aspects of mandible morphology.     

Lack of diet segregation during breeding by male and female Northern Flickers foraging on ants

Sexual size dimorphism can result in reduced competition if it leads males and females to use different foraging techniques or consume different prey items. Among woodpeckers, differences between males and females in bill length are common and may explain foraging differences in this family of birds. Northern Flickers (Colaptes auratus) are ground‐foraging woodpeckers that specialize on ants. However, the overall contribution of ants to their diet and the proportions of particular ant genera in their diet are not well known. To understand the relationship between bill morphology and the consumption of prey items, we compared the bill length and bill width of male and female flickers. We then collected and analyzed fecal samples from breeding flickers (N = 40 males, 33 females) at a study site in central British Columbia, Canada. Bills of male flickers were significantly longer (4%) and wider (5%) than those of females. Of 11 prey types identified, ants made up over 99% of their diet, and the abundance and composition of ant taxa in the diet did not differ between the sexes. We found significant year and time of season effects, with the abundance of Tapinoma sessile and Lasius spp. increasing from May to the end of June and differing between years. This difference in diet composition between years may have been due to changes in the abundance or accessibility of certain ant taxa related to differences in vegetation structure or weather. Nine ant taxa were consumed by flickers and the four most common were T. sessile, Lasius spp.,Myrmica spp., and the Formica fusca species group. The degree of dimorphism in bill size of male and female Northern Flickers in our study was smaller than reported for several species of arboreal‐foraging woodpeckers, suggesting that bill size of ground‐foraging woodpeckers may not be strongly linked to niche separation at the level of prey selection.     

High resource overlap and small dietary differences are widespread in food-limited warbler (Parulidae) communities

Although both interspecific competition and coexistence mechanisms are central to ecological and evolutionary theory, past empirical studies have generally focused on simple (two-species) communities over short time periods. Experimental tests of these species interactions are challenging in complex study systems. Moreover, several studies of ‘imperfect generalists’, consistent with Liem's Paradox, raise questions about the ability of evolved species differences to partition niche space effectively when resources vary considerably across the annual cycle. Here we used a recently developed theoretical framework to combine past research on population-level processes with observational data on resource use to test for ongoing interspecific competition and understand the nature of resource overlap. We compared species diet overlaps and differences in several distinctive communities centred on a focal species, the American Redstart Setophaga ruticilla replicated both spatially and seasonally, in combination with documentation of population regulation to assess the ability of similar species to partition dietary niche space and limit interspecific competition. Our results document high dietary overlap in most of the communities studied, with only subtle differentiation consistent with known species differences in foraging behaviour and morphology. These findings are largely consistent with species foraging as imperfect generalists. However, in contrast to past studies, the high diet overlaps observed here during times of inferred resource scarcity were driven by low-value prey taxa (e.g. small ants) and did not involve truly ‘private’ resources. All of these factors increase the potential negative impacts of interspecific competition, and limit the ability of these birds to avoid competition if food availability deteriorates further than observed in our study, either seasonally or at longer intervals.     

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.     

Selectivity underlies the dissociation between seasonal prey availability and prey consumption in a generalist predator

Generalist predators are capable of selective foraging, but are predicted to feed in close proportion to prey availability to maximize energetic intake especially when overall prey availability is low. By extension, they are also expected to feed in a more frequency‐dependent manner during winter compared to the more favourable foraging conditions during spring, summer and fall seasons. For 18 months, we observed the foraging patterns of forest‐dwelling wolf spiders from the genus Schizocosa (Araneae: Lycosidae) using PCR‐based gut‐content analysis and simultaneously monitored the activity densities of two common prey: springtails (Collembola) and flies (Diptera). Rates of prey detection within spider guts relative to rates of prey collected in traps were estimated using Roualdes’ c_st model and compared using various linear contrasts to make inferences pertaining to seasonal prey selectivity. Results indicated spiders foraged selectively over the course of the study, contrary to predictions derived from optimal foraging theory. Even during winter, with overall low prey densities, the relative rates of predation compared to available prey differed significantly over time and by prey group. Moreover, these spiders appeared to diversify their diets; the least abundant prey group was consistently overrepresented in the diet within a given season. We suggest that foraging in generalist predators is not necessarily restricted to frequency dependency during winter. In fact, foraging motives other than energy maximization, such as a more nutrient‐focused strategy, may also be optimal for generalist predators during prey‐scarce winters.     

Spatial and temporal variation in the diet of the gentoo penguin (<Emphasis Type="Italic">Pygoscelis papua</Emphasis>) at Kerguelen Islands

Benthic divers are dependent on local resources and may therefore adopt different foraging strategies to cope with their energetic requirements in varying situations. We investigated the diet of gentoo penguins (Pygoscelis papua) at Kerguelen Islands, comparing its spatial and temporal variations with the general prey distribution. The study was conducted at four sites over 2 years. In total, 212 stomach contents were collected over the entire breeding season. The diet was composed mainly of neritic fish and crustaceans, with important spatial and seasonal variations. Fish dominated the diet at localities facing the open sea (from 38.0% to 94.6% by mass), whereas crustaceans dominated at the more protected site (84.3% by mass). Fish were more abundant in the winter diet and Euphausia vallentini, the major crustacean species, was more abundant in the summer diet. No inter-year variations were detected. These results are consistent with local prey availability, and highlight the large plasticity of the gentoo penguin diet and foraging behaviour.     

Feeding on the edge: foraging White Ibis target inter‐habitat prey fluxes

Avian population dynamics are influenced by the availability of spatiotemporally variable prey resources, but the conditions producing abundant and accessible prey are not always clear. In the Florida Everglades, wading birds nest in the dry season when receding water levels concentrate prey and facilitate improved foraging efficiency. White Ibis (Eudocimus albus) feed extensively on crayfish in sloughs, and previous studies have demonstrated that crayfish move downgradient from higher elevation, heavily vegetated ridge habitats into adjacent less‐vegetated sloughs when ridges are almost dry. Most White Ibis foraging is thought to occur in sloughs with relatively shallow water (< 19 cm), but crayfish move and their densities peak when water in sloughs is deeper (~ 21–32 cm). We conducted an observational study of White Ibis foraging in drying wetlands to determine if White Ibis restricted their foraging to shallow water or if they foraged in relatively deep water when crayfish were migrating. In a series of large drying wetlands, we used time‐lapse imagery to quantify White Ibis foraging activity over 61 d from February to April 2017 and we also quantified crayfish biomass density in sloughs. Crayfish biomass density peaked when ridges were almost dry. Most White Ibis foraging occurred over 2–3 d when ridges were almost dry and water in sloughs averaged ≥ 29 cm deep. White Ibis selected slough edges for foraging, suggesting that they were capturing crayfish migrating between habitats. Our results point to a new mechanism of prey exploitation driven by inter‐habitat prey flux when ridge habitat dries. Although the results of previous studies suggest that White Ibis will not forage on fish in deeper water (> 25 cm), we found that White Ibis will forage on crayfish in water at those depths. Maintenance of habitat elevational differences and hydro‐patterns that promote crayfish production will be necessary to promote this predator–prey interaction in the ridge‐slough landscape of the Everglades.     

Trophic niche partitioning and diet composition of sympatric fin (Balaenoptera physalus) and humpback whales (Megaptera novaeangliae) in the Gulf of Alaska revealed through stable isotope analysis

Fin and humpback whales are large consumers that are often sympatric, effectively sharing or partitioning their use of habitat and prey resources. Stable carbon and nitrogen isotopes in the skin of fin and humpback whales from two regions in the western Gulf of Alaska, Kodiak, and Shumagin Islands, were analyzed to test the hypothesis that these sympatric baleen whales exhibit trophic niche partitioning within these regions. Standard ellipse areas, estimated using Bayesian inference, suggested that niche partitioning between species is occurring in the Kodiak region but not in the Shumagin Islands. Isotopic mixing models based on stable isotopes from whales and local prey samples, were used to estimate possible diet solutions for whales in the Kodiak region. Comparison of isotopic niches and diet models support niche partitioning, with fin whales foraging primarily on zooplankton and humpback whales foraging on zooplankton and small forage fish. The results of this study show that niche partitioning between sympatric species can vary by region and may be the result of prey availability, prey preferences, or both.     

Using dive behavior and active acoustics to assess prey use and partitioning by fin and humpback whales near Kodiak Island,Alaska

Near the Kodiak Archipelago, fin (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales frequently overlap spatially and temporally. The Gulf Apex Predator‐prey study (GAP) investigated the prey use and potential prey partitioning between these sympatric species by combining concurrent analysis of vertical whale distribution with acoustic assessment of pelagic prey. Acoustic backscatter was classified as consistent with either fish or zooplankton. Whale dive depths were determined through suction cup tags. Tagged humpback whales (n = 10) were most often associated with distribution of fish, except when zooplankton density was very high. Associations between the dive depths of tagged fin whales (n = 4) and the vertical distribution of either prey type were less conclusive. However, prey assessment methods did not adequately describe the distribution of copepods, a potentially significant resource for fin whales. Mean dive parameters showed no significant difference between species when compared across all surveys. However, fin whales spent a greater proportion of dive time in the foraging phase than humpbacks, suggesting a possible difference in foraging efficiency between the two. These results suggest that humpback and fin whales may target different prey, with the greatest potential for diet overlap occurring when the density of zooplankton is very high.     

The influence of prey size,sediment thickness and fish size on consumption in common sole (Solea solea L.)

This study determined prey consumption in common sole as a function of prey size (0–0.5, 1–1.5, 2–2.5 and 4–5 g), sediment thickness (20 cm and 2 cm) and fish size (50 g, 125 g or 300 g). Prey consumption (in numbers of prey eaten per fish per day) was reduced with increasing prey size and sediment thickness, and was increased with increasing fish size (p < .001 for all factors). All 3 factors showed significant two way interactions (p < .001) when expressed in numbers of prey eaten. Prey consumption decreased with prey size when prey could not escape by burying (2 cm of sediment thickness) irrespective of fish size. We suggest that increasing effort to ingest and handle larger prey played a role. Prey consumption increased with fish size when prey could not bury (2 cm of sediment thickness). However, when prey was able to bury (at 20 cm sediment thickness) prey consumption was similar irrespective of fish size (p < .001 for interaction fish size × sediment). This interaction suggests that with increasing fish size there is an increasing mismatch between foraging adaptation and prey burial depth. This may explain the dominance of crustaceans in the diet of adult common sole in nature, despite the high abundance of polychaetes.     

Insect prey foraging strategies in Callicebus oenanthe in northern Peru

Titi monkeys (genus Callicebus) are small-bodied platyrrhines that supplement their predominantly frugivorous diet with variable amounts of leaves, seeds, and/or arthropod prey. Notable interspecific variation in the amount of insect prey in the diet has been observed in Callicebus, ranging from 0% to 20%. In this study, I investigate the degree and type of prey foraging in a little-known species, Callicebus oenanthe inhabiting a fragmented, secondary forest on the foothills of the Andes in northern Peru. I present data on prey type, prey search and capture techniques, substrate/vegetation use, foraging height, prey capture efficiency, and seasonal variation of insect prey foraging in one group of C. oenanthe observed from January to August 2005. Insect prey accounted for 22% of the diet, the highest amount reported for any Callicebus species to date, and insects from at least six different orders were included. C. oenanthe was mainly an investigative forager of hidden prey, manipulating easy-to-open substrates such as rolled up leaves, and hunted ant swarms and larger insects opportunistically. Insect foraging was predominant during the dry season (26%) and decreased during the wet season (13%). The study group foraged mostly in the understory (2-6 m) within vine-laden shrubs and trees, which may conform to an anti-predator strategy of crypticity. Overall the group had an 83% insect capture success rate. These data suggest that insect prey is an important part of the diet of C. oenanthe and may be especially notable during periods of resource scarcity. This study adds to the knowledge concerning insect prey foraging in Callicebus, which can have an important role in defining ecological strategies in the selection of secondary protein food resources within a given ecosystem.     

The relationship between skull morphology, biting performance and foraging mode in Kalahari lacertid lizards

Lizards are a diverse clade in which one radiation consists entirely of sit-and-wait foragers and another consists of wide foragers. Lizards utilizing these two foraging modes are known to differ in diet, but little is known about how feeding morphology relates to diet and/or foraging mode. This study tested the hypothesis that skull morphology and biting performance are related to diet preference, and consequently, coevolve with foraging mode. Four species of lacertid lizard were studied because they vary in foraging mode, their phylogenetic relationships are known and they are well studied ecologically. Using an 'ecomorphological' approach, skull morphology and biting performance were quantified and mapped on to the phylogeny for the species. The results indicate that sit-and-wait species have shorter, wider skulls than the wide foraging species, and that all are significantly different in overall head shape. The sit-and-wait species had similar values for biting performance; however, clear phylogenetic patterns of covariation were not present between sit-and-wait and wide foraging species for either biting performance or skull morphology. Thus, skull morphology and performance have little influence on diet and foraging mode in these species. Instead it is likely that other factors such as seasonal prey availability and/or life history strategy shape foraging mode decisions.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 403–416.     

Prey choice by marten during a decline in prey abundance

Summary We examined variation in diet choice by marten (Martes americana) among seasons and between sexes and ages from 1980–1985. During this period prey populations crashed simultaneously, except for ruffed grouse (Bonasa umbellus) which was common at the beginning and end of the study, and masked shrews (Sorex cinereus) which were abundant in 1983. Marten were catholic in selection of prey and made use of most available mammalian prey, ruffed grouse, passerine birds, berries, and insects. Diet niche was widest during the latter three years when prey was scare, particularly in late winter. Diet niche breadth was negatively correlated with abundance of all common prey species. Proportion of small prey species in the diet was correlated with absolute abundance of those species, but proportion of some large prey was related to their relative abundance. Diet choice varied among years and among seasons. Berries and insects were common in summer diets while large prey, particularly varying hare (Lepus americanus), were more frequent in winter diet than in summer diet. We found little evidence that any small mammal species was a preferred prey. Sexual size dimorphism between the sexes did not affect prey choice, nor did age. Reduced foraging effort in winter resulted in a wider diet niche only when prey was scarce. The only prediction of optimal foraging models fully supported by our data was a wider diet niche with reduced prey abundance. However, among the three most profitable prey species choice was dependent on the absolute abundance of the most profitable type (varying hare). We suggest that marten primarily forage for large prey but employ a strategy which results in encounters with small prey as well. These small prey are eaten as they provide energy at minimal cost, between captures of large prey.     

Interpopulation resource partitioning of Lesser Frigatebirds and the influence of environmental context

Conspecific individuals inhabiting nearby breeding colonies are expected to compete strongly for food resources owing to the constraints imposed by shared morphology, physiology, and behavior on foraging strategy. Consequently, colony‐specific foraging patterns that effectively partition the available resources may be displayed. This study aimed to determine whether intraspecific resource partitioning occurs in two nearby colonies of Lesser Frigatebirds (Fregata ariel). A combination of stable isotope analysis and GPS tracking was used to assess dietary and spatial partitioning of foraging resources during the 2013 and 2014 breeding seasons. These results were compared to vessel‐derived estimates of prey availability, local primary productivity, and estimates of reproductive output to suggest potential drivers and implications of any observed partitioning. Isotopic data indicated a more neritic source of provisioned resources for near‐fledged chicks at an inshore colony, whereas their offshore counterparts were provisioned with resources with a more pelagic signal. Deep pelagic waters (>200 m) had higher availability of a preferred prey type despite a trend for lower primary productivity. Differences in foraging ecology between the two populations may have contributed to markedly different reproductive outputs. These findings suggest environmental context influences dietary and spatial aspects of foraging ecology. Furthermore, the effect of colony‐specific foraging patterns on population demography warrants further research.     

Pikeperch Sander lucioperca trapped between niches: foraging performance and prey selection in a piscivore on a planktivore diet

The foraging behaviour of planktivorous pikeperch Sander lucioperca during their first growing season was analysed. Field data showed that S. lucioperca feed on extremely rare prey at the end of the summer, suggesting the presence of a bottleneck. In experiments, foraging ability of planktivorous S. lucioperca was determined when fish were feeding on different prey types (Daphnia magna or Chaoborus spp.) and sizes (D. magna of lengths 1 or 2·5 mm) when they occurred alone. From these results, the minimum density requirement of each prey type was analysed. The energy gain for three different foraging strategies was estimated; a specialized diet based on either large D. magna or Chaoborus spp. or a generalist diet combining both prey types. Prey value estimates showed that Chaoborus spp. should be the preferred prey, assuming an energy maximizing principle. In prey choice experiments, S. lucioperca largely followed this principle, including D. magna in the diet only when the density of the Chaoborus spp. was below a threshold value. Splitting the foraging bout into different sequences, however, resulted in a somewhat different pattern. During an initial phase, S. lucioperca captured both prey as encountered and then switched to Chaoborus spp. if prey density was above the threshold level. The prey selection observed was mainly explained by sampling behaviour and incomplete information about environmental quality, whereas satiation only had marginal effects. It was concluded that the observed diet based on rare prey items was in accordance with an optimal foraging strategy and may generate positive growth in the absence of prey fish in suitable sizes.     

Predicting the contributions of novel marine prey resources from angling and anadromy to the diet of a freshwater apex predator

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Feeding Performance in Heterochronic Alpine Newts is Consistent with Trophic Niche and Maintenance of Polymorphism

The feeding performances of two heterochronic morphs of the Alpine newt Triturus alpestris were investigated in laboratory experiments. Although both morphs are able to feed in the aquatic habitat, the hydrodynamics of prey capture differ between morphs. In paedomorphs water sucked with prey is expelled behind the mouth through gill bars. In metamorphs, water is expelled by the mouth as gill slits are closed. Feeding performance was better in paedomorphs than in metamorphs when foraging on aquatic crustaceans, but paedomorphs were less successful when foraging on terrestrial invertebrates caught at the water surface. These differences in prey capture success related to prey type allow the two morphs to use specific resources in their aquatic habitat. These results are consistent with previous studies that showed diet differentiation between morphs in natural populations. Such resource partitioning is a factor favouring the maintenance of facultative paedomorphosis in natural populations.