Networks of prey specialization in an Arctic monomorphic seabird

Generalist predator populations are sometimes made up of individuals that specialize on particular prey items. To examine specialization in thick‐billed murres Uria lomvia during self‐feeding we obtained stomach contents and muscle stable isotope values for 213 birds feeding close to five colonies in the Canadian Arctic. Adults were less specialized during self‐feeding than during chick‐provisioning. Nonetheless, particular specialists clustered together within the foraging network. While sexes showed similar levels of specialization, individuals of the same sex clustered together within the foraging network. The significant degree of clustering regardless of sex showed that individuals specializing on one prey item tend to also specialize on another, although network topology varied from colony to colony. Adult muscle stable isotope values correlated with the stable isotope values of the prey found in stomachs, at least at the one colony with relevant prey data, suggesting that specializations are maintained over time. Degree of specialization increased with niche width across the five colonies, but similarity in gastro‐intestinal and bill morphology was independent of dietary similarity. Thus, although individual specialization is thought to play a key role in sympatric speciation through trophic specialization, we found no support for an association between morphology and foraging patterns in our species. We conclude that self‐feeding murres show clustered dietary specialization, and that specialization is highest where diet is most diverse.     

Foraging and feeding ecology of Platanista: an integrative review

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Individual patterns of prey selection and dietary specialization in an Arctic marine carnivore

The cumulative effect of individual‐level foraging patterns may have important consequences for ecosystem functioning, population dynamics and conservation. Dietary specialization, whereby an individual exploits a subset of resources available to the rest of the population, can develop in response to environmental or intrinsic population factors. However, accurate assessment of individual diets may be difficult because analyses of recent food intake may misrepresent foraging variability within a heterogeneous environment. We used quantitative fatty acid signature analysis (QFASA) and a novel index of longitudinal dietary change to examine the individual foraging patterns of 64 polar bears Ursus maritimus successively sampled in Western and Southern Hudson Bay between 1994–2003. Estimated diets varied between and within age and sex classes, with adult male polar bears consuming significantly more bearded seal Erignathus barbatus than adult female or subadult bears, whose diets were dominated by ringed seal Pusa hispida. Among individual adult males, consumption of bearded seal accounted for 0–98% of the diet and bearded seal consumption was positively correlated with individual dietary specialization, as measured by proportional similarity (PS_i) to the rest of the population. Most individual diets were consistent from year‐to‐year and were therefore not a product of short‐term heterogeneity in prey distribution. However, a novel dietary change index indicated that adult male polar bears had the most temporally variable diets with 23% of adult males switching their diet from predominantly ringed seal to predominantly bearded seal or vice versa. We conclude that QFASA is well‐suited to analyses of individual‐level foraging because it reflects an animal's diet over the preceding weeks to months. The subpopulations of bears in this study were near the southern limit of their species range and have experienced negative individual‐ and population‐level impacts related to sea ice loss and climate warming. The tightly constrained diets of some individuals, particularly adult females and subadults, may make them especially sensitive to future climate change.     

What you need is what you eat? Prey selection by the bat Myotis daubentonii

Optimal foraging theory predicts that predators are selective when faced with abundant prey, but become less picky when prey gets sparse. Insectivorous bats in temperate regions are faced with the challenge of building up fat reserves vital for hibernation during a period of decreasing arthropod abundances. According to optimal foraging theory, prehibernating bats should adopt a less selective feeding behaviour – yet empirical studies have revealed many apparently generalized species to be composed of specialist individuals. Targeting the diet of the bat Myotis daubentonii, we used a combination of molecular techniques to test for seasonal changes in prey selectivity and individual‐level variation in prey preferences. DNA metabarcoding was used to characterize both the prey contents of bat droppings and the insect community available as prey. To test for dietary differences among M. daubentonii individuals, we used ten microsatellite loci to assign droppings to individual bats. The comparison between consumed and available prey revealed a preference for certain prey items regardless of availability. Nonbiting midges (Chironomidae) remained the most highly consumed prey at all times, despite a significant increase in the availability of black flies (Simuliidae) towards the end of the season. The bats sampled showed no evidence of individual specialization in dietary preferences. Overall, our approach offers little support for optimal foraging theory. Thus, it shows how novel combinations of genetic markers can be used to test general theory, targeting patterns at both the level of prey communities and individual predators.     

Specialized foraging habits of adult female California sea lions Zalophus californianus

Individual foraging specialization occurs when organisms use a small subset of the resources available to a population. This plays an important role in population dynamics since individuals may have different ecological functions within an ecosystem related to habitat use and prey preferences. The foraging habitat fidelity and degree of specialization of California sea lions (Zalophus californianus) were evaluated by analyzing the stable isotopes values of carbon and nitrogen in vibrissae collected from 16 adult females from the reproductive colony on Santa Margarita Island, Magdalena Bay, Mexico, in 2012 and 2013. Based on the degree of individual specialization in δ¹⁵N, 62.5% of the females assessed can be considered specialist consumers focusing on the same prey or different prey from the same trophic level. The degree of individual specialization in δ¹³C indicated that 100% of the individuals showed fidelity to their foraging habitat as some fed in the lagoon, others foraged along the coast, and a third group preferred prey from the pelagic environment during both the breeding and nonbreeding seasons, suggesting diversification of foraging areas. Foraging area fidelity persisted despite the 2°C increase in the sea surface temperature over the course of the study period.     

Community structure affects trophic ontogeny in a predatory fish

While most studies have focused on the timing and nature of ontogenetic niche shifts, information is scarce about the effects of community structure on trophic ontogeny of top predators. We investigated how community structure affects ontogenetic niche shifts (i.e., relationships between body length, trophic position, and individual dietary specialization) of a predatory fish, brown trout (Salmo trutta). We used stable isotope and stomach content analyses to test how functional characteristics of lake fish community compositions (competition and prey availability) modulate niche shifts in terms of (i) piscivorous behavior, (ii) trophic position, and (iii) individual dietary specialization. Northern Scandinavian freshwater fish communities were used as a study system, including nine subarctic lakes with contrasting fish community configurations: (i) trout‐only systems, (ii) two‐species systems (brown trout and Arctic charr [Salvelinus alpinus] coexisting), and (iii) three‐species systems (brown trout, Arctic charr, and three‐spined sticklebacks [Gasterosteus aculeatus] coexisting). We expected that the presence of profitable small prey (stickleback) and mixed competitor–prey fish species (charr) supports early piscivory and high individual dietary specialization among trout in multispecies communities, whereas minor ontogenetic shifts were expected in trout‐only systems. From logistic regression models, the presence of a suitable prey fish species (stickleback) emerged as the principal variable determining the size at ontogenetic niche shifts. Generalized additive mixed models indicated that fish community structure shaped ontogenetic niche shifts in trout, with the strongest positive relationships between body length, trophic position, and individual dietary specialization being observed in three‐species communities. Our findings revealed that the presence of a small‐sized prey fish species (stickleback) rather than a mixed competitor–prey fish species (charr) was an important factor affecting the ontogenetic niche‐shift processes of trout. The study demonstrates that community structure may modulate the ontogenetic diet trajectories of and individual niche specialization within a top predator.     

Feeding ecology and trophic segregation of two sympatric mesopredatory sharks in the heavily exploited coastal ecosystem of the Adriatic Sea

The ecological roles and trophic interactions of two commercially important mesopredatory shark species, Squalus acanthias and Mustelus punctulatus that co‐occur on the continental shelf of the north‐central Adriatic Sea were investigated. Both shark species are dietary specialists, with a significant dietary overlap recorded only during the spring season. They showed different patterns of feeding as they grew: S. acanthias extended its trophic niche with an increase in size, while M. punctulatus developed a more specialized diet. These two sharks partition food resources and reduce niche overlap by foraging at different trophic levels. Mustelus punctulatus is a crustacean feeder, specialized in foraging on scavenging malacostracans frequently found along trawl tracks or on discards in the Adriatic fishing zone. Conversely, S. acanthias prefers small pelagic fishes, which are commercially exploited and in decline. The different foraging strategies adopted by these two species suggest that they should be managed separately. Dietary specialization, direct competition with humans for prey and their higher intrinsic vulnerability make S. acanthias particularly susceptible to the effects of anthropogenic perturbations.     

Foraging habits in a generalist predator: Sex and age influence habitat selection and resource use among bottlenose dolphins (Tursiops truncatus)

This study examines resource use (diet, habitat use, and trophic level) within and among demographic groups (males, females, and juveniles) of bottlenose dolphins (Tursiops truncatus). We analyzed the δ¹³C and δ¹⁵N values of 15 prey species constituting 84% of the species found in stomach contents. We used these data to establish a trophic enrichment factor (TEF) to inform dietary analysis using a Bayesian isotope mixing model. We document a TEF of 0‰ and 2.0‰ for δ¹³C and δ¹⁵N, respectively. The dietary results showed that all demographic groups relied heavily on low trophic level seagrass‐associated prey. Bayesian standard ellipse areas (SEA_b) were calculated to assess diversity in resource use. The SEA_b of females was nearly four times larger than that of males indicating varied resource use, likely a consequence of small home ranges and habitat specialization. Juveniles possessed an intermediate SEA_b, generally feeding at a lower trophic level compared to females, potentially an effect of natal philopatry and immature foraging skills. The small SEA_b of males reflects a high degree of specialization on seagrass associated prey. Patterns in resource use by the demographic groups are likely linked to differences in the relative importance of social and ecological factors.     

Does body size influence thermal biology and diet of a python (Morelia spilota imbricata)?

Current theory predicts that larger‐bodied snakes not only consume larger prey (compared with smaller individuals), but may also have a different range of prey available to them due to their thermal biology. It has been argued that smaller individuals, with lower thermal inertia (i.e. faster cooling rates at nightfall when air temperature falls and basking opportunities are limited), may be thermally restricted to foraging and hunting during the day on diurnally active prey, and have reduced capacity to hunt crepuscular and nocturnal prey species. This predictive theory was investigated by way of dietary analysis, assessment of thermal biology and thermoregulation behaviour in an ambush forager, the south‐west carpet python (Morelia spilota imbricata, Pythonidae). Eighty‐seven scats were collected from 34 individual pythons over a 3‐year radiotelemetry monitoring study. As predicted by gape size limitation, larger pythons took larger prey; however, 65% of prey items of small pythons were represented by nocturnally active, small mammals, a larger proportion than present in larger snakes. Several measures of thermal biology (absolute body temperature, thermal differential of body temperature to air temperature, maximum hourly heating and cooling rates) were not strongly affected by python body mass. Additionally, body temperature was only influenced by the behavioural choice of microhabitat selection and was not affected by python body size or position, suggesting that these behavioural choices do not allow smaller pythons to vastly increase their temporal foraging window. By coupling dietary analysis, measures of body temperature and behavioural observations of free‐ranging animals, we conclude that, contrary to theoretical predictions, a small body size does not thermally restrict the temporal window for ambush foraging in M. s. imbricata. An ontogenetic or size‐determined switch from ambush feeding to actively foraging on slower prey would account for the differences in prey taken by these animals. The concept of altered foraging behaviour warrants further investigation in this species.     

Feeding rate and attack specialization: the roles of predator experience and energetic tradeoffs

Synopsis Individual mosquitofish, Gambusia affinis, can adopt a broad range of attack selectivities. In part, this variation can be explained by the past experiences of a fish. Individuals selected the more profitable Ceriodaphnia dubia (Cladocera) over less profitable cyclopoid copepods to a greater degree after being exposed to both prey types than did individuals experienced with only one of the prey types. Feeding rate (biomass ingested per unit time) declined with increased attack specialization on the profitable prey (Ceriodaphnia) when such prey were scarce, a result in agreement with assumptions of optimal diet theory. When profitable prey were abundant feeding rate was a bimodal function of the intensity of specialization on profitable prey; fish that specialized on cyclopoid copepods (the less profitable prey type) fed at higher rates than did generalists. This may be the result of antagonistic learning that precluded feeding efficiently on more than one type of prey at a time. The data are consistent with the hypothesis that rejection of unsuitable prey involves a time cost. The two preceeding aspects of foraging behavior, which are absent from most optimal diet models, could lead to failure in predicting the attack specialization of some predators, An additional aspect of the results was the generally weak relationship between feeding efficiency and specialization behavior. This suggests that feeding rate may not have been as tightly linked to the specialization behavior a predator adopts as is assumed by current foraging theory.     

Assessing niche partitioning of co‐occurring sibling bat species by DNA metabarcoding

Niche partitioning through foraging is a mechanism likely involved in facilitating the coexistence of ecologically similar and co‐occurring animal species by separating their use of resources. Yet, this mechanism is not well understood in flying insectivorous animals. This is particularly true of bats, where many ecologically similar or cryptic species coexist. The detailed analysis of the foraging niche in sympatric, cryptic sibling species provides an excellent framework to disentangle the role of specific niche factors likely involved in facilitating coexistence. We used DNA metabarcoding to determine the prey species consumed by a population of sympatric sibling Rhinolophus euryale and Rhinolophus mehelyi whose use of habitat in both sympatric and allopatric ranges has been well established through radio tracking. Although some subtle dietary differences exist in prey species composition, the diet of both bats greatly overlapped (O_jk = 0.83) due to the consumption of the same common and widespread moths. Those dietary differences we did detect might be related to divergences in prey availabilities among foraging habitats, which prior radio tracking on the same population showed are differentially used and selected when both species co‐occur. This minor dietary segregation in sympatry may be the result of foraging on the same prey‐types and could contribute to reduce potential competitive interactions (e.g., for prey, acoustic space). Our results highlight the need to evaluate the spatial niche dimension in mediating the co‐occurrence of similar insectivorous bat species, a niche factor likely involved in processes of bat species coexistence.     

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.     

Stock‐specific variation of trophic position,diet and environmental stress markers in Atlantic salmon Salmo salar during feeding migrations in the Baltic Sea

This study investigated stock‐specific variation in selected ecophysiological variables during the feeding migrations of Atlantic salmon Salmo salar in the Baltic Sea. Oxidative stress biomarkers and EROD (ethoxyresorufin‐O‐deethylase, Cyp1A enzyme) activity were used as indicators of possible environmental stress and stable isotopes as determinants of diet and trophic position. Latvian S. salar stocks Daugava and Gauja had distinct stable‐isotope signatures compared to the other stocks, indicating differences in migration patterns, residency or arrival times, or dietary specialization among stocks. Salmo salar originating from Daugava and Gauja also had lower catalase enzyme activity than the other stocks. Post‐smolts originating from rivers of the Gulf of Finland had elevated EROD activities compared to fish of the same age from Bothnian Bay rivers, which could indicate exposure to organochlorine pollutants. No other stock‐specific differences in oxidative stress biomarkers were found. The study demonstrates how genetic, oxidative stress biomarker, EROD and stable‐isotope data may be combined to study trophic position, prey prevalence and environmental stress of mixed S. salar stocks foraging in the sea.     

Optimization of foraging and diet by the piscivorous Othos dentex (Serranidae)

The aim of this study was to determine the dietary characteristics and mouth morphology of Othos dentex and to use these data, together with in situ observations of feeding behaviour, to elucidate how foraging and diet are optimized by this piscivorous serranid. Seasonal spear and line fishing over reefs in south‐western Australia yielded 426 O. dentex (total length, L_T, 183–605 mm), among which the stomachs of 95 contained food. The food in the stomachs of 76 fish was sufficiently undigested to be seen to contain, almost invariably, a single fish prey, which was typically identifiable to family and often to species. The prey of O. dentex, which were measured (L_T), represented 10 families, of which the Labridae and Pempheridae constituted nearly two‐thirds of the prey volume. Two‐way crossed analysis of similarities of volumetric data for stomach contents showed that the dietary compositions of the different length classes of O. dentex in the various seasons were significantly related to length class of prey, but not to prey family, length class within the various prey families or season. Furthermore, an inverse (Q‐mode) analysis, including one‐way analysis of similarities, showed that the patterns in the prey consumed by the different length classes of O. dentex in the various seasons were related more strongly to length class than prey family. The former trend is exemplified in a shade plot, by a marked diagonality of the length classes of prey with increasing predator size. The ingestion of typically a single teleost prey, whose body size increases as that of O. dentex increases, reduces the frequency required for seeking prey, thus saving energy and reducing the potential for intraspecific competition for food. The ability of O. dentex to ingest large prey is facilitated by its possession of a very large gape, prominent recurved teeth, dorsal and independently‐moveable eyes, cryptic colouration and effective ambush behaviour. Othos dentex has thus evolved very cost‐effective mechanisms for optimizing its foraging and diet.     

Interspecific variation in distributions and diets of coral reef butterflyfishes (Teleostei: Chaetodontidae)

This study examined within‐reef distributions for 19 species of butterflyfishes (Chaetodontidae) at Lizard Island, northern Great Barrier Reef, Australia, and compared spatial patterns of abundance among species with contrasting diets. Spatial variation in abundance of butterflyfishes was most prominent among physiognomic reef zones mainly due to significant zonation of eight species, including four obligate hard‐coral feeders (Chaetodon trifascialis, Chaetodon baronessa, Chaetodon plebeius and Chaetodon lunulatus) and four generalist species (Chaetodon auriga, Chaetodon citrinellus, Chaetodon kleinii and Chaetodon rafflesi). Distributions of obligate hard‐coral feeders were closely associated with spatial variation in percentage cover of scleractinian corals, but no more restricted compared with facultative hard‐coral feeders or non‐coral feeders. Species with highest dietary specialization (C. trifascialis and C. baronessa), however, exhibited the most pronounced zonation patterns and were restricted to habitats with greatest abundance of their preferred prey. While there are conspicuous links between dietary specialization v. spatial patterns in abundance of butterflyfishes, it remains unclear whether dietary specialization is the cause or consequence of more restricted distributions.     

Why do eastern curlews Numenius madagascariensis feed on prey that lowers intake rate before migration?

Eastern curlews Numenius madagascariensis spending the nonbreeding season in eastern Australia foraged on three intertidal decapods: soldier crab Mictyris longicarpus, sentinel crab Macrophthalmus crassipes and ghost‐shrimp Trypaea australiensis. Due to their ecology, these crustaceans were spatially segregated (=distributed in ‘patches’) and the curlews intermittently consumed more than one prey type. It was predicted that if the curlews behaved as intake rate maximizers, the time spent foraging on a particular prey (patch) would reflect relative availabilities of the prey types and thus prey‐specific intake rates would be equal. During the mid‐nonbreeding period (November–December), Mictyris and Macrophthalmus were primarily consumed and prey‐specific intake rates were statistically indistinguishable (8.8 versus 10.1 kJ×min^?1). Prior to migration (February), Mictyris and Trypaea were hunted and the respective intake rates were significantly different (8.9 versus 2.3 kJ×min^?1). Time allocation to Trypaea‐hunting was independent of the availability of Mictyris. Thus, consumption of Trypaea depressed the overall intake rate. Six hypotheses for consuming Trypaea before migration were examined. Five hypotheses: the possible error by the predator, prey specialization, observer overestimation of time spent hunting Trypaea, supplementary prey and the choice of higher quality prey due to a digestive bottleneck, were deemed unsatisfactory. The explanation for consumption of a low intake‐rate but high quality prey (Trypaea) deemed plausible was diet optimisation by the curlews in response to the pre‐migratory modulation (decrease in size/processing capacity) of their digestive system. With a seasonal decrease in the average intake rate, the estimated intake per low tide increased from 1233 to 1508 kJ between the mid‐nonbreeding and pre‐migratory periods by increasing the overall time spent on the sandflats and the proportion of time spent foraging.     

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.     

Habitat selection by the Cape clawless otter (Aonyx capensis) in rivers in the Western Cape Province, South Africa

We radio‐tracked seven Cape clawless otters (Aonyx capensis) (Schinz, 1821) in two rivers in the Western Cape Province, South Africa, providing data on their habitat selection. Habitat type was investigated at a scale that enabled us to separate the effects of types of riparian vegetation, geomorphology and anthropogenic influences. Otters selected areas with boulders and/or reed beds, which provided high crab density and shelter. Direct observations showed that they used two foraging modes depending on the habitat selected. Otters could select open water within c. 8 m of the shore, dive and surface with or without prey. Otherwise hunting involved them moving into shallow water (c. 0.2 m deep), and walking along the substrate feeling for prey with their forefeet. Disturbed possible prey items were then caught with the forefeet.     

Feather corticosterone reveals stress associated with dietary changes in a breeding seabird

Changes in climate and anthropogenic pressures might affect the composition and abundance of forage fish in the world's oceans. The junk‐food hypothesis posits that dietary shifts that affect the quality (e.g., energy content) of food available to marine predators may impact their physiological state and consequently affect their fitness. Previously, we experimentally validated that deposition of the adrenocortical hormone, corticosterone, in feathers is a sensitive measure of nutritional stress in seabirds. Here, we use this method to examine how changes in diet composition and prey quality affect the nutritional status of free‐living rhinoceros auklets (Cerorhinca monocerata). Our study sites included the following: Teuri Is. Japan, Middleton Is. central Gulf of Alaska, and St. Lazaria Is. Southeast Alaska. In 2012 and 2013, we collected “bill loads” delivered by parents to feed their chicks (n = 758) to document dietary changes. We deployed time–depth–temperature recorders on breeding adults (n = 47) to evaluate whether changes in prey coincided with changes in foraging behavior. We measured concentrations of corticosterone in fledgling (n = 71) and adult breeders' (n = 82) feathers to determine how birds were affected by foraging conditions. We found that seasonal changes in diet composition occurred on each colony, adults dove deeper and engaged in longer foraging bouts when capturing larger prey and that chicks had higher concentrations of corticosterone in their feathers when adults brought back smaller and/or lower energy prey. Corticosterone levels in feathers of fledglings (grown during the breeding season) and those in feathers of adult breeders (grown during the postbreeding season) were positively correlated, indicating possible carryover effects. These results suggest that seabirds might experience increased levels of nutritional stress associated with moderate dietary changes and that physiological responses to changes in prey composition should be considered when evaluating the effect of prey quality on marine predators.