The foraging ecology of two pairs of congeneric demersal fish species: importance of morphological characteristics in prey selection

The feeding habits of two sympatric species pairs of demersal fish ( Mullus barbatus-Mullus surmuletus, Serranus cabrilla-Serranus hepatus ) which occupy the shallow coastal area (25–30 m) in Iraklion Bay were investigated from samples collected on a monthly basis (August 1990 to August 1992). Stomach content analyses revealed that all of them were carnivores, feeding mainly on benthic invertebrates, and that each species consumed a narrow range of prey species with no significant dietary overlap. The morphology of their feeding apparatus was compared to examine the effect of any morphological differences on food selection and resource partitioning between the fish species. The species could be distinguished on the basis of the size of their mouth gape, the number of gill rakers and the length of their intestine. This study shows that each species pair follows a different strategy segregating along food niche dimensions. In particular, M. barbatus and M. surmuletus segregate their feeding niche consuming different prey taxa with similar sizes whereas S. cabrilla and S. hepatus differ considerably with respect to the degree to which prey species contribute to their diets coupled with differences in mean prey sizes.     

Feeding characteristics of two tintinnid ciliate species on phytoplankton including harmful species: effects of prey size on ingestion rates and selectivity

The feeding abilities of two tintinnid ciliates, Favella ehrenbergii and Favella taraikaensis, on 10 species of flagellates including harmful marine algae were examined under single prey conditions, and selective feeding of F. taraikaensis on two species of algae of different sizes was investigated under mixed prey conditions. Ingestion rates calculated from the rate of increases of auto-fluorescent particles inside food vacuoles in individuals ranged from 0.5 to 22.1 cells ind(-1) h(-1) for F. ehrenbergii and from 0.8 to 44.9 cells ind(-1) h(-1) for F. taraikaensis on nine species of prey algae. Significant ingestion rates of both Favella species could not be detected on Heterosigma akashiwo, although some individuals of both species were observed ingesting H. akashiwo in the initial incubation period. The relationship between prey cell volume and ingestion rate could be expressed mathematically for both Favella species, indicating that it is possible to estimate the potential feeding activity of each Favella species on specific algae using an equation, and may be applicable to evaluate the food value of prey alga for both Favella species. When F. taraikensis was fed mixtures of H. circularisquama and Pavlova lutheri, significant ingestion rates of H. circularisquama by F. taraikaensis could not be measured when H. circularisquama accounted for less than 20% of the other prey biomass. However, clear selectivity for H. circularisquama was observed when H. circularisquama reached and exceeded 34% of the other prey biomass. Under mixed prey conditions, it is likely that the selectivity of F. taraikaensis is stronger for larger prey compared to prey algae with a size near the lower limit on which F. taraikaensis can feed.     

Swan foraging shapes spatial distribution of two submerged plants,favouring the preferred prey species

Compared to terrestrial environments, grazing intensity on belowground plant parts may be particularly strong in aquatic environments, which may have great effects on plant-community structure. We observed that the submerged macrophyte, Potamogeton pectinatus, which mainly reproduces with tubers, often grows at intermediate water depth and that P. perfoliatus, which mainly reproduces with rhizomes and turions, grows in either shallow or deep water. One mechanism behind this distributional pattern may be that swans prefer to feed on P. pectinatus tubers at intermediate water depths. We hypothesised that when swans feed on tubers in the sediment, P. perfoliatus rhizomes and turions may be damaged by the uprooting, whereas the small round tubers of P. pectinatus that escaped herbivory may be more tolerant to this bioturbation. In spring 2000, we transplanted P. perfoliatus rhizomes into a P. pectinatus stand and followed growth in plots protected and unprotected, respectively, from bird foraging. Although swan foraging reduced tuber biomass in unprotected plots, leading to lower P. pectinatus density in spring 2001, this species grew well both in protected and unprotected plots later that summer. In contrast, swan grazing had a dramatic negative effect on P. perfoliatus that persisted throughout the summer of 2001, with close to no plants in the unprotected plots and high densities in the protected plots. Our results demonstrate that herbivorous waterbirds may play a crucial role in the distribution and prevalence of specific plant species. Furthermore, since their grazing benefitted their preferred food source, the interaction between swans and P. pectinatus may be classified as ecologically mutualistic.     

Prey size,prey perishability and group foraging in a social spider

Summary Selection might favor group foraging and social feeding when prey are distributed in patches that do not last long enough for a solitary individual to consume more than a small fraction of them (Pulliam and Millikan 1982; Pulliam and Caraco 1984). Here we considered the foraging behavior of a social spider, Anelosimus eximius, in light of this ephemeral resource hypothesis. This species builds large webs in which members cooperate to capture a wide variety of different sizes and types of prey, many of which are very large. The capture success of this species was very high across all prey sizes, presumably due to the fact that they foraged in groups. Group consumption times in natural colonies for all prey larger than five mm were less than the time that dead insects remained on the plastic sheets that we used as artificial webs. Solitary consumption estimates, calculated from the rate at which laboratory individuals extracted insect biomass while feeding, were the same as the residence times of insects on artificial webs in the field for insects between 6 and 15 mm in length and were significantly longer than the persistence of insects on plastic sheets for all larger insects. Large prey, that contribute substantially to colony energy supplies, appeared to be ephemeral resources for these spiders that could not be consumed by a single spider in the time they were available. These factors made the food intake of one spider in a group less sensitive to scavenging by others and could act to reinforce the social system of this species.     

Intra- and interspecific dominance hierarchies and variation in foraging tactics of two species of stream-dwelling chars

Aggressive interactions, foraging behavior, habitat use and diet were studied in sympatric populations of white-sported char,Salvelinus leucomaenis, and Dolly Varden,Salvelinus malma, in a Japanese mountain stream. Underwater observations on individuals of both species revealed two distinct behavioral regimes: aggressive drift foragers and non-aggressive benthos foragers. Aggressive drift foragers defended partial territories around focal points from which they made forays to capture invertebrates drifting in the water column. Non-aggressive benthos foragers cruised around and beneath cobble in large foraging ranges that overlapped each other. Intra- and interspecific, size-dependent dominance hierarchies were recognized among aggressive drift foragers, whereas non-aggressive benthos foragers showed no such relationships. Terrestrial invertebrates were the most abundant prey in the diets of drift foragers, whereas a very small proportion of the diet of benthos foragers was made up of these taxa. Benthos foragers showed more complex diet composition than drift foragers. These results suggest that non-aggressive benthos foragers may avoid not only interference but also exploitative competition by using alternative foraging tactics. The proportion of drift foragers to benthos foragers among white-spotted char was more than 35 times that among Dolly Varden. The significant difference in the proportion of each species using the two types of foraging strategy results in interspecific food segregation in sympatric populations.     

Fluctuations in prey density: effects on the foraging tactics of scolopendrid centipedes

Foraging animals are faced with the problem of acquiring information about prey populations and utilizing that information in making foraging decisions. In this paper the effect of variation in prey density on the search tactics and mechanisms of prey density assessment in the centipede Scolopendra polymorpha are examined. Centipedes exhibited a prey density-dependent repertoire of search tactics. After 50 min of exposure to high prey density, centipedes switched from active search to ambush-like tactics, while maintaining a high rate of search at the lowest prey density. An initial period of sampling of prey density was involved in the switch in search behaviour and it is suggested that the encounter rate with prey was the key element in density assessment. When the prey density was changed from low to high, centipedes switched from active search to ambush tactics and when prey density was changed from high to low centipedes switched from ambush to active search within 40 min. Such behaviour may decrease the unreliability of sampling information and the risk involved in foraging decisions in variable environments.     

The foraging ecology of sympatric marine fish in the genus Embiotoca (Embiotocidae): Importance of foraging behavior in prey size selection

Summary Two locally sympatric temperate marine reef fishes, Embiotoca jacksoni and E. lateralis (Embiotocidae), have high taxonomic similarity in diets. Subdivision of gammarid amphipods, their principal prey, was found. E. jacksoni took more tubicolous gammarid amphipods whereas E. lateralis consumed mostly free-living individuals. The species differed considerably with respect to between-individual variability in taxonomic compositions of their diets. Each E. jacksoni closely resembled other conspecifics in this regard while individual E. lateralis displayed very high between-fish variation. The principal interspecific difference in fish diets concerned the sizes of prey items taken. E. jacksoni ate small but very common items and the mean prey weight in their guts did not differ from random collections of available prey. E. lateralis concentrated on large, rarer sizes such that the average prey weight in their guts was much heavier than available or in the diet of E. jacksoni of the same length. Disparate foraging behaviors was a much better indicator of the relative differences in diets of these two fishes than was external fish morphology. E. jacksoni, which can winnow prey items from unwanted debris, was a relatively indiscriminant forager. E. lateralis did not winnow but actively searched for prey. This species was a much more discriminating forager, but displayed much variability in foraging behavior.     

The effects of human presence, flock size and prey density on shorebird foraging rates

Animals may alter their foraging behaviour in the presence of humans because they perceive humans as potential predators. In this study I determined whether people caused shorebirds to reduce feeding rates at a stopover site in coastal British Columbia, Canada. I controlled for prey density and flock size because these variables may influence both the foraging rates as well as the effect of human disturbance on feeding efficiency. Semipalmated plovers decreased feeding rates when there were more people on the beach (multiple regression: F_1,15=5.86, b=0.59, P=0.029, R²=37.6%). For least sandpipers, the effect of human densities on feeding rates depended on flock size (F_1,21=5.97, P=0.023) and amphipod availability (F_1,21=4.98, P=0.037). This study demonstrated the importance of measuring subtle behavioural changes in foraging rates along with key ecological variables in order to assess the true impact of human disturbance on migratory shorebirds.     

Claw morphology, prey size selection and foraging efficiency in generalist and specialist shell-breaking crabs

Claw morphology, and claw-closing forces of four species of intertidal crabs from San Juan Island, Washington were compared and related these findings were related to prey size selection, shell breaking times and total handling times on their snail prey, Littorina sitkana Philippi. Two functional groups of crabs emerged: generalists and specialists on hard-shelled prey. The generalist, Hemigrapsus nudus (Dana), has an omnivorous diet and possesses weak claws with small, fine denticles and mechanical advantage (MA) of the claw's lever system <0.3, while the specialists, Lophopanopeus bellus (Stimpson), Cancer oregonensis (Dana) and C. productus (Randall), consume hard-shelled prey and possess large, powerful claws with broad, blunt molars and MA>0.3. The claws of the generalist, H. nudus, exhibited weaker claw closing forces (5 N) than those of similar sized specialists (>12 N). When crabs of similar weight were offered four size categories of Littorina sitkana, the generalist, Hemigrapsus nudus, exhibited a consistent preference for the smallest size categories, while the three specialists attacked all size classes offered. Hemigrapsus nudus took significantly longer (134 s) than the specialists (30–52 s) to break open a 4 mm L. sitkana. This difference in shell-breaking time between the generalist and the specialists increased with increasing prey size. The rate of successful attacks on increasingly larger L. sitkana decreased with prey size in the generalist (70% on 4 mm, 37% on 6 mm, and 0% on 8 mm snails), but remained high in the specialists (70–100%). Strength limitation of the claws is the best hypothesis to explain the avoidance of large snails by the generalist, H. nudus.     

Seasonality, optimal foraging, and prey coexistence

Several empirical studies suggest that herbivores may promote coexistence between plants by relaxing the strength of resource competition. In contrast, recent mathematical models predict that food-limited herbivory instead cause exclusion through apparent competition, regardless of whether herbivore selectivity is constant or density dependent. This study extends existing theory to consider a strongly seasonal system. Herbivores with fixed diet preferences have the same effect regardless of seasonality, but there is a marked difference when the diet selectivity of herbivores conforms to a simple optimal-foraging model. An optimally foraging herbivore in a seasonal environment is able to promote plant coexistence among many species. The mechanism involves diet switching, occurring over narrow density intervals. For this to have an effect in a nonseasonal model, equilibrium resource densities must be in this interval, which requires close parameter fitting. In seasonal environments, resource densities change through the year and may frequently move across narrow regions in which diet changes occur. The potential of gray-sided voles to promote coexistence between two arctic dwarf shrubs is evaluated in terms of the model. For this system, it is shown that vole herbivory has the potential to reverse competitive dominance.     

Spatial distributions of multiple plant species affect herbivore foraging selectivity

Spatial distribution of food resources is an important factor determining herbivore foraging. Previous studies have demonstrated that clumped distribution of preferred species increases its consumption by herbivores in single‐ or two‐species systems. However, the potential impact of distribution pattern of less preferred species on foraging was ignored. In natural grasslands with high species diversity and complexity, the spatial distribution of preferred species impacts on herbivore foraging may be strongly correlated with the distribution of less preferred species. Our aims were to determine the effect of distribution of both preferred and other plant species on herbivore foraging under conditions close to a native, multi‐species foraging environment, and conceptualize the relationships between spatial distribution of food resources and herbivore consumption. We hypothesized that random distribution of non‐preferred species reduces herbivore consumption of preferred species because the dispersion of less preferred species likely disturbs herbivore foraging. We conducted an experiment using three species with five combinations of clumped and random distribution patterns. Three species Lathyrus quinquenervius, Phragmites australis and Leymus chinensis, were of high, intermediate and low preferences by sheep, respectively. Results showed that distribution of low preferred species, but not that of high preferred one, affected the consumption of preferred species. Sheep obtained higher consumption of high preferred species when low preferred species followed a clumped distribution than a random distribution. Distance between aggregations of high and low preferred species did not affect sheep foraging. It was concluded that the effects of spatial distribution of preferred species on its consumption are dependent on herbivore foraging strategy, and sheep can consume more preferred species when there is a consistent spatial pattern between preferred species and the entire food resource, and that the random dispersion of low preferred species in grassland may reduce herbivore consumption of high preferred species, thus minimizing selective grazing.     

Optimal foraging on perilous prey: risk of bill damage reduces optimal prey size in oystercatchers

Intake rate maximization alone is not always sufficient in explainingprey size selection in predators. For example, bivalve-feedingoystercatchers regularly select smaller prey than expected ifthey aimed to maximize their intake rate. It has been proposedthat to these birds large prey are "risky," in the sense thatbirds may damage their bills when feeding on large bivalves.Large bivalves yield more energy, but according to this hypothesisthis is achieved at the expense of energy yield in the longterm when (1) the risk of bill damage increases with prey sizeand (2) foraging with a damaged bill is less effective. In accordancewith this hypothesis, we show that captive oystercatchers feedingon large cockles experienced a high probability of bill tipdamage, while bill damage was absent when cockles were small.Moreover, among free-living oystercatchers the prevalence ofbill damage was correlated with mean cockle size near the capturesite, and the data on captive birds fit in this pattern. Foodintake of captive oystercatchers feeding exclusively on cockleswas reduced by 23% after bill damage, and free-living birdswith damaged bills had 14 g lower mass. Because lower body masswas associated with higher mortality probability, these resultsindicate long-term costs associated with feeding on large cockles.We conclude that the risk of bill damage can potentially explainwhy oystercatchers avoid large bivalves and that oystercatchersmay maximize long-term intake rate by selecting prey sizes thatare "suboptimal" from a short-term rate-maximizing point ofview.     

Morphological limitations, prey size selectivity, and growth response of juvenile atlantic salmon, Salmo salar

The head and jaw movements involved in capture, buccal manipulation, ingestion and rejection of prey were investigated using sequential photography of juvenile Atlantic salmon feeding in a simulated stream environment. The results are described and discussed and mouth breadth and gill raker spacing are proposed as morphometric limitations to the range of prey sizes available which remains constant at 0·06 · fish fork length ( PFR ).
A recirculatory flume tank was used to study prey size selectivity behavior. Simplified downstream-drifting prey items elicited a variety of responses depending on their physical size. One hundred percent of offered prey of PFR 0·025 were ingested, while 90 % of prey at PFR 0·051 and 100% of prey at PFR 0·105 were rejected. It is demonstrated that fish show negative selection for prey sizes smaller than PFR 0·025 and that prey of this size elicits maximum growth response.
The validity of the proposed morphometric limitations on the available prey sizes is demonstrated by reference to selectivity behaviour and prey size related differential growth.     

The dynamics of prey choice in fish: the importance of prey size and satiation

Over a number of decades the process of prey choice has been investigated using fishes as model predators. Using fishes for the model has allowed the proximate factors that determine how a mobile predator finds and chooses to eat the prey encountered within a variable 3‐D environment to be estimated. During prey choice a number of constraints exist, in particular most fish predators will eat their prey whole thus their jaws and gut create functional limitations once a prey has been attacked. By considering the relationship between the size of the prey and the predator's feeding apparatus and feeding motivation this study explores the link between mechanistic studies and theoretical, optimal foraging based predictions. How the prediction of prey choices made by the fish following prey encounter can be reconciled with what is likely to be found in the fish's stomach is discussed. This study uses a progression of empirical examples to illustrate how the limits of functional constraints and prey choice at different stages of motivation to feed can be taken into account to improve predictions of predator prey choice.     

Sibling cannibalism among juvenile vundu under controlled conditions. I. Cannibalistic behaviour, prey selection and prey size selectivity

Sibling cannibalism among vundu Heterobranchus longifilis larvae started at the age of 4 days, with the prey caught tail-first then swallowed up to the head, which was eventually discarded (type I cannibalism). At 17 days old, this type of cannibalism vanished and was replaced by the ingestion of the whole prey (type II cannibalism), which could only be exerted by predators six times as heavy as their prey. Type II cannibalism consisted of a seemingly opportunistic ambush attack by a formerly passive predator towards a disorientated prey. It required no preliminary aggression or chase, or even contact with the prey, suggesting that the attack was not mediated by the tactile sense, and that cannibalism was independent of aggressive behaviour. When alternative food resources (formulated feed, live tilapia prey) were available, the intensity of cannibalism decreased but pellet-eaters or tilapia predators always achieved lower growth rates than those feeding on conspecifics, suggesting that cannibalism was the most advantageous foraging tactic. Losses to cannibalism among populations of 30-day old juvenile vundu with an initial ratio of 4% of cannibals were as high as 75·5–79·9% over 15 days. Predation peaked during the first days (up to 2·8 prey C⁻¹ day⁻¹), then vanished progressively as surviving prey grew faster than cannibals and escaped their predation. Cannibals preferred consuming the largest prey available with respect to the logistics of cannibalism (body weight ratio of 6·0). This preference for large prey was interpreted both as a foraging tactic aiming to maximize the energetic return, and as foraging strategy enabling the cannibals to exploit their prey as long as possible. Based on these data, comprehensive models of the impact of cannibalism on vundu populations were developed.     

The effect of prey abundance and foraging tactics on the population dynamics of a social, territorial carnivore, the spotted hyena

We used naturally occurring spatial and temporal changes in prey abundance to investigate whether the foraging behavior of a social, territorial carnivore, the spotted hyena ( Crocuta crocuta ), conformed to predictions derived from the ideal free distribution. We demonstrate that hyenas in the Ngorongoro Crater, Tanzania, redistributed themselves from less profitable to more profitable areas, even when this required them to undertake foraging trips to areas beyond their clan territory boundary, or required normally philopatric females to emigrate. As expected for a system with rank related access to food resources in the territory, females of low social status foraged more often outside their territory and were more likely to emigrate than dominant females. Probably because Crater hyenas regularly foraged outside their territories, there was no correlation between clan size and prey density within territories, suggesting that clan sizes may have exceeded the carrying capacity of territories. A substantial decline of the hyena population in the Crater from 385 adults in the mid 1960s to 117 in 1996 was most likely due to a substantial decline of their main prey. The decline in the hyena population was associated with a decline in the size of clans but not in the number of clans. The number of clans probably remained constant due to emigration by females from large clans into vacant areas or clans with no adult females, and because hyenas regularly fed in areas containing concentrations of prey beyond their territory boundary. Between 1996 and 2003 annual recruitment rates of Crater hyenas consistently exceeded annual mortality rates, resulting in an almost doubling of the adult population. This increase was most likely due to an increase in prey abundance, a relatively low level of predation on hyenas by lions ( Panthera leo ), and an absence of high levels of disease related mortality.     

Predation by Neomysis mercedis: effects of temperature, Daphnia magna size and prey density on ingestion rate and size selectivity

1. Neomysis mercedis predation rates on Daphnia magna were determined under laboratory conditions. There were generally no consistent differences between the number of Daphnia ingested at 10 and 14°C. 2. At each temperature, the number of prey consumed increased with mysid size and decreased with Daphnia size. 3. For small prey the relationship between ingestion rate and prey density represented a Type II functional response. However, for larger prey there was no significant relationship between density of prey and consumption by mysids. 4. The pattern of size-selective predation by Neomysis was studied to test the optimal foraging hypothesis. For prey populations with mixed size classes, the smallest size of prey was consumed most frequently but intermediate size prey provided the greatest biomass. These observations are contrary to our predictions based on calculations of profitability of different sizes of prey.