Size-biased allocation of prey from male to offspring via female: family conflicts, prey selection, and evolution of sexual size dimorphism in raptors

In birds with bi-parental care, the provisioning link between prey capture and delivery to dependent offspring is regarded as often symmetric between the mates. However, in raptors, the larger female usually broods and feeds the nestlings, while the smaller male provides food for the family, assisted by the female in the latter part of the nestling period, if at all. Prey items are relatively large and often impossible for nestlings to handle without extended maternal assistance. We video-recorded prey delivery and handling in nests of a raptor with a wide diet, the Eurasian kestrel Falco tinnunculus, and simultaneously observed prey transfer from male to female outside the nest. The male selectively allocated larger items, in particular birds and larger mammals, to the female for further processing and feeding of nestlings, and smaller items, in particular lizards and smaller mammals, directly to the nestlings for unassisted feeding. Hence, from the video, the female appeared to have captured larger prey than the male, while in reality no difference existed. The female’s size-biased interception of the male’s prey provisioning line would maximize the male’s foraging time, and maximize the female’s control of the allocation of food between her own need and that of the offspring. The male would maximize his control of food allocation by capturing smaller prey. This conflict would select for larger dominant females and smaller energy-efficient males, and induce stronger selection the longer the female depends on the male for self-feeding, as a proportion of the offspring dependence period.     

Dynamic changes in prey choice by stickleback during simultaneous encounter with large prey

When a three-spined stickleback Gasterosteus aculeatus encountered prey simultaneously the probability of hanging and the median pursuit time were greater than when prey were encountered sequentially. During simultaneous prey encounter fish did not choose to attack the more profitable prey but instead the nearer prey was handled first except when the difference between the two prey sizes was large. No difference was found in the level of total energetic intake by the fish regardless of prey size pairing. Fish that handled and ate the first prey of a pair in <5 s attacked the second prey with a high probability of success, demonstrating an opportunistic feeding strategy. Importantly however, the fish did not choose to maximize long term energy intake rate by eating both prey, but rather short-term considerations over the course of feeding took precedence. With an empty stomach, the probability of a fish eating ( P _eat) the first prey handled was high regardless of prey size. As stomach fullness increased, the P _eat the first prey handled decreased if it was the larger prey. Hence, the fish were unselective when the stomach was empty but thereafter there was a shift in preference towards the smaller prey. The decision of which prey to attack and eat appeared to be based on short-term energy considerations and the level of stomach fullness. This study demonstrates that feeding on a short-term scale is a crucial factor to take account of when analysing fish feeding during simultaneous prey encounter.     

The feeding success of cattle egrets in flocks

The feeding rates of grouped (<1.5 m from conspecifics) and solo (>5 m from conspecifics) cattle egrets (Bubulcus ibis) in loose flocks away from cows were compared, to test the hypothesis that grouped cattle egrets benefit from feeding on prey flushed inadvertently by nearby conspecifics. The flock feeding rates were also compared to those of grouped and solo egrets near cows, to determine the effects of flock membership on feeding rates. Birds in flocks captured prey faster than those with cows, and tended to capture larger prey, but field observations and captive experiments failed to show that the feeding success of flock members was enhanced by the hypothesized ‘beater’ effect. Increases in prey density, however, always resulted in higher feeding rates, so some cattle egret groups may form in response to local concentrations of prey. Prey size may also play a role in group formation, because birds in the field tended to feed at greater distances from their neighbours when larger prey were captured, regardless of prey density. When small groups did form among cattle egrets feeding on relatively large prey, group members occasionally captured prey items that had been discovered by nearby conspecifics. This behaviour was not observed among birds in dense aggregations, which fed on small, highly abundant prey. These data indicate that there is a potential cost associated with feeding too near others unless the prey are relatively small and abundant.     

Prey capture and processing behaviors vary with prey size and shape in Australian and subantarctic fur seals

When hunting at sea, pinnipeds should adapt their foraging behaviors to suit the prey they are targeting. We performed captive feeding trials with two species of otariid seal, Australian fur seals (Arctocephalus pusillus doriferus) and subantarctic fur seals (Arctocephalus tropicalis). This allowed us to record detailed observations of how their foraging behaviors vary when presented with prey items that cover the full range of body shapes and sizes encountered in the wild. Small prey were captured using suction alone, while larger prey items were caught in the teeth using raptorial biting. Small fish and long skinny prey items could then be swallowed whole or processed by shaking, while all prey items with body depths greater than 7.5 cm were processed by shaking at the water's surface. This matched opportunistic observations of feeding in wild Australian fur seals. Use of “shake feeding” as the main prey processing tactic also matches predictions that this method would be one of the only tactics available to aquatic tetrapods that are unable to secure prey using their forelimbs.     

Parental Sex Differences in Food Allocation to Junior Brood Members as Mediated by Prey Size

Individual offspring within a brood may receive different amounts of provisioning from the male and female parents. Some hypotheses suggest that this bias is the result of an active and adaptive choice by parents. An alternative hypothesis is that feeding biases arise as a result of a constraint of fitting large prey items into small gapes. In an experiment with pied flycatchers, Ficedula hypoleuca , we tested for sex-biased allocation to junior nestlings in asynchronous broods and whether this could be explained by active parental choice or by passive allocation according to prey size and gape size. In both control broods and broods with experimentally increased degree of asynchrony, prey types did not differ between parents but females brought smaller prey than males at younger but not older nestling stages. At younger but not older nestling stages, the majority of feeds to junior nestlings were from females, and the smaller nestlings consumed smaller prey than older siblings. However, there was no evidence of active preference of small nestlings by females as parents did not differ in the tendency to bypass a begging senior nestling in order to feed a junior nestling. Provisioning rates by females were lower than those by males when nestlings were young and we suggest that foraging time constraints caused by the need to brood offspring result in females bringing smaller prey than males. In turn, the larger prey brought by males was more often transferred to larger offspring after the smaller ones failed to swallow it. In such cases, 'preferential' feeding of small nestlings by females may simply be a passive side effect of foraging constraints and gape-size limitations.     

The effects of turbidity on prey consumption and selection of zooplanktivorous Gasterosteus aculeatus L.

It is well documented that reduced visibility caused by elevated turbidity can affect feeding of fish, yet the extent to which selective zooplanktivory is altered in turbid conditions remains ambiguous. In this study, we examined the influence of natural sediment-induced turbidity on the overall prey consumption and selective predation of a common brackish water littoral zooplanktivore, the particulate feeding three-spined stickleback (Gasterosteus aculeatus L.). We hypothesized that the effects of turbidity on prey consumption and prey type selection would be pronounced due to the vision-oriented feeding of this species and that these effects would differ between genders. Using aquarium experiments with three different groups of cladocerans and copepods varying in size and behavior, we studied prey consumption and selectivity of this key planktivore in varying turbidity treatments. Our results indicated significantly decreased total prey consumption in the high turbidity treatments, as well as altered selective feeding on copepods and an enhanced preference for larger cladocerans. We found gender-dependent differences in prey consumption, which are consistent with observations of other visually feeding fish with sexual size dimorphism. We conclude that high turbidity, such as that occurring in shallow coastal areas, may affect selective feeding in vision-oriented zooplanktivores and that these effects may be gender-related.     

Feeding habits of larval Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret (Israel) II. Experimental study

Interactions between the larvae of Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret, Israel, and their zooplankton prey were studied experimentally. Prey species preference and size selectivities were measured. Larvae were hatched in the lab from eggs collected in the field, and fed different food items in various concentrations. The food items included lake zooplankton, algae, and commercial pellets. It was shown that small, first feeding larvae (7–8.5 mm SL) prefer small bodied zooplankters (< 180 µ). The effect of these food sources on larval growth was measured. It was found that larval Mirogrex grew at a higher rate when fed zooplankton prey sized from 63 µ–250 µ. Food items smaller than 63 µ, larger than 250 µ and Scenedesmus sp., produced less than optimal growth rates. The importance of Mirogrex feeding habits and their potential influence on the Kinneret ecosystem is considered.     

Australian Fur Seals (Arctocephalus pusillus doriferus) Use Raptorial Biting and Suction Feeding When Targeting Prey in Different Foraging Scenarios

Foraging behaviours used by two female Australian fur seals (Arctocephalus pusillus doriferus) were documented during controlled feeding trials. During these trials the seals were presented with prey either free-floating in open water or concealed within a mobile ball or a static box feeding device. When targeting free-floating prey both subjects primarily used raptorial biting in combination with suction, which was used to draw prey to within range of the teeth. When targeting prey concealed within either the mobile or static feeding device, the seals were able to use suction to draw out prey items that could not be reached by biting. Suction was followed by lateral water expulsion, where water drawn into the mouth along with the prey item was purged via the sides of the mouth. Vibrissae were used to explore the surface of the feeding devices, especially when locating the openings in which the prey items had been hidden. The mobile ball device was also manipulated by pushing it with the muzzle to knock out concealed prey, which was not possible when using the static feeding device. To knock prey out of this static device one seal used targeted bubble blowing, where a focused stream of bubbles was blown out of the nose into the openings in the device. Once captured in the jaws, prey items were manipulated and re-oriented using further mouth movements or chews so that they could be swallowed head first. While most items were swallowed whole underwater, some were instead taken to the surface and held in the teeth, while being vigorously shaken to break them into smaller pieces before swallowing. The behavioural flexibility displayed by Australian fur seals likely assists in capturing and consuming the extremely wide range of prey types that are targeted in the wild, during both benthic and epipelagic foraging.     

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.     

Sexual divergence in diets and morphology in Fijian sea snakes Laticauda colubrina (Laticaudinae)

In the Fiji Islands, female yellow‐lipped sea kraits (Laticauda colubrina) grow much larger than males, and have longer and wider heads than do conspecific males of the same body length. This morphological divergence is accompanied by (and may be adaptive to) a marked sex divergence in dietary habits. Adult female sea kraits feed primarily on large conger eels, and take only a single prey item per foraging bout. In contrast, adult males feed upon smaller moray eels, and frequently take multiple prey items. Prey size increases with snake body size in both males and females, but the sexes follow different trajectories in this respect. Female sea kraits consume larger eels relative to predator head size and body length than do males. Thus, the larger relative head size of female sea kraits is interpreted as an adaptation to consuming larger prey items. Our results are similar to those of previous studies on American water snakes (natricines) and Australian file snakes (acrochordids), indicating that similar patterns of sex divergence in dietary habits and feeding structures have evolved convergently in at least three separate lineages of aquatic snakes.     

School size affects individual feeding success in three-spined sticklebacks (Gastevosteus aculeatus L.)

The effect of school size on the feeding success of individual three-spined sticklebacks was studied. We found that the proportion of fish feeding on benthic prey increased with school size and that fish in large schools tended to start feeding sooner than fish in small schools. The total number of strikes also increased in larger schools. Despite this evidence for a foraging benefit associated with school membership we propose that improved feeding returns do not alone explain stickleback schooling.     

Predation on the Invasive Copepod,Pseudodiaptomus forbesi,and Native Zooplankton in the Lower Columbia River: An Experimental Approach to Quantify Differences in Prey-Specific Feeding Rates

Invasive planktonic crustaceans have become a prominent feature of aquatic communities worldwide, yet their effects on food webs are not well known. The Asian calanoid copepod, Pseudodiaptomus forbesi, introduced to the Columbia River Estuary approximately 15 years ago, now dominates the late-summer zooplankton community, but its use by native aquatic predators is unknown. We investigated whether three species of planktivorous fishes (chinook salmon, three-spined stickleback, and northern pikeminnow) and one species of mysid exhibited higher feeding rates on native copepods and cladocerans relative to P. forbesi by conducting `single-prey’ feeding experiments and, additionally, examined selectivity for prey types with `two-prey’ feeding experiments. In single-prey experiments individual predator species showed no difference in feeding rates on native cyclopoid copepods (Cyclopidae spp.) relative to invasive P. forbesi, though wild-collected predators exhibited higher feeding rates on cyclopoids when considered in aggregate. In two-prey experiments, chinook salmon and northern pikeminnow both strongly selected native cladocerans (Daphnia retrocurva) over P. forbesi, and moreover, northern pikeminnow selected native Cyclopidae spp. over P. forbesi. On the other hand, in two-prey experiments, chinook salmon, three-spined stickleback and mysids were non- selective with respect to feeding on native cyclopoid copepods versus P. forbesi. Our results indicate that all four native predators in the Columbia River Estuary can consume the invasive copepod, P. forbesi, but that some predators select for native zooplankton over P. forbesi, most likely due to one (or both) of two possible underlying casual mechanisms: 1) differential taxon-specific prey motility and escape responses (calanoids > cyclopoids > daphnids) or 2) the invasive status of the zooplankton prey resulting in naivety, and thus lower feeding rates, of native predators feeding on invasive prey.     

A predator's costs of overcoming the confusion-effect of swarming prey

Swarms of prey are known to have a confusion effect on predators that increases with swarm density. Hungry sticklebacks (Gasterosteus aculeatus) overcome the confusion (Heller & Milinski 1979) except when frightened by a predator of their own (Milinski & Heller 1978). This study investigates whether the costs of overcoming the confusion consist of decreased attention for predators. Hungry sticklebacks detected a predator of their own less often when they fed upon a prey swarm of high density than when they attacked a low density swarm. They also overlooked the predator more frequently when they fed with a high rate than with a low one in the same high density. Thus, there is a cost of attacking high density swarms. These results confirm two necessary conditions of a model for balancing feeding rate and predation risk (Milinski & Heller 1978). The decreased attention is assumed to be due to the limited channel capacity of the nervous system in processing sensory information.     

A forceful upper jaw facilitates picking-based prey capture: biomechanics of feeding in a butterflyfish,Chaetodon trichrous

Biomechanical models of feeding mechanisms elucidate how animals capture food in the wild, which, in turn, expands our understanding of their fundamental trophic niche. However, little attention has been given to modeling the protrusible upper jaw apparatus that characterizes many teleost species. We expanded existing biomechanical models to include upper jaw forces using a generalist butterflyfish, Chaetodon trichrous (Chaetodontidae) that produces substantial upper jaw protrusion when feeding on midwater and benthic prey. Laboratory feeding trials for C. trichrous were recorded using high-speed digital imaging; from these sequences we quantified feeding performance parameters to use as inputs for the biomechanical model. According to the model outputs, the upper jaw makes a substantial contribution to the overall forces produced during mouth closing in C. trichrous. Thus, biomechanical models that only consider lower jaw closing forces will underestimate total bite force for this and likely other teleost species. We also quantified and subsequently modeled feeding events for C. trichrous consuming prey from the water column versus picking attached prey from the substrate to investigate whether there is a functional trade-off between prey capture modes. We found that individuals of C. trichrous alter their feeding behavior when consuming different prey types by changing the timing and magnitude of upper and lower jaw movements and that this behavioral modification will affect the forces produced by the jaws during prey capture by dynamically altering the lever mechanics of the jaws. In fact, the slower, lower magnitude movements produced during picking-based prey capture should produce a more forceful bite, which will facilitate feeding on benthic attached prey items, such as corals. Similarities between butterflyfishes and other teleost lineages that also employ picking-based prey capture suggest that a suite of key behavioral and morphological innovations enhances feeding success for benthic attached prey items.     

Feeding habits of the bluemouth, Helicolenus dactylopterus dactylopterus (Delaroche, 1809) (Pisces: Sebastidae) in the Portuguese coast

In order to investigate the feeding habits of Helicolenus dactylopterus dactylopterus along the continental Portuguese coast, a total of 619 individuals were sampled of which 60% contained food in their stomach and 35% had more than one prey item. Among the 81 prey items that were identified in the stomachs, benthic and benthopelagic prey prevail on this species diet. Acantephyra sp, Pasiphaea sp, mysidacea, and teleostei n.i. were the prey with the higher percent index of relative importance (%IRI) value. Three length groups (5?C20?cm, 21?C27?cm, and 28?C48?cm) were defined through cluster analysis of the mean abundance of prey items. A permutational MANOVA detected significant differences in the diet and stomach fullness index for TLG, season, and maturation stage. Smaller fishes had a generalized diet, feeding mainly on mysidacea changing their diet above 20?cm TL, where a major consumption of natantia was found. The larger individuals, >28?cm TL, present a less generalized diet with pisces as dominant prey group. Seasonally, natantia and pisces were the principal prey groups during spring and winter, respectively, while mysidacea and other crustaceans were predominant during the rest of the year. Mysidacea were also the main prey group for immature individuals while natantia and pisces were the principal prey groups to the other maturity stages. The results of this study indicate that H. d. dactylopterus has a diverse diet focused on small crustaceans such as misyds and as specimens grow shrimps and fishes become more consumed, with larger specimens having a more specialized diet. The different nutritional needs during spawning season also seemed to influence the feeding habits of H. d. dactylopterus.     

Optimal foraging and risk of claw damage: How flexible are shore crabs in their prey size selectivity?

Understanding the behavioural mechanisms that underlie prey size preference of predators is an essential component of unravelling the processes that govern predator-prey dynamics. In marine systems, despite being able to consume larger and more profitable prey, many molluscivorous predators show a preference for smaller, less profitable prey, most likely to minimize the risk of damaging feeding extremities. Here we assessed the flexibility of this prey size preference. We observed that shore crabs (Carcinus maenas) that were food deprived, and which were offered mussels (Mytilus edulis) of different sizes in dichotomous preference tests, preferred smaller, less profitable mussels. The same result was observed for crabs foraging with a conspecific competitor. Only crabs that were conditioned to feed on the larger, most profitable mussels shifted their prey size preference and ranked the most profitable mussels as highest. Although shore crabs showed flexibility in prey size preference, through which they would be able to cope with environmental variability, our results in general emphasize preference for smaller prey. We discuss the possibility that crabs maximize their long-term feeding rate, in which case it can be optimal to select these smaller mussels.     

Age-specific differences in the winter foraging strategies of rooks Corvus frugilegus

Summary Foraging efficiency and intraspecific competition were compared between wild adult and immature rooks Corvus frugilegus with respect to flock size. Behavioural time budgets, and observations of prey selection and prey energetic values revealed that adult rooks in large flocks (> 50 individuals) consumed smaller, less profitable prey, but allocated more time to feeding and fed at a faster rate and with greater success than adults in small flocks. By contrast, immature rooks in flocks of more than 30 individuals allocated proportionally less time to feeding, fed at a lower rate and fed with no increase in success rate than when foraging in smaller flocks. Agonistic encounters and the avoidance of adults by immature rooks appeared responsible for such inefficient foraging. Hence immature rooks showed a preference for smaller flocks (< 50 individuals) with low adult: immature ratios while adults preferred larger flocks (> 50 individuals). We discuss the possible influence of competitive disadvantages on immature rook distribution, flock composition and post-natal dispersal.     

Feeding ecology of the Bay Goby (Pisces: Gobiidae). Effects of behavioral,ontogenetic, and temporal variation on diet

The ontogenetic, diel, seasonal, and yearly variations in gut fullness, diet, and prey diversity for a California estuarine gobiid (Lepidogobius lepidus Girard) were examined. Also the feeding behavior of this species was described.Small (<50 mm, SL) and large (? 50 mm, SL) gobies consumed similar prey types in different proportions. Major prey items were polychaetes, harpacticoid copepods, gammarid amphipods, molluscs, and other crustaceans. Diets of large and small gobies were not significantly correlated, and larger fish had a more diverse diet. Small fish fed at all times while larger gobies fed primarily at night. Changes in diet may be related to differential prey preferences, feeding chronologies, and increases in fish size.Both large and small gobies displayed seasonal differences in diet and prey diversity. Year-to-year changes in diet also were noted for both size classes. The bay goby uses different feeding behaviors to capture sedentary and motile prey and appears to forage opportunistically. This behavior is probably advantageous in an environment which fluctuates drastically.     

Age-Related Changes in Nestling Diet of the Cooperatively Breeding Green Woodhoopoe

In many socially monogamous bird species, parents of altricial young respond to the increasing demands of growing nestlings by increasing their feeding rate and the size of prey items delivered and by altering the types of prey provided. In some cooperatively breeding species, similar changes in feeding rate and prey size have been documented. However, potential changes in the types of prey delivered, both as nestlings age and by different group members, remain largely unexplored. Moreover, studies rarely compare the diet fed to nestlings with that eaten by the provisioning adults themselves. Here, I show that green woodhoopoe ( Phoeniculus purpureus ) nestlings receive a smaller proportion of spiders and larger proportions of caterpillars and centipedes as they grow older. Both male and female adults delivered a higher proportion of spiders to young nestlings than they ate while self-feeding, probably in response to particular nutritional requirements of the chicks. However, only males altered the proportions of caterpillars and centipedes delivered, providing smaller proportions to young nestlings than eaten themselves. These prey items may be too large for young nestlings to handle, and males may make a greater adjustment in provisioning diet than females because they collect more caterpillars and centipedes than do females. Although there were sex differences in provisioning diet, there were no differences between same-sex breeders and helpers in terms of the overall proportions of prey delivered or the changes with nestling age. Hence, individuals of different reproductive status may be following the same provisioning rules, at least in terms of prey type.     

Diel feeding periodicity, daily ration and prey selection of a riverine population of juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

The diel feeding periodicity, daily ration and prey selection of juvenile chinook salmon, Oncorhynchus tshawytscha , were studied in relation to the available prey. Maximum dry weight of food intake occurred about dawn, when mayflies were the major prey, but the greatest number of freshly eaten prey occurred during the afternoon, when chironomids and terrestrial dipterans predominated. Feeding activity at night was low, with smaller mayflies comprising up to 50% of the prey. During the day the young salmon fed selectively on chironomids and the larger mayflies, while trichopterans and terrestrial taxa were under-represented in the diet. Food consumption over the 24-h period averaged 8.3% of the fish dry body weight. Prey abundance in the drift explained about 50% of the composition of the diet. Although the fish selected larger mayflies, size apparently was not a main criterion for selection because chironomids, although smaller than mayflies, were also frequently eaten. Previous dietary experience of the fish and the diel pattern of prey abundance appear to best explain the selective feeding of juvenile chinook salmon.