DIET OF HARBOR PORPOISES IN THE KATTEGAT AND SKAGERRAK SEAS: ACCOUNTING FOR INDIVIDUAL VARIATION AND SAMPLE SIZE

Stomach contents of 112 bycaught harbor porpoises ( Phocoena phocoena ) collected between 1989 and 1996 in the Kattegat and Skagerrak seas were analyzed to describe diet composition and estimate prey size, to examine sample size requirements, and to compare juvenile and adult diets. Although porpoises preyed on a variety of species, only a few contributed substantially to the diet. Atlantic herring ( Clupea harengus ) was the dominating prey species for both juveniles and adults. Our results, in combination with those from previous studies, suggest that where herring is a dominant food source, porpoises prey primarily on size classes containing mature or maturing individuals. Further, we also show that Atlantic hagfish ( Myxine glutinosa ) may be an important food resource, at least for adult porpoises. Examination of sample size requirement showed that, depending on the taxonomic level used to describe the diet, a minimum of 35–71 stomachs are needed to be confident that all common prey species will be found.     

Prey attack and predators defend: counterattacking prey trigger parental care in predators

That predators attack and prey defend is an oversimplified view. When size changes during development, large prey may be invulnerable to predators, and small juvenile predators vulnerable to attack by prey. This in turn may trigger a defensive response in adult predators to protect their offspring. Indeed, when sizes overlap, one may wonder "who is the predator and who is the prey"! Experiments with "predatory" mites and thrips "prey" showed that young, vulnerable prey counterattack by killing young predators and adult predators respond by protective parental care, killing young prey that attack their offspring. Thus, young individuals form the Achilles' heel of prey and predators alike, creating a cascade of predator attack, prey counterattack and predator defence. Therefore, size structure and relatedness induce multiple ecological role reversals.     

ONTOGENETIC SHIFT IN PREY SIZE SELECTION AND ITS RELATION TO PROFITABILITY IN NUCELLA LAPILLUS

According to the energy maximization premise an animal shouldchoose its diet to maximize its net energy gain per unit handlingtime (E/T_b). Previous studies on adult marine gastropods haveshown this index of profitability to be a monotonicaUy increasingfunction of prey size, yet they select smaller, apparently sub-optimalprey. Laboratory experiments were used to investigate the profitabilityof different sized mussels, as prey, to juvenile Nucella lapillususing two separate criteria. First the ability to promote growthand second the more usual energy gain per unit handling time. The major difference between the two measures of prey valuewas that E/T_h predicted an optimum prey size larger than thegrowth rate model. Selection experiments were used to comparepreferences exhibited by developing Nucella with the spectraof profitability defined from the two models. In general thewhelks' choice of prey size was best predicted by the growthrate model of prey value. Field studies illustrated a functionalrelationship between size of mussel eaten and size of whelk,the mean size of mussel eaten again conformed more closely tomaximization of growth rate. The energetic return in the E/T_h model is generally taken asthe amount of food ingested, however gross growth efficiencywas found to be a decreasing function of ingestion rate. Sinceingestion rate was an increasing function of prey size, theE/T_h model overestimated the value of large prey by ignoringthe efficiency with which food is used by the predator.     

Accelerating the onset of piscivory: intersection of predator and prey phenologies

Piscivorous fishes tend to be able to consume other fishes early in development and generally experience a dramatic increase in growth after the ontogenetic diet shift to piscine prey. Hence, an acceleration of the onset of piscivory may be favoured strongly by natural selection. Temperate freshwater piscivores, for example, becomes piscivorous at a relatively young age by spawning in advance of, and thereby achieving a size advantage over, the young of their piscine prey. Research in various North American estuaries suggests that young-of-the-year (YOY) bluefish Pomatomus saltatrix , an offshore-spawning estuarine-dependent marine fish, may accelerate the onset of piscivory by being advected to higher latitudes and timing their estuarine entry with the appearance of small coastal fishes. This hypothesis was tested by: (i) determining the annual recruitment date of YOY bluefish and their prey; and (ii) examining the diet and prey size preferences, and predator size-prey size relationships, of YOY bluefish in two different estuarine systems: Great South Bay, and the lower Hudson River. Results suggest that the relationships between bluefish and their prey are determined by a complex interplay between recruitment timing of both predator and prey, prey size availability, predator selectivities, and the timing of vernal warming. It is concluded that YOY bluefish migration into northern estuaries at an advanced size provides them with a predatory size advantage over their principal piscine prey thereby facilitating an early diet shift to piscivory white minimizing the time spent as planktivores.     

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.     

Prey profitability for adult Grey Herons Ardea cinerea and the constraints on prey size when feeding young nestlings

Patterns of handling time and profitability are examined for adult Grey Herons feeding on carp, eels and catfish. Handling times generally increased with prey size but were influenced markedly by the morphology and behaviour of the prey. Profitability was highest for carp (max. 0.9 g/s for 15–20 cm fish), lowest for catfish (max. 0.05 g/s) and intermediate for eels (max. 0.1 g/s). Nestlings were unable to ingest the sizes of fish most profitable for the adults to consume until aged 20 days; by the age of 30 days, they could consume the full size-range of prey taken by the adults. In order to feed their young chicks, adults must therefore either select smaller prey, or break their large prey into smaller pieces. The diet of nestling Grey Herons in the Camargue is examined for evidence to support or refute the former hypothesis.
Young nestlings (≤20 days) regurgitated smaller carp than old nestlings (> 20 days). Comparison of prey types in the diet of the two groups showed that small prey species occurred significantly more often in the diet of young chicks, while the converse was true for larger prey species. The occurrence of particular prey types in the diet only of young chicks suggests that adults may forage in more marginal, shallower water (where small prey are probably more abundant) to meet the requirements of their brood during the early part of the nestling phase. The second hypothesis, that the adults break down large prey into smaller pieces, was not examined, although evidence from other studies suggests that this does occur; both mechanisms may therefore be important.     

Foraging ecology of the California gnatcatcher deduced from fecal samples

The California gnatcatcher is a threatened species essentially restricted to coastal sage scrub habitat in southern California. Its distribution and population dynamics have been studied intensely, but little is known about its diet. We identified arthropod fragments in 33 fecal samples of the California gnatcatcher to gain insight into its foraging ecology and diet. Fecal samples were collected from adult males, adult females, fledglings, and nestlings. Leaf- and planthoppers (Homoptera) and spiders (Araneae) predominated numerically in samples. Spider prey was most diverse, with eight families represented. True bugs (Hemiptera) and wasps, bees, and ants (Hymenoptera) were only minor components of the gnatcatcher diet. Gnatcatcher adults selected prey to feed their young that was larger than expected given the distribution of arthropod size available in their environment, and chicks were provisioned with larger prey items and significantly more grasshoppers and crickets (Orthoptera) and spiders than adults consumed themselves. Both adults and young consumed more sessile than active prey. Further studies are needed to determine whether arthropods sampled in coastal sage scrub that are common in fecal samples are good indicators of California gnatcatcher habitat. Received: 30 December 1998 / Accepted: 28 April 1999     

Contrasting mortality in young freshwater leeches and triclads

A guild of leeches and triclads coexist and are the most numerous invertebrate predators on the stony shores of productive British lakes. Populations of all species are food-limited. Mortality of recruited young is considerably higher in leech than in triclad populations, and this paper investigates reasons for this. In particular, the feeding success of young leeches and triclads in relation to prey species, prey size, prey condition (alive or crushed), spatial heterogeneity (with or without the presence of stones or gravel), and the presence or absence of other young or adults predators (leeches or triclads) of the same or different species are investigated in the laboratory. Feeding success by young leeches and triclads on crushed prey without the presence of stones was high, but declined dramatically in leeches but not triclads when stones were present. Young leeches and triclads were inept at capturing live prey, of a small or large size, with the exception of soft-bodied prey such as oligochaetes. Feeding success by young predators on live prey was not increased by the presence of other young predators of the same or different species. With only a few exceptions, the presence of adult leeches, and to a much lesser extent adult triclads, increased the feeding success, growth and survival of young leeches and triclads. It is concluded that the high mortality of young leeches, compared to triclads, in field populations is due to their inability to locate damaged food in an environment with spatial heterogeneity due to a poorly developed chemosensory system. High and low levels of juvenile morality are accompanied by high and low reproductive rates in leech and triclad populations, respectively. It is unusual for a food limited population to have a high level of recruitment, but it is speculated that the characteristically high reproductive output in parasitic leeches, from which predaceous leeches are derived or have affinities, has been retained to counterbalance high juvenile mortality rates.     

Foraging Decision Rules and Prey Species Preferences of Northwestern Crows (Corvus caurinus)

Categorization of similar prey types and the application of decision rules by dietary generalists can enhance the efficiency of foraging decisions and facilitate the inclusion of novel prey types in the diet. While considerable research attention has been directed toward investigation of these concepts in invertebrates, few have assessed categorization and decision rules used by generalist vertebrate predators. In this study, we experimentally investigated decision rules and prey preferences of northwestern crows (Corvus caurinus) feeding on littleneck clams (Tapes philippinarum) and whelks (Nucella lamellosa). We presented crows with three species‐size combinations: small clams (2.0–2.9 cm length) paired with large whelks (4.0–4.9 cm), small clams paired with medium whelks (3.0–3.9 cm), and large clams (4.0–4.9 cm) with large whelks. Profitability estimates based on observations of crows feeding on these prey species indicated that clams were always the more energetically profitable option; however, in prey choice trials crows consistently selected the heavier prey species, regardless of differences in profitability. These results show that crows apply a general decision rule according to which they select heavier prey items when feeding on hard‐shelled prey requiring similar handling techniques, and that while such decision rules may approximate optimal choices they may not always follow predictions based solely on prey profitability. We discuss these results in the context of behavioural flexibility of generalist predators, and predicting impacts of intertidal avian predators on prey populations.     

Evaluation of Trail-Cameras for Analyzing the Diet of Nesting Raptors Using the Northern Goshawk as a Model

Diet studies present numerous methodological challenges. We evaluated the usefulness of commercially available trail-cameras for analyzing the diet of Northern Goshawks (Accipiter gentilis) as a model for nesting raptors during the period 2007–2011. We compared diet estimates obtained by direct camera monitoring of 80 nests with four indirect analyses of prey remains collected from the nests and surroundings (pellets, bones, feather-and-hair remains, and feather-hair-and-bone remains combined). In addition, we evaluated the performance of the trail-cameras and whether camera monitoring affected Goshawk behavior. The sensitivity of each diet-analysis method depended on prey size and taxonomic group, with no method providing unbiased estimates for all prey sizes and types. The cameras registered the greatest number of prey items and were probably the least biased method for estimating diet composition. Nevertheless this direct method yielded the largest proportion of prey unidentified to species level, and it underestimated small prey. Our trail-camera system was able to operate without maintenance for longer periods than what has been reported in previous studies with other types of cameras. Initially Goshawks showed distrust toward the cameras but they usually became habituated to its presence within 1–2 days. The habituation period was shorter for breeding pairs that had previous experience with cameras. Using trail-cameras to monitor prey provisioning to nests is an effective tool for studying the diet of nesting raptors. However, the technique is limited by technical failures and difficulties in identifying certain prey types. Our study also shows that cameras can alter adult Goshawk behavior, an aspect that must be controlled to minimize potential negative impacts.     

Winter diet of the little auk (Alle alle) in the Northwest Atlantic

The little auk (Alle alle) is one of the most numerous seabirds in the world, but their winter prey selection has never been thoroughly studied. In the present study, we analyzed the proventricular contents of 205 little auks caught in coastal areas off southwest Greenland during December–February and off Newfoundland in March. Large Calanoid copepods are known to be the main prey during summer. We found krill to become the crucial winter prey in both areas, followed by Themisto spp. and young capelin (Mallotus villosus). No difference was found between the diet of juvenile and adult birds. Copepods constituted around 1 % of the diet and close to all copepods were in birds caught near Cape Farwell in December. These findings provide new and important insight into the forage ecology of the little auk, and they support other studies showing that large Calanoid copepods in the Arctic decent for winter hibernation at depths that are below the diving range of the little auks. More studies to determine offshore diet and annual variation are, however, needed in order to get a more complete picture.     

Optimal diet selection,frequency dependence and prey renewal

This paper extends existing models of frequency-dependent diet selection by considering the optimal diet selection of a predator feeding upon prey populations which can be depleted but are also capable of renewal (e.g. immigration, growth, or reproduction). This model and existing models which include prey depletion, predict partial-preference and a generic diet preference for the commonest prey types (apostatic selection). Unlike previous diet selection models, it is found that the optimal diet selection of an individual predator can be to favour the rarest prey type (anti-apostatic selection) when encounter rates are high, even if the individual prey do not differ in their nutritional value. Studies have demonstrated that predators generally show apostatic selection, even when all prey have the same nutritional value. Anti-apostatic selection has also been observed when prey are crowded, and therefore at high density, consistent with the idea of high encounter rates. This anti-apostatic diet selection has previously been proposed as evidence for the use of prey search images by a predator, or variation in individual prey preference. In this paper it is suggested that prey renewal is a further factor, often confounded in experiments, which could favour anti-apostatic selection.     

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

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

The use of integrated constructed wetlands (ICW) for the treatment of separated swine wastewaters

Haplochromine cichlids used to be the main prey of the introduced Nile perch, Lates niloticus, in Lake Victoria. After depletion of the haplochromine stocks at the end of the 1980s, Nile perch shifted to the shrimp Caridina nilotica and to a lesser degree to its own young and the cyprinid Rastrineobola argentea. In the present study, we investigated the Nile perch diet in the northern Mwanza Gulf after resurgence of some of the haplochromine species and compared it with data collected in the same area in 1988/1989. It became clear that haplochromines are again the major prey of Nile perch. The dietary shift from invertebrate feeding (shrimps) to feeding on fish (haplochromine cichlids) occurs at a smaller size than it did when Nile perch were taking primarily dagaa and juvenile Nile perch as their fish prey. The apparent preference for haplochromines as prey has reduced the degree of cannibalism considerably, which may have a positive impact on Nile perch recruitment.     

Body-size evolution on islands: are adult size variations in tiger snakes a nonadaptive consequence of selection on birth size?

Mean adult size has been used as the traditional measure of body size to explain trends of insular gigantism and dwarfism in a wide array of taxa. However, patterns of variation in body size at birth have received surprisingly little attention, leaving open the possibility that adult body-size differences are nonadaptive consequences of selection acting on neonate body size. Here I used an empirical and correlative approach to test this hypothesis in a mosaic of 12 island and mainland snake populations in Australia. Data collected on 597 adult and 1,084 neonate tiger snakes showed that (1) both adult and neonate mean body sizes varied strongly across populations; (2) prey diversity and size convincingly explained birth-size variations: birth size-notably, gape size-correlated with prey size; (3) neonate snout-vent length was significantly correlated with neonate gape size; and (4) neonate snout-vent length was significantly correlated with adult snout-vent length. Postnatal growth rates recorded under common-garden conditions differed across populations and were correlated with mean prey size. These data collectively suggest that (1) prey size is the main driver for the evolution of body size at birth in gape-limited predators, (2) adult size variations may reflect selective forces acting on earlier life stages, and (3) adult size variations may also reflect resource availability during ontogeny (notably, prey diversity).     

Using molecular and crowd-sourcing methods to assess breeding ground diet of a migratory brood parasite of conservation concern

Breeding ground food availability is critical to the survival and productivity of adult birds. The common cuckoo Cuculus canorus is a brood-parasitic Afro-Palearctic migrant bird exhibiting long-term (breeding) population declines in many European countries. Variation in population trend between regions and habitats suggests breeding ground drivers such as adult food supply. However, cuckoo diet has not been studied in detail since before the most significant population declines in Europe began in the mid-1980s. 20th century studies of cuckoo diet largely comprised field observations likely to carry bias towards larger prey taxa. Here we demonstrate the potential value of 1) using high-throughput DNA sequencing of invertebrate prey in faeces to determine cuckoo diet with minimal bias towards large prey taxa, and 2) using crowd-sourced digital photographs from across Britain to identify lepidopteran cuckoo prey taxa during recent years post-decline (2005–2016). DNA analysis found a high frequency of Lepidoptera, including moths of family Lasiocampidae, prominent within the past literature, but also grasshoppers (Orthoptera) and flies (Diptera) that may be overlooked by field observation methodologies. The range of larval lepidopteran prey identified from photographs largely agreed with those previously documented, with potential signs of reduced diversity, and identities of key adult prey taxa were supported by molecular results. Notably, many identified cuckoo prey taxa have shown severe declines due to agricultural intensification, suggesting this has driven spatial patterns of cuckoo loss. Landscape-scale, lowland rewilding interventions provide opportunities to understand the scale of reversal of previous agricultural intensification that may be necessary to restore prey populations sufficiently to permit recolonization by cuckoos.     

Feeding habits of grey gurnard,Eutrigla gurnardus (L., 1758), along the Catalan coast (northwestern Mediterranean)

The stomach content of 589 specimens of Eutrigla gurnardus (collected at quarterly intervals) have been analyzed to determine diet according to fish size and season. The results show that crustaceans (% No = 93%, %Weight = 81%) are the basic food and that teleostean fishes are quite important in the adult diet. The importance of nycthemeral migrators decreases significantly as size increases, whilst the importance of benthic prey increases inversely. In every size, the feeding intensity is greater at midday than in the morning. Larger prey are eaten in winter than in summer although feeding intensity is lower in winter.     

Comparative morphology and diet of young cichlids in the dammed Sinnamary river, French Guiana, South America

The growth of most mensural characters of Krobia guianensis and Crenicichla saxatilis during early development was explained by a split regression indicating inflection in allometry at specific standard lengths. Double-centred PCA revealed morphological transformations during ontogeny mostly under the influence of the maximum body depth and the maximum caudal peduncle width, the area of the caudal fin and the horizontal diameter of the eye contributing also, but to a lesser extent. Young K. guianensis switched to larger prey at the end of the period when the growth of most of their studied mensural characters demonstrated an inflection, but young C. saxatilis changed their diet at a much smaller size than that at which an inflection occurred. However, in the lower reservoir sites, most young juveniles of K. guianensis and C. saxatilis did not switch to larger items during their ontogeny and micro-crustaceans occurred more frequently in their diet. It is hypothesized that the low net energy gained when eating such small prey may explain why the young of both species were thinner at these sites than in their undisturbed habitats. If food resources of young cichlids do not improve in the Petit-Saut reservoir, it is hypothesized that these species will develop stunted populations there.     

Prey size of single-prey loaders as an indicator of prey abundance

In this paper I am concerned with the behaviour of seabirds that bring back just one prey item at a time to their young. I use a simple model from central place foraging theory to show that the size of fish that a parent bird brings back may increase or decrease with an increase in the abundance of fish. This means that it may not be possible to use the size of fish that is fed to the young as an indicator of prey abundance.     

The food of the Nile perch, Lates niloticus (L.), after the disappearance of the haplochromine cichlids in the Nyanza Gulf of Lake Victoria (Kenya)

After the disappearance of the haplochromine species in the Nyanza Gulf of Lake Victoria as a result of predation by Lates niloticus , the latter has turned its attention to aquatic invertebrates and other fish. Changes in the diet of the Nile perch with increase in its size have been observed: young L. niloticus preyed mostly on invertebrates, including crustaceans and various small aquatic insects; large, immature L. niloticus supplemented the invertebrate diet with both young and small fish; adults above 80 cm total length were mainly piscivorous. L. niloticus feeds on fish prey of about one third its own length.
The tendency of L. niloticus to switch from one prey item to another, depending on availability, is reported; e.g., in the Nyanza Gulf, the prey diet has shifted from the haplochromine to Caridina nilotica and L. niloticus juveniles.