The food and feeding ecology of Blue petrel (Halobaena caerulea) and Dove prion (Pachyptila desolata)

The diet of two small burrow dwelling petrels, Dove prion, Pachyptila desolata and Blue petrel Halobaena caerulea , was studied at the subantarctic island of South Georgia by quantitative analysis of 246 food samples regurgitated by adults about to feed chicks. Crustaceans, and particularly krill Euphausia superba , predominate in both species' diet but fish is considerably more important to the Blue petrel. Other dietary differences between the species chiefly involve the relative proportions of krill, copepods (nearly exclusive to Dove prion) and seven amphipod species taken.
There is no overlap in the species' chick feeding periods but food differences are likely to reflect prey selection rather than resource availability; differences in prey size are ascribed to use of different feeding techniques particularly relating to bill structure and function. Data on frequency of chick feeding, proportion of oil and presence of pumice in food samples are used to indicate that the Blue petrel feeds further away from the breeding colony than the Dove prion. The significance of the various adaptive differences is discussed.     

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

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

COMPARISON OF THE DIET OF BREEDING AND NONBREEDING CAPE GANNETS MORUS CAPENSIS

Berruti, A. 1991. Comparison of the diet of breeding and nonbreeding Cape Gannets Morus capensis. Ostrich 62:8-12.

The effects of breeding status on the diet composition of Cape Gannets Morus capensis were tested at two colonies in the western Cape, South Africa. At both colonies, there were no significant differences in the species composition of the diet, but the mean energy content of regurgitations of breeders was significantly greater. Although the regurgitations of breeders were heavier and contained more fish at both colonies, only the number of fish per regurgitation from breeders at one colony was significantly greater. Birds feeding chicks at Malgas Island took significantly shorter fish than nonbreeders. The small effect of breeding status on prey species composition shows that temporal and geographical variation in prey availability is more important than breeding status in affecting diet cornposition of the Cape Gannet.     

Diet and feeding habits of Economidichthys pygmaeus (Perciformes: Gobiidae) in Lake Pamvotis,NW Greece

The purpose of the study was to define the diet composition, feeding preferences, seasonal, size and sex related changes in diet, feeding strategy and diel cycle of Economidichthys pygmaeus. Important features of the overall feeding patterns of the goby include (i) feeding activity, (ii) bottom habits and (iii) consumption of food items to identify the dietary breadth. Stomach contents of 533 specimens, 13.42–48.65 mm total length (TL), collected by hand net (2 mm mesh size) from October 2006 to September 2007 were analyzed. The percentage of 76 empty stomachs (14.26%) did not vary significantly with season. The food composition suggested that the goby is a carnivorous fish. Prey items identified in the stomachs belonged to four major groups: Crustaceans, Insects, Mollusca and Plants. Copepods were the most important (%IRI = 57.51), especially in fish larger than 23 mm TL. Chironomids (%IRI = 43.23) constituted the main prey for fish <23 mm TL. Diet composition showed little seasonal variation. Copepoda dominated the diet in autumn and winter (43.4%; 82.4%) and were replaced by Chironomidae larvae in spring (39.7%) and summer (47.7%). During the reproductive season, large males show a narrow food spectrum as opposed to large females, reflecting the different activity pattern exhibited by males and females during the breeding season, since the former supports parental care. According to the modified Costello graphical method, specialist individuals form the goby population that feeds on two preferential prey types (e.g. Copepods, Chironomids). Nevertheless, they consume some occasional prey (e.g. Cladocera, Insects, Gastropods). The species foraged chiefly at night and early morning, during lower light intensity. The highest feeding activity recorded at night (02.00 hours) and during early morning (08.00 hours) could be related to the cryptic behaviour displayed by the species.     

Unchanging diet in a stable colony: contemporary and past diet composition of black-legged kittiwakes <Emphasis Type="Italic">Rissa tridactyla</Emphasis> at Helgoland,south-eastern North Sea

In contrast to the situation at the west coast of the North Sea, the breeding colony of black-legged kittiwakes Rissa tridactyla at Helgoland in the south-eastern North Sea did not exhibit severe declines since 1990 but instead numbers increased and only lately stabilised. Declines at the west coast of the North Sea were attributed to a lower abundance and lower quality of the key prey, sandeels. We hypothesised that kittiwakes at Helgoland do not rely as heavily on sandeels as their conspecifics. We analysed stomach contents of nestlings and adults of 2001, 2002, 2004 and 2006. In concordance with earlier studies of the 1980s and 1990s, young whiting Merlangius merlangus was the most important prey species in 2001, 2002 and 2004. Clupeids and sandeels were consumed in lower proportions. While earlier studies suggested whiting to originate from fisheries discards, evidence now supports that kittiwakes prey upon whiting in areas of hydrographic fronts. No whiting was recovered in samples of 2006 and the proportion of fish prey was low. Main prey items were polychaete worms (Nereidae), which were presumably consumed as swarming Heteronereis stages. An observed strong rise in water temperature in summer 2006 might have influenced food availability of kittiwakes by inducing swarming of Nereidae. Overall, kittiwakes breeding on Helgoland showed a positive population trend for several decades while mainly feeding on whiting.     

The effects of diet and stocking density on the growth and behaviour of pond pre-reared pikeperch under intensive conditions

Experiments were carried out between 1999 and 2001 on pikeperch fingerlings using two feeds at four stocking densities to evaluate the success of training 35‐day‐old pond‐reared pikeperch, Stizostedion lucioperca L., to lifeless feed in intensive culture. Training periods of 28 days for each of three cohorts (1999–2001) were applied to introduce pikeperch from zooplankton to minced feed. In a preliminary study (1999), two feeds (live prey and minced fish meat) were compared at two stocking densities. Two higher densities were investigated in 2000 and 2001 and matched with the minced fish diet. Different feeds had significant effects on feed consumption and, as a consequence, on body weight gain; fish fed live prey showed highest growth rates. Unexpectedly, the stocking density had no significant effect on growth, and at the highest density there was a significantly betterfeed conversion rate. Stocking density and feed type significantly influenced the feeding behaviour. Three characteristic foraging techniques are described.     

Consequences of foraging trip duration on provisioning behaviour and fledging condition of common murres Uria aalgae

We examine the provisioning constraints of a pursuit‐diving seabird in a cold ocean regime by comparing the behaviour of common murres Uria aalge rearing chicks at two colonies in the Northwest Atlantic during 1998‐2000. Funk Island is the largest (340,000–400,000 breeding pairs) and most offshore (60 km) colony of common murres in eastern Canada. Seventy‐five percent of the Northwest Atlantic population of common murres breeds on this island. Great Island is one island within the Witless Bay Ecological Reserve, which is the second largest breeding aggregation (100 000 breeding pairs) and is located near‐shore (2 km). The primary forage fish species in Newfoundland waters is capelin Mallotus villosus, which spawns on or near coastal beaches during summer. Therefore, the two study colonies differ in their distance to food resources and colony size. It is within this natural context that we compare: (1) prey types and frequency of delivery (amount of prey), (2) parental time budgets, and (3) the mass and condition (mass/wing length) of fledglings at both colonies. Similarly sized female capelin (100–150 mm) were delivered to chicks at both colonies. Foraging time per day per parent, a proxy of foraging effort, was similar at both colonies (Great Island: 5.1 h; Funk Island: 5.5 h), as was the percentage of time spent with mates (Great Island: 12.3%; Funk Island: 10.9%). Foraging trips, however, were longer at Funk Island (4.1 h) than at Great Island (2.9 h). This resulted in lower feeding rates of chicks (0.17 feeds per h) and poorer condition of fledglings (2.9 g/mm) at Funk Island compared to those at Great Island (0.22 feeds per h; 3.9 g/mm). We hypothesize that provisioning efforts are constrained at Funk Island by (1) distant food resources and increased competitor density, resulting in longer foraging trip durations and (2) the time spent paired with mates at the colony, which may reflect a minimum time required to maintain breeding sites due to higher breeding densities at Funk Island compared to Great Island. Demographic consequences of this poor fledgling condition at Funk Island are unknown, but fledglings may sufficiently accelerate growth at sea due to their closer proximity to an important nursery area. If fledgling survival is compromised, however, the lower potential for growth at Funk Island will impact the entire Northwest Atlantic population of common murres.     

The diet of Common Guillemot Uria aalge chicks provides evidence of changing prey communities in the North Sea

Local differences in feeding conditions have been suggested as a cause of regional variation in seabird demography but multi‐colony comparisons of diet are rare. In UK waters the main fish eaten by seabirds during the breeding season belong to three families: Ammodytidae, Clupeidae and Gadidae. Climate change and fishing are affecting these fish stocks and so probably impact on predators such as seabirds. We used standardized observations of prey brought in for chicks to make the first integrated assessment of the diet of Common Guillemot Uria aalge chicks at a UK scale. Chick diet varied markedly among the 23 colonies sampled between 2006 and 2011. Sandeels (Ammodytidae), probably Lesser Sandeels Ammodytes marinus, were the commonest prey. Their contribution to the diet varied both latitudinally and among marine regions, with the proportion significantly higher for a given latitude on the west coast compared to the east. The non‐sandeel component of the diet showed latitudinal changes, with small clupeids, probably Sprats Sprattus sprattus, predominant at southern colonies whereas juvenile gadids were the main alternative to sandeels in the north. Comparison of our Guillemot chick diet with data collected 15–30 years earlier suggests that the proportion of sandeels in the diet has decreased at colonies bordering the North Sea. No significant change was apparent in Atlantic colonies but historical data were limited. The early years of our study coincided with a population explosion of Snake Pipefish Entelurus aequoreus in the Northeast Atlantic and North Sea. Pipefish were recorded in Guillemot chick diet at several northern and northwestern colonies in 2006 and 2007 but have been absent since 2009. Spatial and temporal variation in chick diet accorded broadly with patterns expected as a result of rising sea temperatures and impacts of fishing. Guillemot chick diet could potentially be a useful indicator of changes in the distribution and abundance of forage fish.     

Intercolony variations in movement patterns and foraging behaviors among herring gulls (Larus argentatus) breeding in the eastern Wadden Sea

Herring gulls (Larus argentatus) are opportunistic predators that prefer to forage in the intertidal zone, but an increasing degree of terrestrial foraging has recently been observed. We therefore aimed to analyze the factors influencing foraging behavior and diet composition in the German Wadden Sea. Gulls from three breeding colonies on islands at different distances from the mainland were equipped with GPS data loggers during the incubation seasons in 2012–2015. Logger data were analyzed for 37 individuals, including 1,115 foraging trips. Herring gulls breeding on the island furthest from the mainland had shorter trips (mean total distance = 12.3 km; mean maximum distance = 4.2 km) and preferred to feed on the tidal flats close to the colony, mainly feeding on common cockles (Cerastoderma edule) and shore crabs (Carcinus maenas). In contrast, herring gulls breeding close to the mainland carried out trips with a mean total distance of 26.7 km (mean maximum distance = 9.2 km). These gulls fed on the neobiotic razor clams (Ensis leei) in the intertidal zone, and a larger proportion of time was spent in distant terrestrial habitats on the mainland, feeding on earthworms. δ¹³C and δ¹⁵N values were higher at the colony furthest from the mainland and confirmed a geographical gradient in foraging strategy. Analyses of logger data, pellets, and stable isotopes revealed that herring gulls preferred to forage in intertidal habitats close to the breeding colony, but shifted to terrestrial habitats on the mainland as the tide rose and during the daytime. Reduced prey availability in the vicinity of the breeding colony might force herring gulls to switch to feed on razor clams in the intertidal zone or to use distant terrestrial habitats. Herring gulls may thus act as an indicator for the state of the intertidal system close to their breeding colony.     

Sex‐specific prey partitioning in breeding piscivorous birds examined via a novel,noninvasive approach

Piscivorous birds frequently display sex‐specific differences in their hunting and feeding behavior, which lead to diverging impacts on prey populations. Cormorants (Phalacrocoracidae), for example, were previously studied to examine dietary differences between the sexes and males were found to consume larger fish in coastal areas during autumn and winter. However, information on prey partitioning during breeding and generally on sex‐specific foraging in inland waters is missing. Here, we assess sex‐specific prey choice of Great Cormorants (Phalacrocorax carbo) during two subsequent breeding seasons in the Central European Alpine foreland, an area characterized by numerous stagnant and flowing waters in close proximity to each other. We developed a unique, noninvasive approach and applied it to regurgitated pellets: molecular cormorant sexing combined with molecular fish identification and fish‐length regression analysis performed on prey hard parts. Altogether, 364 pellets delivered information on both, bird sex, and consumed prey. The sexes differed significantly in their overall prey composition, even though Perca fluviatilis, Rutilus rutilus, and Coregonus spp. represented the main food source for both. Albeit prey composition did not indicate the use of different water bodies by the sexes, male diet was characterized by higher prey diversity within a pellet and the consumption of larger fish. The current findings show that female and male cormorants to some extent target the available prey spectrum at different levels. Finally, the comprehensive and noninvasive approach has great potential for application in studies of other piscivorous bird species.     

Cormorant (Phalacrocorax carbo [L]) populations and patterns of abundance at breeding and feeding sites in Northern Ireland,with particular reference to Lough Neagh

Breeding cormorants, Phalacrocorax carbo, in Northern Ireland remain seasonally dependent on a coastal environment where they can be censused with accuracy, but numbers in other habitats and at other times of the year are less certain. This study establishes the long term and regional patterns of abundance at breeding and feeding localities, which might in turn be related to diet.Birds were regularly observed and counted at a variety of feeding sites, and some aspects of their breeding success and fledgling diet evaluated at the largest N.Ireland breeding colony.Numbers of breeding birds increased dramatically over a period of eight years but recently show signs of declining. There is likely to be a dynamic relationship between populations of a tapeworm (Ligula intestinalis L.), the numbers of roach (Rutilus rutilus [L.]) and cormorant populations feeding at Lough Neagh, the largest lake in the British Isles. While roach are not a major component in the diet of fledglings, the large numbers of cormorants feeding at the perimeter of L. Neagh suggest that changes in roach populations will affect birds most acutely during or following the winter. We suggest that this might result in a reduction in the proportion of the population volunteering to breed in the subsequent season, but require further data. The longer term effects of one fish species on cormorant populations are unlikely to be critical, since these birds are highly opportunist.In other habitats cormorant numbers are either very stable (estuary) or variable (river) depending on the seasonal and annual availability of their prey. There is no evidence for a systematic seasonal shift in habitat, as suggested by other studies.     

Prey capture and selection throughout the breeding season in a deep‐diving generalist seabird,the thick‐billed murre

Generalist seabirds forage on a variety of prey items providing the opportunity to monitor diverse aquatic fauna simultaneously. For example, the coupling of prey consumption rates and movement patterns of generalist seabirds might be used to create three‐dimensional prey distribution maps (‘preyscapes’) for multiple prey species in the same region. However, the complex interaction between generalist seabird foraging behaviour and the various prey types clouds the interpretation of such preyscapes, and the mechanisms underlying prey selection need to be understood before such an application can be realized. Central place foraging theory provides a theoretical model for understanding such selectivity by predicting that larger prey items should be 1) selected farther from the colony and 2) for chick‐feeding compared with self‐feeding, but these predictions remain untested on most seabird species. Furthermore, rarely do we know how foraging features such as handling time, capture methods or choice of foraging location varies among prey types. We used three types of animal‐borne biologgers (camera loggers, GPS and depth‐loggers) to examine how a generalist Arctic seabird, the thick‐billed murre Uria lomvia, selects and captures their prey throughout the breeding season. Murres captured small prey at all phases of a dive, including while descending and ascending, but captured large fish mostly while ascending, with considerably longer handling times. Birds captured larger prey and dove deeper during chick‐rearing. As central place foraging theory predicted, birds travelling further also brought bigger prey items for their chick. The location of a dive (distance from colony and distance to shore) best explained which prey type was the most likely to get caught in a dive, and we created a preyscape surrounding our study colony. We discuss how these findings might aid the use of generalist seabirds as bioindicators.     

Feeding Habits and Interspecific Feeding Associations of Caranx Latus (Carangidae) in a Subtropical Reef

I studied the feeding behavior and diet of the carangid jack Caranx latus in a subtropical reef at Búzios Island on the southeastern Brazilian coast. Caranx latus foraged alone or in small groups of up to three individuals during daytime employing two main feeding styles: searching for prey while swimming in mid-water and following individuals of other fish species which disturbed the substrate while foraging among the rocks. The labrid wrasse Bodianus rufus was the main fish species followed by the jack. The jack feeds on crustaceans and fishes some of which are benthic rocky bottom dwellers and are caught during following. The behavioral flexibility of C. latus enables this fish to consume both crustaceans from the water column and benthic rocky dweller fishes. There is little dietary overlap between C. latus and the followed B. rufus. The interspecific feeding associations could be regarded as advantageous for C. latus allowing access to a broader range of prey categories and increasing the food intake through the consumption of large benthic fish prey     

Assessment of Grey Heron predation on fish communities: the case of the largest European colony

The fish predation rate by Grey Heron Ardea cinerea was studied during two breeding seasons (1987–88) in the largest European colony at the Lake of Grand-Lieu (Loire-Atlantique, France). The herons' diet was compared to the available fish population of its main feeding area, the marsh of Bourgneuf (16000 ha) which is composed of former salt pans and meadows drained by a dense network of shallow ditches. This study is the first attempt to assess the predation exerted by an ardeid colony on a fish community over such an extensive natural environment. It also provides the first data about the abundance and the structure of fish communities in shallow coastal dyked marshes. For this purpose, two different sampling methods were used according to the water's salinity. In fresh waters, electrofishing was used as the removal method, and density estimates were calculated with Carle & Strub estimator (1978). Fish were caught in randomly selected stations (sections of ditches enclosed by two 5 mm mesh nets). In brackish waters, pools and ditches were drained. The distribution of the herons at the feeding areas was determined by direct observations, by counting flights from the colony, and by radio-tracking. The diet was investigated by observing adult herons on the foraging areas, and by analyzing the prey regurgitations of the young at the nests. The global food consumption was assessed from Marion (1988), according to the birds' activity determined during 5 years of radio-tracking. Altogether, at least 39 species of fish were available in the herons' feeding area (during the reference period, 87–88) and the mean fish biomass was 270 kg per ha of open water, or 30 kg per ha of marsh (open water = 11.2% of the marsh area). The fish community was dominated by eel Anguilla anguilla (145 kg ha^–1, 50,8% of the total biomass), and catfish Ictalurus melas (40 kg ha^–1, 14%). Except for small and inaccessible species (living in the deepest parts of the marsh), heron diet was very similar to fish species composition of the community occurring in the marsh. The catfish was the species captured most frequently by the heron (45% of the mass), the eel was second with 28% of the mass. The catfish was probably over represented in the diet considering that they are caught in catfish-dumps created by professional fishermen at Grand-Lieu lake, in order to reduce the density of this undesirable species. Inversely, small species such as Gasterosteus aculeatus were not found in the diet whereas they are very numerous in the marsh. On average herons of Grand-Lieu colony catch 1.92 kg of fish per ha of marsh (6% of the fish standing crops in the marsh) during the breeding season, the main predation period.     

Feeding habits of Schizothorax oconnori Lloyd, 1908 in the Yarlung Zangbo River,Tibet

Seasonal, ontogenetic and sexual feeding habits of Schizothorax oconnori in the Yarlung Zangbo River were studied using percentages by number, weight and index of relative importance (IRI). The Schoener overlap index was applied to compare diets related to season, fish size and sex. The feeding intensity exhibited seasonal trends, with a minimum food intake in summer. Overall, 103 prey taxa belonging to seven prey categories (diatoms, green algae, cyanophytes, other algae, small invertebrates, macroinvertebrates and remains) were identified in 136 guts. Highest feeding diversity and evenness were recorded in summer, while lowest values of both indices occurred in autumn. S. oconnori fed almost exclusively on periphyton. Diatoms were the most important prey in terms of number, weight and IRI, followed by macroinvertebrates; other prey were also occasionally ingested. S. oconnori is a generalized and opportunistic feeder relying upon a wide trophic spectrum, but mainly feeds on diatoms as they are the most prevalent periphyton. There were high diet overlaps among seasons, fish sizes and between sexes. S. oconnori showed no dietary preference for any of the other prey available in the periphyton community.     

Diet and diving behaviour of European Storm Petrels Hydrobates pelagicus in the Mediterranean (ssp. melitensis)

Capsule Unlike Atlantic populations, which feed on krill, Mediterranean populations feed mainly on pelagic fish Gymnammodites cicerellus.

Aims To determine the diet and dive depth of the Mediterranean subspecies of European Storm Petrels Hydrobates pelagicus melitensis.

Methods Analysis of regurgitates of adults arriving at the colony for chick feeding and by determination of dives depth using the capillary tube method.

Results The main prey is Gymnammodites cicerellus, a pelagic fish. Storm Petrels dive for their prey and can reach up to 5 m in depth. They also make short foraging trips just outside the colony where they capture Opossum Shrimps Misydacea.

Conclusions European Storm Petrels in the Mediterranean exploit pelagic fish which are taken by diving. This contrasts with the Atlantic populations which feed mainly on krill. Mediterranean birds also feed on Opossum Shrimps Mysidacea during short foraging trips made at night just outside the colony. Differences in diet between long and short foraging trips may be because adults have to forage for both themselves and their chicks.     

Seasonal and annual differences in the foraging ecology of two gull species breeding in sympatry and their use of fishery discards

Niche segregation between similar species will result from an avoidance of competition but also from environmental variability, including nowadays anthropogenic activities. Gulls are among the seabirds with greater behavioural plasticity, being highly opportunistic and feeding on a wide range of prey, mostly from anthropogenic origin. Here, we analysed blood and feather stable isotopes combined with pellet analysis to investigate niche partitioning between Audouin's gull Larus audouinii and yellow‐legged gull Larus michahellis breeding in sympatry at Deserta Island, southern Portugal, during 2014 and 2015. During the breeding season there was considerable overlap in the adults’ diet, as their stable isotope values of blood and primary feather (P1) did not differ, and their pellets were comprised mainly by marine fish species. However, Audouin's gulls presented higher occurrences of pelagic fish, while yellow‐legged gulls fed more on demersal fish, insects, and refuse. SIAR mixing models also estimated a higher proportion of demersal fish in the diet of yellow‐legged gulls. We also found differences between the two gull species in chicks’ feathers, suggesting that Audouin's gull adults selected prey with lower carbon isotope values to feed their young. Secondary feather (S8) of Audouin's gull presented higher isotope values compared to yellow‐legged gulls, indicating different foraging areas (δ¹³C) and/ or trophic levels (δ¹⁵N) between the two species in the non‐breeding season. During both the all‐year and non‐breeding periods the yellow‐legged gull showed a broader isotopic niche width than Audouin's gull in 2013, and in 2014 the two gull species exhibited different isotopic niche spaces. Our study suggests that both gull species foraged in association with fisheries during the breeding season. In this sense, a discard ban implemented under the new European Union Common Fisheries Policy may lead to a food shortage, therefore future research should closely monitor the population dynamics of Audouin's and yellow‐legged gulls.     

Fish in the diet of the Antarctic shag Phalacrocorax bransfieldensis at Laurie Island, South Orkney Islands

The analysis of 29 stomach contents collected from a colony close to Pirie Peninsula, South Orkney Islands, from 2 January to 18 February 1995, showed that fish were by far the most important prey of the Antarctic shag Phalacrocorax bransfieldensis, followed by octopods, decapods and gammarids. The fish Notothenia coriiceps constituted the bulk of the diet; however, its importance decreased by the end of the study when Lepidonotothen nudifrons was the most important prey. The composition of the stomach contents varied throughout the breeding season according to changes in the chicks' energetic demands; as chicks grew older the adults increased the mass of the loads carried to the nests, preying on larger fish, a phenomenon also observed at the South Shetland Islands. Received: 3 July 1996 / Accepted: 9 March 1997     

Changing handling times during feeding and consequences for prey size selection of 0+ zooplanktivorous fish

The interrelationship of fish size, prey size and handling time within a 15-min feeding period was studied in three size groups of 0 + roach, Rutilus rutilus, and bleak, Alburnus alburnus. Four size classes of cladoceran prey were used to measure changes in feeding rate and handling time from initial rapid feeding to sustained feeding. Observed differences in increase of handling time between prey size classes led to a change in the prey profitability ranking of those size classes within the first 2 min of the experiments. A 1-min feeding period is interpreted as reflecting an intermediate motivational status between extreme hunger and satiation. The use of average handling times for this period revealed a substantial change in prey profitability estimates compared to previous studies which used handling times based on short-term (a few seconds up to 1 min) feeding. It is not the largest prey items a fish can handle and swallow that are most profitable, but prey of intermediate size. By this approach a closer fit between expectations derived from optimal foraging theory and empirical data on prey size selection of 0 + zooplanktivorous fish is qualitatively achieved. Optimal prey size was found to be close the mouth gape width in small fish of 15 mm standard length, decreasing to 50% of mouth gape width in fish of 40 mm standard length.     

Cost of morphological specialization: feeding performance of the two morphs in the trophically polymorphic cichlid fish,Cichlasoma citrinellum

Summary The feeding performance on soft and hard prey of two morphs of the trophically polymorphic Neotropical cichlid fish, Cichlasoma citrinellum, was investigated in the laboratory. The molariform morphs, specialized to feed on hard prey, are able to crack snail shells that are twice as hard as those cracked by the papilliform morphs. During ecological bottlenecks in food resources this ability should allow molariform morphs to exploit alternate, less preferred prey sources that are not available to papilliform morphs. Analysis of stomach contents revealed that molariform morphs feed significantly more often on hard snails than do papilliform morphs (Meyer 1989a). The performance advantage of the trophically specialized morphs when feeding on hard prey is countered by their less efficient performance on soft diets. The morphologically generalized papilliform morph feeds more efficiently on soft prey. The abundance of preferred soft prey, seasonal fluctuations in prey availability and the frequency of ecological bottlenecks may determine the relative abundance of these two morphs in natural populations in Nicaraguan lakes.