Feeding behavior of Geoffroy's marmoset (Callithrix geoffroyi) in an Atlantic forest fragment of South-eastern Brazil

We report on the diet and feeding behaviour of a group of Geoffroy's marmosets (Callithrix geoffroyi) in an Atlantic forest fragment in south-eastern Brazil, studied during the period February 1993 to Januaray 1994. Major food categories were gums (68.6%) fruits (15%), and small animal prey (invertebrates 14.6% and vertebrates 0.8%). Dietary changes were observed between the wet and dry seasons, although they were not statistically significant. Insects and gums were consumed throughout the year but fruits were more prevalent in the diet in the wet season. Plant species exploited for their gums includedInga stipularis, followed byAcacia paniculata, Paulinia carpopodia, andBauhinia angulosa. Chemical analysis of the gum of the four species most used all presented high values for carbohydrates and crude proteins. Fruits of an unidentified species of Myrtaceae (sp. 2) were consumed the most. Animal prey consisted mainly of insects, arachnids, snails, and in three cases, lizards and frogs, with orthopterans being the most prevalent insect prey. This study demonstrates thatC. geoffroyi efficiently exploits resources typically available in secondary and disturbed forest habitat. The main threat to the species is forest destruction, degradation, and fragmentation, but the management of small forest fragments may be an effective corservation strategy.     

Seasonal food habits of corsac and red foxes in Mongolia and the potential for competition

Competition often occurs between sympatric species that exploit similar ecological niches. Among canids, competition may be reduced by partitioning resources such as food, time, and habitat, but the mechanisms of coexistence remain poorly understood, particularly among fox species. We described the food habits of two foxes that live sympatrically across northern and central Asia, the corsac fox (Vulpes corsac) and red fox (V. vulpes), by analyzing scats collected during a field study in Mongolia. We analyzed 829 corsac and 995 red fox scats collected from April 2005 to August 2007 and tested the extent to which food partitioning occurred. The diets of both species consisted mainly of insects followed by rodents, but also included birds, reptiles, large mammal remains (carrion), plant material (including fruits and seeds), and garbage. Despite high overlap in the proportion of food items consumed, differences existed between species in overall diet with corsacs more frequently consuming beetles, but proportionally fewer crickets and large mammal remains than red foxes. We detected interspecific differences during the pup rearing and dispersal seasons, when prey was abundant, but not during the breeding season, when prey was scarce and diet overlap highest. Each species’ diet also differed seasonally and exhibited moderate overall breadth. Corsacs consumed proportionally more beetles and rodents during pup rearing and crickets during dispersal relative to other seasons, whereas red foxes consumed proportionally more crickets during pup rearing and dispersal and more rodents and large mammals during pup rearing and breeding relative to other seasons. Our results suggest that partitioning of food resources during most of the year facilitates coexistence, and that the potential for competition is highest during winter months.     

Dietary niche differentiation of five sympatric species of Platycephalidae

The dietary composition and partitioning of food resources between five sympatric species of Platycephalidae inhabiting the coastal waters of New South Wales, Australia was investigated. Samples were collected monthly between March and November 2007 onboard commercial ocean prawn trawlers based in the ports of Yamba and Newcastle. Monthly percentage weight contribution of 12 prey categories was analysed to determine if diet was influenced by the variables: species, location, depth, size and maturity. Of the 959 stomachs from the five species examined, 28–54% contained prey. All Platycephalid species primarily consumed teleosts, however the diversity of prey and the proportion each prey type contributed to the overall diet varied substantially between species. Platycephalus caeruleopunctatus, P. longispinis, P. richardsoni and Ambiserrula jugosa were generalist carnivores and consumed prey from a wide variety of phyla including teleosts, crustaceans, polychaetes, molluscs and echinoderms. In contrast, Ratabulus diversidens were primarily piscivorous. Partitioning of prey resources between species was more evident in waters at Yamba than at Newcastle. Differences in diet between locations were considered a result of differential prey exploitation rather than shifts in the suite of prey consumed. Dietary composition was observed to be influenced by size, maturity status and depth however these differences were not observed for all species.     

Seasonal pattern in food gathering of the weaver ant Oecophylla longinoda (Hymenoptera: Formicidae) in mango orchards in Benin

Prey capture and food scavenging activities of Oecophylla longinoda were monitored through regular weekly samplings during two consecutive years (2009–2010) in a large mango orchard of the Borgou Department of Benin, West Africa, a main mango production area located in the Sudan agro-ecological zone. In both years, interspecific competition with other ants occurred mainly during the dry season (January to March) resulting in increased captures of Formicinae, Myrmicinae and Ponerinae. More prey was caught during the rainy season (end April to end October) than during the rest of the year, with Diptera and Coleoptera prey peaking in May and June, respectively, along with the mango season. As insect prey quickly decreased during November to December weaver ants increasingly collected seeds and plant debris. A total of 241 species of insects were captured including 61 species (25.3%) associated with mango and a few with cashew, among which 48 mango pest species (78.7% of species associated with mango tree). Only five species (2.1%) of beneficial insects were captured. It is concluded that the presence of O. longinoda colonies is beneficial to perennial tree cropping systems such as mango and cashew.     

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

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

Ontogenetic variations in the type and size of prey consumed by juvenile coho,Oncorhynchus kisutch,and chinook,O. tshawytscha,salmon

Synopsis Stomach contents of juvenile coho,Oncorhynchus kisutch, and chinook,O. tshawytscha, salmon collected in purse seines off the coast of Washington and Oregon were examined for variations related to predator size. There was a general trend toward increasing consumption of fish with increasing body size, due mainly to the increase in northern anchovy biomass consumed by the larger salmon. Most of the major prey taxa showed significant differences among the size classes examined for both salmon species. There was a direct relationship between predator and prey size for both coho and chinook, but considerable variation was found in prey length consumed within each size class. Prey width did not provide as good a fit as prey length for either species. In general, coho consumed larger fish prey in relation to their body length than chinook but there were substantial differences by month or year of collection.     

Feeding ecology of a vertebrate assemblage inhabiting a stream of NW Spain (Riobo; Ulla basin)

The food resource use of a stream in NW Spain by fish (Salmo trutta L. and Anguilla anguilla L.), birds (Cinclus cinclus L. and Motacilla cinerea L.) and mammals (Galemys pyrenaicus G. and Neomys anomalus C.) was studied. Data on seasonal diets and stream benthos prey were used to determine prey selection patterns.Caddisfly larvae are the main resource for Cinclus and Galemys, but these predators also consumed other benthic prey. Salmo fed on a wide range of benthic invertebrates, emergent pupae and terrestrial prey, whereas Anguilla consumed primarily benthic invertebrates, especially Lumbricids. Neomys fed mainly on terrestrial prey (Gasteropods and Lumbricids), but also consumed aquatic prey. Motacilla captured aquatic insects both in larval and aerial stages, as well as terrestrial prey.Both prey availability and selection led to seasonal differences in the use of food resources. All species showed a marked prey selection of aquatic taxa. Prey size plays an important role in this selection, most species consuming the largest of available prey sizes. In spite of the fact that all species feed upon freshwater invertebrates, substantial resource partitioning was observed in all seasons. This partitioning may be attributable to morpholological and physiological differences. Nevertheless, Anguilla and Galemys, two quite different animals, did feed on the same prey much of the time.     

Evidence of Large Change in Unionid Mussel Abundance from Selective Muskrat Predation,as Inferred by Shell Remains Left on Shore

Three species of unionid mussels (Pyganodon grandis, Lampsilis siliquoidea, and Toxolasma parvus) were preyed on by muskrats (Ondatra zibethicus) that left shell remains at feeding sites (middens) along the shore of Lake LaSalle (West) in western New York State, USA. Mussels became established within five years of the construction of this small artificial lake, and individuals of all three species have grown rapidly to large size. The abundance of prey in middens in 1988–1990 and a marked decline in subsequently deposited shells indicate how dramatically muskrats had exploited this food source. They consumed > 2700 individuals of P. grandis (by far the most common prey item) through 1990, yet only 20 newly deposited shells were found in 1993. At a midden where prey depletion was directly catalogued during 1988, larger specimens of P. grandis were consumed first, suggesting size selection. L. siliquoidea has been least affected by predation, and was the mussel most often found alive in the littoral zone. No live T. parvus were ever collected, and this species is known only from middens. Complete collections of shell remains from the shoreline through 1999 suggest that the mussel population may be recovering slowly. Muskrat predation may be more important in influencing mussel population structure than previously realized, and the effects can be dramatic spatially and temporally.     

Diet of the nominate Southern Grey Shrike Lanius meridionalis meridionalis in the north of its range (Mediterranean France)

Capsule In this region the diet is mainly cold-blooded prey, mostly insects such as beetles.

Aims To describe the diet of this newly separated, poorly documented and endangered species.

Method Diet was inferred from pellet analysis, collected during a single breeding and winter period in the steppe of the Crau.

Results A total of 5409 prey were identified from 257 pellets. Vertebrates were seldom taken, except by adults (small passerines) during the fledgling period. High seasonal differences were found. Hymenoptera were largely consumed in autumn, Arachnida in autumn and winter, Orthoptera in summer and autumn and Lepidoptera larvae in winter and spring and by fledglings. Nevertheless, Coleoptera were ingested in large proportions all year round. Carabidae were the main prey in winter and Melolonthidae were especially important for adults during the nestling period, as were Cetoniidae for the fledglings.

Conclusion Small mammals and small birds were less exploited in France and Spain (L. m. meridionalis) than in Israel (L. m. elegans or L. m. aucheri), whereas the opposite might be expected, following a north–south climatic gradient. Thus, the nominate subspecies L. m. meridionalis differed in diet from L. m. elegans or L. m. aucheri.     

Diet and foraging effort of Adélie penguins in relation to pack-ice conditions in the southern Ross Sea

We investigated the diet and aspects of foraging effort among Adélie penguins (Pygoscelis adeliae) breeding at three colonies on Ross Island, in the southwestern Ross Sea – Capes Royds, Bird and Crozier – during the chick-provisioning period of three austral summers, 1994–1995, 1995–1996 and 1996–1997. During the study period, pack-ice cover differed in waters offshore of these colonies, by colony, seasons and year. Diet differed among colonies only slightly. The fish Pleuragramma antarcticum was the most important prey, especially during years or periods within years when little pack ice was present. With respect to krill, which composed the remainder of diet, juvenile Euphausia crystallorophias were consumed predominantly in a year of heavy pack-ice cover; more adult krill were consumed in 2 years when pack ice was sparse. Foraging trip duration differed by colony, season and year and was related directly to distance from the colony to the nearest pack ice. The amount of food brought to chicks increased as trip duration increased, to a point (2 days), but then decreased as duration increased further (up to 4 days). On the basis of data on mass of parents and of meal sizes to chicks, it appeared that on the longest trips more of the food gathered by parents was used for self maintenance; on the longest trips, parents lost body mass. Successful foraging during chick rearing, the period when adult foraging is most intense, appears to depend on the proximity of pack ice to nesting colonies for this penguin species. Received: 1 October 1997 / Accepted: 25 April 1998     

Dietary Dependence of Predatory Arthropods on Volant Aquatic Insects in Tropical Stream Riparia

The dietary dependence on volant aquatic insects of eight species of predatory arthropods from three different orders was determined by stable isotope analyses in combination with three‐source, two‐isotope (C and N) Bayesian mixing models. The predators were collected from riparian zones along three streams in tropical Hong Kong during both the wet and dry seasons. Dietary importance of aquatic insects varied according to predator hunting modes, and showed a consistent pattern across all sites during the wet season. The web‐building tetragnathid spider (Orsinome diporusa) had the greatest reliance (~40–55%) on this water‐to‐land subsidy, followed by two species of damselflies (40–50%), three cursorial spiders (Lycosidae, Pisauridae, and Sparassidae: 32–51%) and two neustic gerrids (17–36%). Such reliance also varied according to the microhabitat preferences of different cursorial spiders. Four species of predators (gerrids and cursorial spiders) that were active year‐round showed generally consistent reliance on aquatic insects between seasons, which probably reflected the observed lack of seasonal variability in the relative proportions of aquatic and terrestrial prey. There was a marked overlap in isotopic signatures of aquatic and terrestrial prey at all sites which, combined with the absence of data on the extent to which isotopic fractionations may vary among individual species of prey and predators, contributes some uncertainty to the estimates of dietary compositions derived by mixing models. The findings of the present study are thus likely to be indicative rather than definitive.     

Amphipod (Gammarus minus) responses to predators and predator impact on amphipod density

Recent theoretical work suggests that predator impact on local prey density will be the result of interactions between prey emigration responses to predators and predator consumption of prey. Whether prey increase or decrease their movement rates in response to predators will greatly influence the impact that predators have on prey density. In stream systems the type of predator, benthic versus water-column, is expected to influence whether prey increase or decrease their movement rates. Experiments were conducted to examine the response of amphipods (Gammarus minus) to benthic and water-column predators and to examine the interplay between amphipod response to predators and predator consumption of prey in determining prey density. Amphipods did not respond to nor were they consumed by the benthic predator. Thus, this predator had no impact on amphipod density. In contrast, amphipods did respond to two species of water-column predators (the predatory fish bluegills, Lepomis macrochirus, and striped shiners, Luxilus chrysocephalus) by decreasing their activity rates. This response led to similar positive effects on amphipod density at night by both species of predatory fish. However, striped shiners did not consume many amphipods, suggesting their impact on the whole amphipod “population” was zero. In contrast, bluegills consumed a significant number of amphipods, and thus had a negative impact on the amphipod “population”. These results lend support to theoretical work which suggests that prey behavioral responses to predators can mask the true impact that predators have on prey populations when experiments are conducted at small scales. Received: 21 March 1997 / Accepted: 15 December 1997     

Feeding Habits of Two Large Insects from a Desert Stream: Abedus herberti (Hemiptera: Belostomatidae) and Thermonectus marmoratus (Coleoptera: Dytiscidae)

Feeding preference experiments were conducted to determine the feeding habits of Abedus herberti (Heteroptera, Belostomatidae) and Thermonectus marmoratus (Coleoptera, Dytiscidae), two large insects in Sycamore Creek, an intermittent Sonoran desert stream, Arizona, U.S.A. Numbers of live versus dead prey consumed were tested between and across three prey sizes. Five prey species were offered simultaneously (5 live and 5 dead specimens) in each size class. We found that A. herberti preferred live prey of small and medium size, but it chose mainly dead prey in the large size class. These results fitted the model of size-selective predation (Zaret, 1980). Size dependent predators selected prey of increased size, according to their visibility, but only up to where difficulty in handling and probability of escape affect successful consumption. Snails were the most preferred prey of A. herberti. By contrast, T. marmoratus consumed only dead prey of all sizes, but it preferred soft organisms with thin cuticle, such as immature larvae of some mayflies, beetles, dragonflies or fishes.     

Shifts of food habit and jaw position of white-spotted charr after damming

Stomach fullness of white-spotted charr, Salvelinus leucomaenis did not differ significantly between above-dam (allopatric with masu salmon) and below-dam (sympatric with masu salmon) areas, but stomach contents did. Juveniles (1+) below the dam consumed relatively more aquatic insects (benthic prey), and those above the dam consumed relatively more terrestrial insects (drifting prey), but there was no difference in the diet of >2+ fish between areas. The mouths of juveniles above the dam were more terminal than those of fish below the dam. The adaptive significance of these differences is discussed in relation to differences in fish assemblages. 1999 The Fisheries Society of the British Isles     

Aquatic prey switching and urban foraging by the White Ibis Eudocimus albus are determined by wetland hydrological conditions

Prey availability is known to limit reproduction of some species of nesting birds, but identifying the primary prey types of a species with a flexible diet can be challenging. For the White Ibis Eudocimus albus, a tactile feeding, medium‐sized wading bird, nestling prey composition is suggested to depend on landscape water depths/availability of foraging habitat at the time of nesting and on historical drying events affecting prey production. We collected and compared inter‐ and intra‐annual diet variation of White Ibis chicks reared in the Everglades over two years that were independently identified as being relatively good (2006) and poor (2007) nesting seasons. We collected 127 nestling boluses and analysed the temporal variation in biomass of eight functional prey groups using multivariate techniques. The boluses from 2006 in the central Everglades were dominated by fish, but in 2007, after fish had been reduced by the previous year of drying, the boluses from the same region were more variable and dominated by garbage (i.e. scavenging). Analysis of five different collections taken from a different colony in the northern Everglades indicated that boluses were characterized by crayfish and had fewer fish or less garbage when landscape water depths were relatively higher and more preferred habitat was available. At lower landscape water depths in 2007 the bolus composition shifted away from crayfish towards small fish and urban food (terrestrial insects and garbage). Our results support the suggestion of depth‐dependent diets; prey composition depends on the current landscape water levels around the colonies, and also suggests that previous drying events can lead to increased reliance on alternative food sources. White Ibis partially compensated for unavailable aquatic prey with alternative urban foods, but their nesting success appears to have suffered.     

Interannual variation in the diet of non-breeding male Antarctic fur seals, <Emphasis Type="Italic">Arctocephalus gazella</Emphasis>, at Isla 25 de Mayo/King George Island

The diet of non-breeding male Antarctic fur seals, Arctocephalus gazella, was investigated at Stranger Point, King George Island, through the analysis of scats during three consecutive summer seasons (1996, 1997, 1998). Overall, fish and krill were the most frequent prey occurring, respectively, in an average of 82.9% and 78.8% of samples (n = 131), followed by penguins (22.8%) and cephalopods (17.8%). Myctophids constituted almost 90% of the fish predated, with Electrona antarctica and Gymnoscopelus nicholsi being the most abundant and frequent species consumed. All fish taxa identified were krill feeding species suggesting that seals foraged mainly on a krill and a fish community associated with krill aggregations. However, a seasonal change was observed in the relative proportions of the different prey taxa, with a progressive decrease with time in the occurrence of krill and a concomitant increase of fish, penguins and squid. Possible influence of the strong 1997/98 ENSO event is discussed.     

Feeding ecology of the hibernating primate Cheirogaleus medius: how does it get so fat?

The fat-tailed dwarf lemur (Cheirogaleus medius; Cheirogaleidae) lives in the tropical forests of Madagascar. It is exceptional among primates, because this small nocturnal lemur hibernates during the dry season from the end of April until October. This field study was carried out to assess seasonal changes in body mass and behavioral traits (locomotor activity, feeding ecology) of C. medius that enable this species to accumulate large quantities of fat within weeks. Feeding observations of 36 radio-collared individuals were combined with a mark-recapture study in western Madagascar during the rainy seasons from 1995 to 1998. Pulp of fruit species not consumed, as well as fruit species eaten by C. medius was analyzed for lipid and sugar contents. Body mass changes of recaptured individuals revealed that C. medius may double its body mass within a few weeks. Most of the increase in body mass took place between March and April after infants had been weaned and directly before the onset of hibernation. Locomotor activity was significantly reduced during the period of body mass gain. During the entire study, fruits, nectar, and flowers of 34 different plant species comprised about 80% of the diet, supplemented by a seasonally varying intake of animal prey. Lipid contents of consumed fruits were comparatively low throughout the study period. No difference in lipid content was detectable between fruit species consumed and not consumed by C. medius. During prehibernation fattening, C. medius consumed mostly fruit species with a high sugar content, significantly higher than that of fruit species that were not consumed. The results suggest that a reduction in locomotor activity as well as a preference for fruit species with a high sugar content enable C. medius to deposit large quantities of fat in a few weeks. Received: 15 March 1999 / Accepted: 30 June 1999     

Nutritional composition of actual and potential insect prey for the Kasekela chimpanzees of Gombe National Park,Tanzania

Humans, all great ape species, and some lesser apes consume insects. Insects can provide comparable nutritional yields to meat on a gram‐for‐gram basis and may serve as an important source of energy, fat, protein, minerals, and vitamins for hominoids. Although potential insect prey are abundant in ape habitats, patterns of insectivory are not consistent across species or populations. Efforts to understand these patterns are complicated by a lack of nutritional data. We collected samples of insects consumed by the Kasekela chimpanzee community of Gombe National Park, Tanzania, as well as of some insects found within the community range and ignored by these chimpanzees but known to be preyed upon by Pan elsewhere. We determined the gross energy (GE), estimated metabolizable energy (ME), fat, protein, fiber, and ash content of these samples following standard methodologies. We use these data to test the hypothesis that Kasekela chimpanzees choose insect prey (at least in part) based on energy and/or macronutrient content. On a fresh‐weight, per‐gram basis, the insect prey consumed by Kasekela chimpanzees had significantly higher fat and lower ash content than other assayed insects, and on a fresh‐weight, per‐foraging‐unit (“per‐insect,” “per‐dip,” or “per‐nest”) basis were significantly higher in GE, fat, and protein. On a per‐gram basis, the assayed insects were generally comparable in energy and macronutrients to wild vertebrate meat. We conclude that Kasekela chimpanzees do favor insects that are high in energy, fat, and protein, and that the potential macronutrient yields from some forms of insectivory are not trivial. © 2012 Wiley Periodicals, Inc.     

Prey choice by marten during a decline in prey abundance

Summary We examined variation in diet choice by marten (Martes americana) among seasons and between sexes and ages from 1980–1985. During this period prey populations crashed simultaneously, except for ruffed grouse (Bonasa umbellus) which was common at the beginning and end of the study, and masked shrews (Sorex cinereus) which were abundant in 1983. Marten were catholic in selection of prey and made use of most available mammalian prey, ruffed grouse, passerine birds, berries, and insects. Diet niche was widest during the latter three years when prey was scare, particularly in late winter. Diet niche breadth was negatively correlated with abundance of all common prey species. Proportion of small prey species in the diet was correlated with absolute abundance of those species, but proportion of some large prey was related to their relative abundance. Diet choice varied among years and among seasons. Berries and insects were common in summer diets while large prey, particularly varying hare (Lepus americanus), were more frequent in winter diet than in summer diet. We found little evidence that any small mammal species was a preferred prey. Sexual size dimorphism between the sexes did not affect prey choice, nor did age. Reduced foraging effort in winter resulted in a wider diet niche only when prey was scarce. The only prediction of optimal foraging models fully supported by our data was a wider diet niche with reduced prey abundance. However, among the three most profitable prey species choice was dependent on the absolute abundance of the most profitable type (varying hare). We suggest that marten primarily forage for large prey but employ a strategy which results in encounters with small prey as well. These small prey are eaten as they provide energy at minimal cost, between captures of large prey.     

Intra- and interannual variation in the diet of the Magellanic penguin (Spheniscus magellanicus) at Martillo Island,Beagle Channel

Knowledge of seabirds’ diet at each breeding site and its temporal variation is key to understanding and evaluating how changes in marine resources affect each seabird population. In this study, we determined the diet of Magellanic penguins (MP, Spheniscus magellanicus) at Martillo Island, accounting for sex, breeding stage and year. We analyzed a total of 144 stomach contents during three consecutive breeding seasons (2006–2007, 2007–2008 and 2008–2009) and stages (incubation, early and late chick-rearing). MP fed mainly on fuegian sprat (Sprattus fuegensis), which represented 75 % of the biomass consumed by birds during the entire study. The next important prey was squat lobster (Munida gregaria), followed by Patagonian squid (Loligo gahi). Both sexes consumed similar prey items. We observed variation in diet relative composition among breeding years and stages. Fuegian sprat consumption decreased throughout the years whereas squat lobster increased. Penguins consumed a higher proportion of squat lobster and Patagonian squid during the incubation stage than in the chick-rearing stages, whereas fuegian sprat was almost the only prey item consumed during the late chick-rearing stage. MPs show certain flexibility in the use of resources probably as a response to changes in prey populations. Variability in the diet among different reproductive stages could be related to changes in the distribution and abundance of their main prey near the colony during the breeding season together with changes in the energy requirements of seabirds.