Foraging movements and habitat niche of two closely related seabirds breeding in sympatry

As central-place foragers, pelagic seabirds are constrained by spatiotemporal heterogeneity to find productive marine areas and compete for prey. We analysed 97 foraging trips to study the movement and oceanographic characteristics of foraging habitats of two different—yet closely related—species of shearwaters (Scopoli’s shearwater Calonectris diomedea and Cory’s shearwater C. borealis) breeding in sympatry in the Mediterranean. We combined various methodological approaches (GPS-tracking, species distribution modelling and stable isotope analysis) to explore the foraging strategies of these two species. Isotopic results suggested that trophic habits of both shearwater species were similar, mainly based on pelagic fish consumption. Foraging areas of both species were characterized by shallow waters near the colony. Both shearwater species exploited persistent productive marine areas. The foraging areas of the two species broadly overlapped during the incubation period, but during chick-rearing period, Scopoli’s shearwaters apparently foraged in different areas than Cory’s shearwaters.     

Oceanographic characteristics of areas used by Cory’s shearwaters during short and long foraging trips in the North Atlantic

Many breeding seabird species are central-place foragers and constrained to find productive prey patches within their foraging ranges. We assessed how different populations of a pelagic seabird species, the Cory’s shearwater Calonectris diomedea, breeding in oceanic and neritic conditions, cope with these constraints in the North Atlantic, during both incubation and chick-rearing periods. We analysed 237 foraging trips to study the movements and oceanographic characteristics of foraging habitats of seven different populations of Cory’s shearwaters. Generally, oceanic populations exhibited higher foraging effort, by travelling more time and to more distant areas, and larger home ranges and feeding areas, than the neritic population (i.e. breeding on an island within the Portuguese continental platform). On their short trips (i.e. ≤4 days), birds from the different populations fed mostly in shallower waters around the colony. During long trips (i.e. ≥5 days), feeding areas of both oceanic and neritic populations were characterized by high concentration values of chlorophyll-a, low sea-surface temperature and shallower habitats, with oceanic populations of the Azores exploiting areas north of the islands over known seamounts and frontal regions. Birds from other oceanic population (Selvagens) also exploited the African continental shelf system on their long trips. The home ranges of the different populations overlapped widely, but there was a general spatial segregation in terms of the core feeding areas at the population level. Core feeding areas and areas of foraging overlap between different populations should be important to inform conservation management measures, such as the definition of Marine Important Bird Areas for seabirds over the North Atlantic.     

Parent–offspring dietary segregation of Cory’s shearwaters breeding in contrasting environments

In pelagic seabirds, who often explore distant food resources, information is usually scarce on the level of trophic segregation between parents and their offspring. To investigate this issue, we used GPS tracking, stable isotopes and dietary information of Cory’s shearwaters Calonectris diomedea breeding in contrasting environments. Foraging trips at Selvagem Grande (an oceanic island) mainly targeted the distant African coast, while at Berlenga island (located on the continental shelf), shearwaters foraged mainly over nearby shelf waters. The degree of isotopic segregation between adults and chicks, based on δ¹³C, differed markedly between the two sites, indicating that adult birds at Selvagem fed their chicks with a mixture of shelf and offshore pelagic prey but assimilated more prey captured on coastal shelf waters. Isotopic differences between age classes at Berlenga were much smaller and may have resulted from limited dietary segregation or from age-related metabolic differences. The diet of shearwaters was also very different between the two colonies, with offshore pelagic prey only being detected at Selvagem Grande. Our findings suggest that spatial foraging constraints influence resource partitioning between pelagic seabirds and their offspring and can lead to a parent–offspring dietary segregation.     

Behavioural inertia places a top marine predator at risk from environmental change in the Benguela upwelling system

In variable environments, organisms are bound to track environmental changes if they are to survive. Most marine mammals and seabirds are colonial, central-place foragers with long-term breeding-site fidelity. When confronted with environmental change, such species are potentially constrained in their ability to respond to these changes. For example, if environmental conditions deteriorate within their limited foraging range, long-lived species favour adult survival and abandon their current breeding effort, which ultimately influences population dynamics. Should poor conditions persist over several seasons, breeding-site fidelity may force animals to continue breeding in low-quality habitats instead of emigrating towards more profitable grounds. We assessed the behavioural response of a site-faithful central-place forager, the Cape gannet Morus capensis, endemic to the Benguela upwelling system, to a rapid shift in the distribution and abundance of its preferred prey, small pelagic shoaling fish. We studied the distribution and the abundance of prey species, and the diet, foraging distribution, foraging effort, energy requirements, and breeding success of gannets at Malgas Island (South Africa) over four consecutive breeding seasons. Facing a rapid depletion of preferred food within their foraging range, Cape gannets initially increased their foraging effort in search of their natural prey. However, as pelagic fish became progressively scarcer, breeding birds resorted to scavenging readily available discards from a nearby demersal fishery. Their chicks cannot survive on such a diet, and during our 4-year study, numbers of breeding birds at the colony decreased by 40% and breeding success of the remaining birds was very low. Such behavioural inflexibility caused numbers of Cape gannets breeding in Namibia to crash by 95% following over-fishing of pelagic fish in the 1970s. In the context of rapid environmental changes, breeding-site fidelity of long-lived species may increase the risk of local or even global extinction, rendering these species particularly vulnerable to global change.     

Seabird use of discards from a nearshore shrimp fishery in the South Atlantic Bight,USA

Shrimp trawling is common throughout the southeastern and Gulf of Mexico coasts of the USA and is the primary contributor to fisheries discards in these regions. Tens of thousands of nearshore seabirds nest near shrimp trawling grounds in the USA, but to date, there has been no assessment of the relationship between seabirds and shrimp trawlers. We examined the taxonomic composition of bycatch, rate at which seabirds scavenged bycatch, and energy density of discarded bycatch in a nearshore commercial shrimp fishery. Bycatch was primarily comprised of demersal fish that are not typically accessible to the plunge-diving and surface-feeding seabirds that occur in the area. Hence, seabird diets in the region appear to be broadened taxonomically by the availability of discards. Results from discard experiments indicated that 70% of the nearly 5,500 items discarded by hand were scavenged by seabirds and that the fate of a discarded item was most strongly predicted by its taxonomic order. Laughing gulls scavenged the greatest proportion of discards, although brown pelicans were the only species to scavenge more discards than predicted based upon their abundance. Because this is the first such study in the region, it is difficult to ascertain the extent or intensity of the impact that discards have on nearshore seabirds. Nonetheless, our results suggest that it will be difficult for managers to clearly understand fluctuations in local seabird population dynamics without first understanding the extent to which these species rely upon discards. This may be especially problematic in situations where seabird populations are recovering following natural or anthropogenic stressors.     

Diets of the demersal fishes on the shelf off Iwate,northern Japan

Diets of demersal fishes were determined on the shelf (ca. 130 m deep) off Iwate, Japan. Samples were taken in three different types of habitat, an artificial reef (AR) site, a natural reef (NR) site, and sandymud bottom (SB) site, from May 1987 to September 1991, mostly every two months. A total of 67 prey items were recognized in the stomachs of 45 predator fish species. The most important preys were the pelagic fishes Sardinops melanostictus and Engraulis japonicus, which comprised 37% wet wt of the overall stomach contents. The percentage of pelagic fishes was highest at AR site, where fish density was highest. The dominant ten species could be divided into five feeding types. The pelagic fish feeders Physiculus maximowiczi and Gadus macrocephalus fed mainly on S. melanostictus. The dietary breadth of P. maximowiczi was wide, while that of Gadus macrocephalus was narrow. The pelagic crustacean feeder Theragra chalcogramma mostly consumed Themisto japonica and euphausiids and showed the least dietary overlap with other fishes. Benthic fish feeders were Hemitripterus villosus and Liparis tanakai. The benthic crustacean feeders Alcichthys alcicornis and Hexagrammos otakii consumed benthic crustaceans as well as pelagic and benthic fishes and showed the largest dietary breadth. The benthic invertebrate feeders Gymnocanthus intermedius, Dexistes rikuzenius and Tanakius kitaharai fed mainly on polychaetes and benthic crustaceans. But Gymnocanthus intermedius consumed a significant proportion of pelagic fishes. Ontogenetic dietary shift was recognized for these fishes. Pelagic fishes were consumed more intensively by larger individuals, especially true of A. alcicornis, Theragra chalcogramma and Gadus macrocephalus. Predominancy of the two most adundant species, P. maximowiczi and A. alcicornis, may be supported by their wide dietary breadth and the significant proportion of pelagic fish in their diets. Interspecific dietary overlap was low in most cases suggesting that food resources were well partitioned, although some high overlap was observed among the pelagic fish feeders, A. alcicornis, and Gymnocanthus intermedius, and among the benthic invertebrate feeders. Interspecific competition seemed more likely in the benthic invertebrate feeders than in the pelagic fish feeders partly because of superabundance of the pelagic prey S. melanostictus.     

Diet of the Humboldt penguin (<Emphasis Type="Italic">Spheniscus humboldti</Emphasis>) in northern and southern Chile

The diet of the Humboldt penguin (Spheniscus humboldti) was examined and compared in two colonies in Chile. Field work was conducted on Pan de Azúcar Island in northern Chile in the breeding season 1998/1999 and on the Puñihuil Islands in southern Chile over two successive breeding seasons during 1997/1998 and 1998/1999. Penguin diet was studied by stomach-pumping birds and analysed by species composition, size and mass of prey. Fish were the dominant prey item at both sites, the contribution of cephalopods and crustaceans varying between sites. The fish prey consisted predominantly of school fish, but there were clear latitudinal differences in fish prey taken. Penguins in the northern colony consumed primarily garfish (Scomberesox saurus), while birds at the southern colony of Puñihuil fed primarily on anchovy (Engraulis ringens), Araucanian herring (Strangomera bentincki) and silverside (Odontesthes regia). The results showed significant differences in terms of numbers of fish taken between the two breeding seasons at Puñihuil. In 1997/1998 penguins consumed almost exclusively anchovy, while they fed primarily on silversides in the successive year. Almost all prey, except stomatopods, were characterised as being pelagic species that occur in relatively inshore water, consistent with the foraging behaviour of Humboldt penguins. The dependence of Humboldt penguins on commercially exploited, schooling prey species makes the species particularly susceptible to changes in prey stocks, due to non-sustainable fisheries management.Communicated by O. Kinne, Oldendorf/Luhe     

Inter- and intra-year variation in foraging areas of breeding kittiwakes (Rissa tridactyla)

While seabird conservation efforts have largely focused on protection from threats at the colony (e.g. reducing disturbance and predation), attention is increasingly being given to implementing protection measures for foraging areas at sea. For this to be effective, important foraging areas must be identified. Although numerous studies have examined seabird foraging behaviour, information is still lacking on the variability in area utilisation within and among breeding seasons. GPS devices were attached to adult black-legged kittiwakes breeding at an expanding North Sea colony (55°20′N, 1°32′W) during both incubation and chick-rearing in 2012 and during chick-rearing in 2011, to determine whether foraging areas remained consistent and to identify the oceanographic characteristics of areas used for foraging. The type and size of prey items consumed at different stages of the breeding cycle was also examined. During incubation (April–May 2012), kittiwakes foraged substantially further from the colony and fed on larger sandeels than when feeding chicks, and there was significant inter-annual variation in foraging areas used during the chick-rearing period (June–July 2011 and 2012). Foraging areas were characterised by cooler sea surface temperatures and areas of high chlorophyll a concentration, although association with specific oceanographic features changed within the breeding season and between years. These results emphasise the importance of considering how foraging areas and reliance on specific oceanographic conditions change over time when seeking to identify important marine areas for seabirds.     

Can variations in the spatial distribution at sea and isotopic niche width be associated with consistency in the isotopic niche of a pelagic seabird species?

This study tested for fluctuations on short-term consistency (within about 1 month) in the isotopic niche of a pelagic seabird species. Short-term consistency in the isotopic niche was assessed using a wide-ranging apex predator, the Cory’s shearwaters Calonectris diomedea, along a 3-year study (2010–2012), during both the pre-laying and chick-rearing periods, with markedly inter- and intra-annual differences in the foraging spatial distribution at sea and isotopic niche width. We used individual movement data and stable isotope data, analysed using recent metrics based in a Bayesian framework, of 69 adults breeding on a small neritic island in the North Atlantic (39°24′N, 009°30′W). As expected, our results confirm that isotopic niche expansion could arise via increased variation in spatial distribution at sea among individuals. Results suggest fluctuations on short-term consistency in the isotopic niche of Cory’s shearwaters related to their different foraging patterns among periods and, ultimately, to presumably temporal changes in the availability and predictability of food resources. Short-term consistency in the isotopic niche was higher and persistent during periods when the population showed an intermediate isotopic niche width and absent when isotopic niche was either smaller or larger during the study period. These results suggest that consistency in the isotopic niche is an important characteristic of this population during the breeding period that may fluctuate depending on resources availability and should be important to understand the dynamics of foraging ecology of pelagic seabirds in general.     

Diets of clupeoid,stomiatoid and salmonoid fish of the Rockall Trough,northeastern Atlantic Ocean

The trophic interactions of species of fish of the continental slopes have not been investigated previously in detail. The present study examines the diets of the clupeoid, stomiatoid and salmonoid species occurring demersally and pelagically in the Rockall Trough to the west of Scotland and Ireland. Pelagic fish were collected between the surface and about 2 500-m depth between 1973 and 1978. Demersal fish were sampled at 250-m intervals of depths between 500- and 2 900-m depth during the years 1975 and 1981. Of the 28 species caught, 18 were strictly pelagic and dominated by the stomiatoids while five were strictly demersal. The remaining five species occurred in both the pelagic and demersal environment. The commonest clupeoid was Alepocephalus bairdii, which is a dominant species within the demersal fish associations at depths of 750 to 1 250m. This species, along with the deeper-living A. agassizi, and 3 other alepocephalid species feed primarily on benthopelagic prey but also exploit the epibenthos. The stomiatoid species, such as Cyclothone microdon, C. braueri, Maurolicus muelleri and Argyropelecus hemigymnus, dominate the associations of pelagic fish in the Trough. They feed on different proportions of ostracods, copepods and amphipods. Of the salmonoids, only Argentina silus and Bathylagus euryops occurred commonly and both feed on benthopelagic fauna, the latter at much greater depths than the former. The principal factor acting to prevent direct competition between species is the modal centres of bathymetric distribution of the species. Other contributing factors are briefly discussed.     

Age and sexual differences in the exploitation of two anthropogenic food resources for an opportunistic seabird

The availability of food resources has been suggested as a major factor in the substantial increase in reproductive output, survival, recruitment and, ultimately, population growth rates in most organisms. In fact, the artificial increase in food availability resulting from human activities has been suggested as a factor in the substantial increase in population size of several seabirds in recent decades. In the present study, our primary aim was to estimate the importance of the main natural prey and two alternative feeding resources, fishery discards and the invasive American crayfish Procambarus clarkii, for an opportunistic seabird, the Audouin’s gull Larus audouinii. We also assessed the influence of age and sex in the use of those three types of food. For this purpose, we compared the analyses of δ¹⁵N and δ¹³C in blood of male and female adults of known age and chicks with those in their potential prey. Our results reveal sex-related and age-related differences in the consumption of fish discards, small pelagic fish and American crayfish. Differences in the diet of males and females and also between adults and chicks could be related to different nutritional requirements. Age differences were probably related to their different foraging proficiency and the tendency of young breeders to opportunistically exploit anthropogenically derived food. This study illustrates the importance of considering the age and sex of individuals to obtain feasible dietary information and to understand how the exploitation of food of human origin could affect population growth.     

Can bottom trawling indirectly diminish carrying capacity in a marine ecosystem?

Bottom trawling is associated with reduced biomass and production in the marine benthic community. Abundance of hard-bodied organisms such as bivalves, crustaceans and echinoderms typically declines in favour of soft-bodied opportunists such as polychaetes. Trawling effects vary with habitat; impact and recovery time are typically greater for more complex substrates/communities and those with lower rates of natural disturbance. Benthic organisms represent the prey base of a large component of the demersal fish assemblage. Hence, trawling-induced change in benthic community structure and function may exert an indirect effect on feeding success and growth of important commercially exploited fishes such as plaice Pleuronectes platessa. We present habitat-specific mixed effects models of plaice length as a function of age, bottom-trawling effort, population density and near-bottom temperature, with sampling year and area, and fish sex incorporated as random effects. Across an observed gradient of trawling effort in the Celtic Sea, plaice on gravel habitat showed significant declines in length at age while plaice on sand habitat showed significant increases in length at age. Contrasting trawling effects likely reflect dietary differences between habitats. Plaice on sand substrates are known to consume predominately polychaetes, which may proliferate at moderate trawling intensity in this habitat. Conversely, plaice on gravel substrates are reported to consume more of the fragile organisms such as echinoderms and bivalves that show marked declines with bottom trawling. An indirect effect of trawling on prey availability and growth of demersal fish has substantial implications for fisheries sustainability via reduced ecosystem carrying capacity and production of commercial fish.     

Evaluating impacts of forage fish abundance on marine predators

Forage fish—small, low trophic level, pelagic fish such as herrings, sardines, and anchovies—are important prey species in marine ecosystems and also support large commercial fisheries. In many parts of the world, forage fish fisheries are managed using precautionary principles that target catch limits below the maximum sustainable yield. However, there are increasing calls to further limit forage fish catch to safeguard their fish, seabird, and marine mammal predators. The effectiveness of these extra-precautionary regulations, which assume that increasing prey abundance increases predator productivity, are under debate. In this study, we used prey-linked population models to measure the influence of forage fish abundance on the population growth rates of 45 marine predator populations representing 32 fish, seabird, and mammal species from 5 regions around the world. We used simulated data to confirm the ability of the statistical model to accurately detect prey influences under varying levels of influence strength and process variability. Our results indicate that predator productivity was rarely influenced by the abundance of their forage fish prey. Only 6 predator populations (13% of the total) were positively influenced by increasing prey abundance and the model exhibited high power to detect prey influences when they existed. These results suggest that additional limitation of forage fish harvest to levels well below sustainable yields would rarely result in detectable increases in marine predator populations.     

Foraging behavior of a generalist marine top predator,Japanese cormorants (<Emphasis Type="Italic">Phalacrocorax filamentosus</Emphasis>), in years of demersal versus epipelagic prey

To examine the behavioral adjustment of a generalist marine top predator to variability of their prey, we studied the foraging behavior of Japanese cormorants (Phalacrocorax filamentosus) breeding at Teuri Island, Hokkaido, in years of contrasting demersal and epipelagic prey composition. We used radio telemetry and ship-based surveys to determine behavior and at-sea distribution during three summers (1996–1998). The cormorants fed on epipelagic anchovy (Engraulis japonicus) and sandlance (Ammodytes personatus) in 1998 (year of epipelagic diet), while they fed on benthic rock fish (Sebastes spp.) and flatfish (Pleuronectidae) and nearshore-living naked sandlance (Hypophychus dybowskii), as well as epibenthic greenling (Hexagrammidae) in 1996 and 1997 (year of demersal diet). Cormorants engaged in larger feeding groups, visited more feeding sites, and stayed at each feeding site for a shorter period in the year of epipelagic diet than in the years of demersal diet. The cormorants made long foraging trips and fed in the mainland coastal habitat, distant from the colony, in the years of demersal diet. Individual radio-tracked birds fed over the wide area between the islands and mainland, in the year of epipelagic diet, while most individuals specialized in mainland or island coastal habitats in the years of demersal diet. Behavioral adjustment of Japanese cormorants might allow them to exploit both unpredictable epipelagic and predictable benthic prey efficiently.Communicated by T. Ikeda, Hakodate     

Restoring piscivorous fish populations in the Laurentian Great Lakes causes seabird dietary change

Ecosystem change often affects the structure of aquatic communities thereby regulating how much and by what pathways energy and critical nutrients flow through food webs. The availability of energy and essential nutrients to top predators such as seabirds that rely on resources near the water's surface will be affected by changes in pelagic prey abundance. Here, we present results from analysis of a 25-year data set documenting dietary change in a predatory seabird from the Laurentian Great Lakes. We reveal significant declines in trophic position and alterations in energy and nutrient flow over time. Temporal changes in seabird diet tracked decreases in pelagic prey fish abundance. As pelagic prey abundance declined, birds consumed less aquatic prey and more terrestrial food. This pattern was consistent across all five large lake ecosystems. Declines in prey fish abundance may have primarily been the result of predation by stocked piscivorous fishes, but other lake-specific factors were likely also important. Natural resource management activities can have unintended consequences for nontarget ecosystem components. Reductions in pelagic prey abundance have reduced the capacity of the Great Lakes to support the energetic requirements of surface-feeding seabirds. In an environment characterized by increasingly limited pelagic fish resources, they are being offered a Hobsonian choice: switch to less nutritious terrestrial prey or go hungry.     

Inter-annual changes in diet and foraging trip lengths in a small pelagic seabird, the thin-billed prion Pachyptila belcheri

Central place foragers are constrained in their foraging distribution by the necessity to return to their nest site at regular intervals. In many petrels that feed on patchily distributed prey from the sea surface over large foraging areas, alternating long and short foraging trips are used to balance the demands of the chick with the requirements of maintaining adult body condition. When the local conditions are favourable for prey density and quality, adults should be able to reduce the number of long foraging trips. We studied the flexibility in foraging trip lengths of a small pelagic petrel, the thin-billed prion Pachyptila belcheri, over three breeding seasons with increasingly favourable, cold-water conditions. During a warm-water influx in February 2006, chicks were fed less frequently and adults carried out foraging trips of up to 8 days. When conditions became more favourable with colder water temperatures in 2007 and 2008, thin-billed prions decreased trip lengths, more often attended their chick every day, and long foraging trips of six to eight days were not registered during 2008. Chick growth rates mirrored this, as chicks grew poorly during 2006, intermediate during 2007 and best during 2008. Thin-billed prions preyed mainly on squid during incubation and mainly on amphipods and euphausiids during chick-rearing. In the poorest season only, the diet was substantially supplemented with very small copepods. Together, the present results indicate that during warm-water conditions, thin-billed prions had difficulties in finding sufficient squid, amphipods or euphausiids and were forced to switch to lower trophic level prey, which they had to search for over large ocean areas.     

Underwater and above-water search patterns of an Arctic seabird: reduced searching at small spatiotemporal scales

How predators vary search patterns in response to prey predictability is poorly known. For example, marine invertebrates may be predictable but of low energy value, while fish may be of higher energy value but unpredictable at large (pelagic schools) or small (solitary benthics) spatial scales. We investigated the search patterns of the thick-billed murre (Uria lomvia), an Arctic seabird feeding on invertebrates, pelagic fish, or benthic fish. Foraging ranges at the Coats Island colony are generally smaller (<240 min per trip) than at larger colonies, and many birds specialize in foraging tactics and diet. Underwater search times for benthic fish were higher than for pelagic fish or invertebrates while above-water search times for pelagic fish were higher than for benthic fish or invertebrates. There were few stops during trips. Total trip time, flying time, number of flights, and number of dives were intercorrelated and increased with prey energy content, suggesting that longer trips involved fewer prey encounters due to selection of higher-quality, but rarer, prey items. Flight times were not Lévy-distributed and seabirds may have used area-restricted searches. The high degree of specialization, apparent absence of information center effects, and reduced above-water searching times may be linked to the relatively small colony size and the resulting short commuting distances to feeding areas, leading to greater prey predictability. We concluded that prey predictability over various scales affected predator search patterns.     

Diets of fishes of the upper continental slope of eastern Tasmania: content,calorific values,dietary overlap and trophic relationships

Diets of 15 species of demersal and pelagic fishes on the upper continental slope (420 to 550 m) were determined, based on samples taken every two months over 13 mo (April 1984 to April 1985) off eastern Tasmania. The calorific contribution of each prey item to the diets was determined. The fish could be divided into four trophic categories: pelagic piscivores, epibenthic piscivores, epibenthic invertebrate feeders and benthopelagic omnivores. Dietary overlap between the groups was low. The pelagic piscivores Apogonops anomalus, Trachurus declivis, Brama brama, Lepidopus caudatus and Macruronus novaezelandiae primarily consume the shelf-break myctophid Lampanyctodes hectoris; their diet is narrow, with a large overlap between species. The epibenthic piscivores Deania calcea and Genypterus blacodes both take a greater variety of prey, but have little dietary overlap. The fish feeding on epibenthic invertebrates, Coelorinchus sp. 2 and Centriscops humerosus, obtain most of their energy from benthic Crustacea and Ophiuroidea, supplemented with Lampanyctodes hectoris; the diet is broad, with little overlap. Among the benthopelagic omnivores (Cyttus traversi, Coelorinchus sp. 4, Lepidorhynchus denticulatus, Neocyttus rhomboidalis, Helicolenus percoides, Epigonus denticulatus and E. lenimen), most diets are broad and show slight overlap. All but E. denticulatus consume significant quantities of Lampanyctodes hectoris as well as Crustacea, particularly Polychaeta, Euphausiacea and Pyrosoma atlanticum. Seasonal changes in diet occurred in G. blacodes, T. declivis, Lepidopus caudatus, Coelorinchus sp. 4, Lepidorhynchus denticulatus, H. percoides, E. denticulatus and E. lenimen; these were related to changes in abundance of particular prey species, not to alterations in feeding habits. Only three species, Lepidopus caudatus, Coelorinchus sp. 2 and H. percoides, showed significant diel feeding periodicity. Ontogenetic dietary changes were evident in Cyttus traversi, Coelorinchus sp. 2, Lepidorhynchus denticulatus and H. percoides. Cyttus traversi and H. percoides progressively changed from crustaceans to fish as their size increased. The diets of size classes within species showed little overlap, except for Lepidorhynchus denticulatus, which eats chiefly euphausiids and Lampanyctodes hectoris at all sizes. In addition to describing the diets and trophic relationships of 90% of the fish biomass, the results emphasize the importance to the entire fish community of mesopelagic food resources, particularly Lampanyctodes hectoris. Many benthopelagic species undertake extensive vertical migrations in search of prey, thus playing a major role in the transport of energy from midwater regions to the benthos of the continental slope.     

Diet and nest attendance of incubating and chick-rearing northern fulmars (Fulmarus glacialis) in Shetland

We examined diets and nest attendance patterns of northern fulmars (Fulmarus glacialis) on Foula and Unst, Shetland, UK, during the breeding seasons of 1998 and 1999. Mean foraging trip duration, derived from nest attendance patterns, differed between incubation (32.5 h) and chick-rearing (early stage: 11.2 h, mid-stage: 20.4 h). It was influenced by chick age and obviously also by prey availability. The numbers of fulmars attending the colonies were influenced by wind direction, wind speed, time of day and state of reproduction. Diet samples were collected by regurgitations of adult and young fulmars. Diet was analysed by identifying prey items on the basis of otoliths, vertebrae, premaxillae (fish), cephalopod beaks and fragments of arthropod exoskeletons. Adult fulmars and chicks fed upon a wide range of prey types. Gadoid fish, including Norway pout (Trisopterus esmarkii), were the most common prey in the diet. Fish offal was found in 32% of regurgitates, clupeids in 15%. In contrast, the proportions of sandeels were very low (1%). This is different from previous studies where sandeels formed a large part of fulmar diet at Shetland in summer. Differences in food composition of incubating and chick-rearing fulmars were found for fish, but not for offal, crustaceans or squid. The percentage of non-discard fish (Clupeidae, Ammodytidae, Isospondylae) was significantly different between regurgitates from incubating and chick-rearing fulmars. Samples from chick-rearing adults contained non-discard items more frequently than regurgitates from incubating birds. The results strongly indicate that fulmars select to feed their chicks on energy-rich clupeids.     

Intra-sex vocal interactions in two hybridizing seabird species (Puffinus sp.)

Hybridization between two closely related species breeding in sympatry depends on the effectiveness of both inter-sex (sexual attraction) and intra-sex (competitors’ repulsion) components of communication. As birds primarily communicate acoustically, several studies have investigated interspecific vocal interactions between the sexes and their consequences on sympatric zones with a focus on songbird species. Here, we investigate these issues on intra-sex vocal interactions occurring during incubation in two non-songbird sister species, the Yelkouan Puffinus yelkouan and the Balearic Puffinus mauretanicus shearwaters. We compared the acoustical structure of calls and the behaviors obtained in response to same-sex Yelkouan and Balearic calls across allopatric parental populations from each species and a sympatric-hybridized population. Acoustic analyses showed that, for both sexes, calls have species-specific characteristics while hybrids have intermediate acoustic features compared to their parental species. Playback experiments showed that despite their vocal differences, both species interact and reply to each other. Remarkably, incubating Yelkouan and hybrid individuals resulted in equal intra-sex responses to either species while incubating Balearic birds led to lower territorial responses to Yelkouan than to conspecific calls. Moreover, incubating Balearic males less readily responded to same-sex Yelkouan calls than Balearic females. These results may have fitness consequences: Yelkouan birds, and especially males, would be more likely to win same-sex disputes against Balearic incubating birds than against Yelkouan incubating birds.