Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

A three-isotope approach to disentangling the diet of a generalist consumer: the yellow-legged gull in northwest Spain

The widespread omnivory of consumers and the trophic complexity of marine ecosystems make it difficult to infer the feeding ecology of species. The use of stable isotopic analysis plays a crucial role in elucidating trophic interactions. Here we analysed δ¹⁵N, δ¹³C and δ³⁴S in chick feathers, and we used a Bayesian triple-isotope mixing model to reconstruct the diet of a generalist predator, the yellow-legged gull (Larus michahellis) that breeds in the coastal upwelling area off northwest mainland Spain. The mixing model indicated that although chicks from all colonies were fed with a high percentage of fish, there are geographical differences in their diets. While chicks from northern colonies consume higher percentages of earthworms, refuse constitutes a more important source in the diet of chicks from western colonies. The three-isotope mixing model revealed a heterogeneity in foraging habitats that would not have been apparent if only two stable isotopes had been analysed. Moreover, our work highlights the potential of adding δ³⁴S for distinguishing not only between terrestrial and marine prey, but also between different marine species such as fish, crabs and mussels.     

Influence of nutrients in the feeding ecology of seagrass (Posidonia oceanica L.) consumers: a stable isotopes approach

Nitrogen inputs to coastal environments can considerably alter the abundance of primary producers. However, how herbivores modify their trophic signatures and adjust to changes in food resource conditions remains controversial. Here, we assess the effect of nutrient availability on the diet shifts of the two main Mediterranean herbivores, the Sparid fish Sarpa salpa L. and the sea urchin Paracentrotus lividus (Lmk.) that feed mostly on the seagrass Posidonia oceanica L. (Delile), epiphytes and benthic macroalgae. To do this, we (1) investigate the patterns of isotopic composition (δ¹³C and δ¹⁵N signatures) of the two herbivores and their potential food sources in three areas of contrasting nutrient conditions and, (2) we assess the diet shift along this nutrient gradient by estimating the isotopic nutrient enrichment (i.e., the contribution of δ¹³C and δ¹⁵N signatures in consumers’ tissues relative to potential food sources). Food web signatures of δ¹³C were similar among the three study sites, and no patterns of δ¹³C shift were observed in their diets. In contrast, there was a consistent increase in N contents and δ¹⁵N along the nutrient gradient for all primary producers and their consumers. The rate of δ¹⁵N enrichment was also clearly distinctive between the two herbivores: in P. lividus it increased by 61% along the nutrient gradient, while in S. salpa it remained constant. Our results suggest that sea urchins behave as facultative omnivores and feed on vegetable or mixed diets depending on the trophic status of the system. It is unclear, however, if this modification is behavioral or the consequence of mere changes in the availability of food items, as animal epiphytes (e.g., hydrozoans, bryozoans and ascidians) can also became more abundant on seagrass leaves under increased nutrient conditions. In contrast, adult fish appear to feed on vegetal material independent of nutrient availability in the ecosystem.     

The trophic link between squid and the emperor penguin <Emphasis Type="Italic">Aptenodytes forsteri</Emphasis> at Pointe Géologie,Antarctica

Cephalopod beaks retrieved from stomachs of dead emperor penguin chicks at Pointe Géologie, Terre Adélie, provide information on taxonomic and size composition of the penguin’s squid diet, on the trophic range of the squid species preyed upon and on the fractional trophic impact of the penguin on the whole food web. Emperor penguins prey upon four squid species (Psychroteuthis glacialis, Kondakovia longimana, Gonatus antarcticus, Alluroteuthis antarcticus) and do not take squid larger than 480 mm mantle length. Larger squid live either below the penguin’s diving range or are beyond its handling capacity. Nitrogen stable isotope ratios indicate that squids cover a range of about two trophic levels (2.5–8‰ δ¹⁵N). The impact of the emperor penguin, however, concentrates on the upper part of this range, about 68% of its squid prey being >6‰ δ¹⁵N. The principal components of the emperor’s diet, fish, krill and squid, differ distinctly in average trophic level. Consequently the trophic position of the emperor penguin changes accordingly with diet composition and may differ by almost one trophic level between different emperor penguin colonies.     

Isotopic niches of emperor and Adélie penguins in Adélie Land,Antarctica

The emperor and Adélie penguins are the only two species of penguins that co-occur at high-Antarctic latitudes. We first measured and compared their isotopic niches on the same year in Adélie Land in spring, when the two species co-exist. Emperor and Adélie penguins segregated by their blood isotopic signatures, with adult δ¹³C values (−24.5 ± 0.2 and −25.4 ± 0.2‰, respectively) suggesting that emperor penguins foraged in more neritic waters than Adélie penguins in spring. At that time, difference in their δ¹⁵N values (4.1‰, 12.0 ± 0.4 vs. 7.9 ± 0.1‰) encompassed more than one trophic level, indicating that emperor penguins preyed mainly upon fish (and squids), while Adélie penguins fed exclusively on euphausiids. Second, we compared the food of breeding adults and chicks. The isotopic signatures of adults and chicks of emperor penguins were not statistically different, but δ¹⁵N value of Adélie penguin chicks was higher than that of adults (10.2 ± 0.8 vs. 9.0 ± 0.2‰). The difference showed that adult Adélie penguins captured higher trophic level prey, i.e. higher-quality food, for their chicks. Third, the isotopic signatures of Adélie penguins breeding in Adélie Land showed that adults fed on Antarctic krill in oceanic waters in spring and shifted to neritic waters in summer where they preyed upon ice krill for themselves and upon fish and euphausiids for their chicks. A comparison of isotopic niches revealed large overlaps in both blood δ¹³C and δ¹⁵N values within the community of Antarctic seabirds and pinnipeds. The continuum in δ¹⁵N values nevertheless encompassed more than one trophic level (5.2‰) from Adélie penguin and crabeater seal to the Weddell seal. Such a broad continuum emphasizes the fact that all Antarctic seabirds and marine mammals feed on varying proportions of a few crustacean (euphausiids) and fish (Antarctic silverfish) species that dominate the intermediate trophic levels of the pelagic neritic and oceanic ecosystems.     

Spatio-temporal variations in the diet and stable isotope composition of the Argentine hake <Emphasis Type="Italic">Merluccius hubbsi</Emphasis> Marini, 1933 of the continental shelf of southeastern Brazil

The feeding ecology of Merluccius hubbsi was investigated in 2 regions of SE Brazil. The major food sources for the hakes were fish, crustaceans, and squid. In the upwelling system of Cabo Frio, the diet was very similar in the summers of 2001/2002 and spring 2002; fish were the most important prey followed by crustaceans. In Ubatuba, euphausiids were an important prey during the winter 2001 (100 m), while in the summer 2002, fish and amphipods predominated in the diet in the shallower site (40 m) and squid in the deeper site (100 m). The hakes showed temporal differences in stable isotope signatures in both regions, while C:N ratios varied only in Cabo Frio. δ¹⁵N and δ¹³C (bulk and corrected for lipid content) increased with fish length, which seems to be related to the increasing importance of fish and decreasing importance of euphausiids and amphipods in the diet of larger hakes. The mean trophic level of 3.7 for M. hubbsi was estimated using δ¹⁵N of bivalves as baseline and the fractionation of 3.4‰ between trophic levels.     

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.     

Comparative foraging ecology and ecological niche of a superabundant tropical seabird: the sooty tern Sterna fuscata in the southwest Indian Ocean

Over 6-million pairs of sooty terns Sterna fuscata breed once a year in the southwest Indian Ocean, mostly on three islands of the Mozambique Channel (Europa, Juan de Nova and Glorieuses) and in the Seychelles region. Seasonal reproduction in either winter or summer is the dominant strategy in the area, but non-seasonal reproduction also occurred in some places like at Glorieuses Archipelago. The feeding ecology of the sooty tern was investigated during the breeding seasons to determine whether terns showed significant differences in their trophic ecology between locations. Regurgitations were analyzed to describe the diet of individuals when breeding, and stable isotopes and mercury concentrations were used to temporally integrate over the medium-term of the trophic ecology of both adults and chicks. Overall, the diet was composed of fish, flying squid and fish larvae in different proportions. At Europa and Aride in the Seychelles, where winter reproduction occurs, large epipelagic prey like flying fish or squid dominated the diet. At Juan de Nova, sooty terns reproduce in summer and rely mostly on fish larvae. At Glorieuses (non-seasonal breeding), the diet was intermediate with fish larvae and flying squid being important prey items. The stable-carbon and nitrogen isotope values in blood confirm the differences observed in dietary analysis, and demonstrate different feeding strategies between colonies. δ¹³C values of feathers showed spatial segregation between birds from the Mozambique Channel and the Seychelles region. Terns from the Seychelles had also higher δ¹⁵N values. Feather δ¹³C values also suggest a significant shift from summer to wintering habitat for birds from Juan de Nova. This study emphasizes the high phenotypic plasticity of the species, which may explain its numerical dominance in all tropical waters of the World’s Ocean.     

Eutrophication and trophic structure in response to the presence of the eelgrass <Emphasis Type="Italic">Zostera noltii</Emphasis>

In estuaries, eelgrass meadows contribute to fundamental ecosystem functions of estuaries, providing food to several predators and buffering the negative effects of eutrophication. We asked whether the presence of the eelgrass Zostera noltii decreased the nitrogen concentration in the overlying water, affected the sources of nitrogen sequestrated by primary producers and changed the benthic and pelagic food web structures. We also studied the importance of these food webs in providing food to fish. We compared bare sediment to sediment covered by a Z. noltii meadow, and examined nutrient concentrations in the water column and δ¹⁵N in primary producers as indicators of anthropogenic inputs of nutrients. We then measured both δ¹³C and δ¹⁵N in the tissues of plants and consumers to establish food web structures. There were no differences in the concentrations and sources of nitrogen between sites. Rather, δ¹⁵N values indicated anthropogenic inputs of N (e.g. sewage discharges, agriculture) in both sites. There were no major differences in the structure of the planktonic food web, which was in part sustained by particulate organic matter and supported most predator fish, and in the structure of the benthic food web. Nonetheless, there were differences in the sources of food for omnivore consumers and for the detritivore Scrobicularia plana. Overall, the benthic food web did not use food derived from the eelgrass or macroalgae deposited on the substratum. Suspension feeders used particulate and sediment organic matter, whereas the δ¹³C and δ¹⁵N values of the other consumers indicated a likely contribution of benthic microalgae. Furthermore, in both habitats we found large variability in the isotope signatures of benthic macrofauna consumers, which did not allow distinguishing clearly different trophic groups and indicated a high level of omnivory and a mixed diet opportunistically making use of the availability of food in the surroundings.     

Use of multiple tools to assess the feeding preference of coastal dolphins

The feeding preferences of the coastal dolphins Pontoporia blainvillei and Sotalia guianensis in south-eastern Brazil (21o18′S–22o25′S) were assessed through the prey’s index of relative importance (IRI), total mercury concentration (Hg_tot), and stable isotopic (δ¹⁵N and δ¹³C) to compare their efficiency in the discrimination of prey contribution to the predators’ diet. The IRI was the best tool to describe the dolphins’ preference, while Hg_tot and δ¹⁵N seemed to be efficient as a trophic marker when the diet is made up of prey of varying sizes, as observed in S. guianensis. Both dolphins presented lighter δ¹⁵N than their prey species, which is an unusual pattern. However, as the sample size to isotope ratios analysis was small, especially to the dolphins, the results should be considered with caution, and further studies are necessary to corroborate these findings. The δ¹³C values characterized a typical coastal food chain, confirming the preferential area of these species.     

Stable isotope analysis in two sympatric populations of bottlenose dolphins <Emphasis Type="Italic">Tursiops truncatus</Emphasis>: evidence of resource partitioning?

Skin and muscle from 43 bottlenose dolphins (38 juveniles/adults, 5 calves) stranded in NW Spain were analysed to determine whether stable isotope ratios (δ¹³C and δ¹⁵N) could be used to assess dietary variation, habitat segregation and population substructure. Results were compared with published stomach contents data. Stable isotope ratios from 17 known prey species were also determined. Isotope ratios of the main prey (blue whiting, hake) varied significantly in relation to fish body size. Dolphin calves showed significant heavy isotope enrichments compared to adult females. Excluding calves, δ¹⁵N decreased with increasing dolphin body size, probably related to an ontogenetic shift in diet towards species at lower trophic levels, e.g. on blue whiting as suggested by stomach content results. Bottlenose dolphins were divided into two putative populations (North, South) based on previous genetic studies, and values of δ¹³C and δ¹⁵N differed significantly between these two groups, confirming the existence of population structuring.     

Age-related shifts in the diet composition of southern elephant seals expand overall foraging niche

Southern elephant seals are important apex predators in a highly variable and unpredictable marine environment. In the presence of resource limitation, foraging behaviours evolve to reduce intra-specific competition increasing a species’ overall probability of successful foraging. We examined the diet of 141 (aged 1–3 years) juvenile southern elephant seals to test the hypotheses that differences between ages, sexes and seasons in diet structure occur. We described prey species composition for common squid and fish species and the mean size of cephalopod prey items for these age groups. Three cephalopod species dominated the stomach samples, Alluroteuthis antarcticus, Histioteuthis eltaninae and Slosarczykovia circumantarcticus. We found age-related differences in both species composition and size of larger prey species that probably relate to ontogenetic changes in diving ability and haul-out behaviour and prey availability. These changes in foraging behaviour and diet are hypothesised to reduce intra-specific food competition concomitant with the increase in foraging niche of growing juveniles.     

Stable isotopic evidence for trophic groupings and bio-regionalization of predators and their prey in oceanic waters off eastern Australia

Muscle tissue was collected for stable isotope analysis (SIA) from the main fish predators and their fish and cephalopod prey from oceanic waters off eastern Australia between 2004 and 2006. SIA of δ¹⁵N and δ¹³C revealed that the species examined could be divided into three main trophic groups. A “top predator” group consisted mainly of large billfish (Xiphias gladius and Tetrapturus audax), yellowfin (Thunnus albacares), bigeye (T. obesus) and southern bluefin (T. maccoyii) tunas and sharks; with mako (Isurus oxyrinchus) the highest. Below this tier was a second group composed of mid-trophic level fishes including albacore tuna (Thunnus alalunga), lancet fish (Alepisaurus ferox), mahi mahi (Coryphaena hippuris) and ommastrephid squid. Underlying both groups was a grouping of small fishes including myctophids, small scombrids and nomeids as well as surface fishes including macrorhamphosids. These groupings were based largely on mean animal size which showed a positive linear relation to δ¹⁵N (r ² = 0.58). Some species showed significant ontogenetic variation in either δ¹⁵N (swordfish, lancet fish, yellowfin and albacore tuna) or δ¹³C (mako shark). We also noted a consistent latitudinal change in δ¹⁵N and δ¹³C at ~28°S for the top predator species, particularly albacore and yellowfin tuna. The differences were consistent with a change from oligotrophic Coral Sea to nutrient rich Tasman Sea waters. These differences suggest that predatory fishes may have extended residence time in distinct regions off eastern Australia.     

Food sources used by sediment meiofauna in an intertidal Zostera noltii seagrass bed: a seasonal stable isotope study

In an intertidal Zostera noltii Hornem seagrass bed, food sources used by sediment meiofauna were determined seasonally by comparing stable isotope signatures (δ¹³C, δ¹⁵N) of sources with those of nematodes and copepods. Proportions of different carbon sources used by consumers were estimated using the SIAR mixing model on δ¹³C values. Contrary to δ¹⁵N values, food source mean δ¹³C values encompassed a large range, from −22.1 ‰ (suspended particulate organic matter) to −10.0 ‰ (Z. noltii roots). δ¹³C values of copepods (from −22.3 to −12.3 ‰) showed that they use many food sources (benthic and phytoplanktonic microalgae, Z. noltii matter). Nematode δ¹³C values ranged from −14.6 to −11.4 ‰, indicating a strong role of microphytobenthos and/or Z. noltii matter as carbon sources. The difference of food source uses between copepods and nematodes is discussed in light of source accessibility and availability.     

Isotopic evidence of distinct feeding ecologies and movement patterns in two migratory predators (yellowfin tuna and swordfish) of the western Indian Ocean

Ecologists primarily use δ¹⁵N values to estimate the trophic level of organisms, while δ¹³C, and even recently δ¹⁵N, are utilized to delineate feeding habitats. However, many factors can influence the stable isotopic composition of consumers, e.g. age, starvation or isotopic signature of primary producers. Such sources of variability make the interpretation of stable isotope data rather complex. To examine these potential sources of variability, muscle tissues of yellowfin tuna (Thunnus albacares) and swordfish (Xiphias gladius) of various body lengths were sampled between 2001 and 2004 in the western Indian Ocean during different seasons and along a latitudinal gradient (23°S to 5°N). Body length and latitude effects on δ¹⁵N and δ¹³C were investigated using linear models. Both latitude and body length significantly affect the stable isotope values of the studied species but variations were much more pronounced for δ¹⁵N. We explain the latitudinal effect by differences in nitrogen dynamics existing at the base of the food web and propagating along the food chain up to top predators. This spatial pattern suggests that yellowfin and swordfish populations exhibit a relatively unexpected resident behaviour at the temporal scale of their muscle tissue turnover. The body length effect is significant for both species but this effect is more pronounced in swordfish as a consequence of their different feeding strategies, reflecting specific physiological abilities. Swordfish adults are able to reach very deep water and have access to a larger size range of prey than yellowfin tuna. In contrast, yellowfin juveniles and adults spend most of their time in the surface waters and large yellowfin tuna continue to prey on small organisms. Consequently, nitrogen isotopic signatures of swordfish tissues are higher than those of yellowfin tuna and provide evidence for different trophic levels between these species. Thus, in contrast to δ¹³C, δ¹⁵N analyses of tropical Indian Ocean marine predators allow the investigation of complex vertical and spatial segregation, both within and between species, even in the case of highly opportunistic feeding behaviours. The linear models developed in this study allow us to make predictions of δ¹⁵N values and to correct for any body length or latitude differences in future food web studies.     

Stable isotopes identify age- and sex-specific dietary partitioning and foraging habitat segregation in southern giant petrels breeding in Antarctica and southern Patagonia

We examined the isotopic signatures (δ¹³C, δ¹⁵N) of adult body feathers from southern giant petrels Macronectes giganteus collected at two breeding colonies in Antarctica (Potter Peninsula and Cape Geddes) and one in southern Patagonia (Observatorio Island), as well as in whole blood collected from adults of both sexes at each Antarctic colonies and from chicks at Potter Peninsula. As body feather moult is a continuous process in giant petrels, feathers provide an integrated annual signal of an adult’s diets and foraging habitats. In contrast, the stable isotope values of adult and chick blood are reflective of their diets during the breeding season. We found that sex-specific dietary segregation in adults breeding in Antarctica was notable during the breeding season (blood samples) but absent when examined across the entire year (feather samples). In addition, blood stable isotope values differed between chicks and adults, indicating that adults provision their offspring with a relatively higher amount of penguin and seal prey that what they consume themselves. This finding confirms previous work that suggests that chicks are preferentially fed with prey of presumably higher nutritional value such as carrion. Finally, based on isotopic differences between major oceanographic zones in the Southern Ocean, our data indicate population-specific differences in foraging distribution, with Antarctic populations move seasonally between Antarctic and subantarctic zones, while Patagonian populations likely forage in subtropical waters and in continental shelf habitats year-round.     

Food of emperor penguins (Aptenodytes forsteri) in the western Ross Sea, Antarctica

The diet of the emperor penguin Aptenodytes forsteri in the western Ross Sea during spring was investigated by analysis of stomach contents sampled at three different localities. At Cape Washington, emperor penguins feeding chicks consistently preyed on fishes (89 to 95% by mass) and crustaceans (5 to 11%) over the four spring seasons examined. By far the commonest prey was the Antarctic silverfish Pleuragramma antarcticum (89% of the fish prey); the remainder of fish prey were mainly unidentified juveniles of different species of channichthyid fishes. Three species dominated the crustacean part of the diet, i.e. the gammarid amphipods Abyssorchomene rossi/plebs (30% of the crustacean prey) and Eusirus microps (22%), together with the euphausiid Euphausia crystallorophias (24%). At Coulman Island and Cape Roget, fishes, mainly P. antarcticum, formed the bulk of the food (88 and 93% by mass, respectively), crustaceans were minor prey (2.5 and 0.4%), and the squid Psychroteuthis glacialis accounted for a small but significant part of the food (3.5 and 0.8%). This study emphasizes the importance of the small, shoaling pelagic fish Pleuragramma antarcticum as a key link between zooplankton and top predators, including seabirds, in the food web and marine ecosystem of the Ross Sea. Received: 20 May 1997 / Accepted: 8 October 1997     

Temporal and spatial variation in the δ<Superscript>15</Superscript>N and δ<Superscript>13</Superscript>C values of fish and squid from Alaskan waters

To test the hypothesis that stable isotope ratios from marine organisms vary, the δ¹⁵N and δ¹³C values from fish and squid collected in Alaskan waters were measured across years (1997, 2000, and 2005), seasons, geographic locations, and different size/age classes, and between muscle tissue and whole animals. Temporal, geographic, and ontogenetic differences in stable isotope ratios ranged from 0.5–2.5‰ (δ¹⁵N) to 0.5–2.0‰ (δ¹³C). Twenty-one comparisons of stable isotope values between whole organisms and muscle tissue revealed only four small differences each for δ¹⁵N and δ¹³C, making costly and space prohibitive collection of whole animals unnecessary. The data from this study indicate that significant variations of stable isotope values from animals in marine systems necessitates collection of prey and predator tissues from the same time and place for best interpretation of stable isotope analysis in foraging ecology studies.     

Central place foraging by breeding Cook’s petrel Pterodroma cookii: foraging duration reflects range, diet and chick meal mass

Pelagic seabirds are central place foragers during breeding and variation in foraging trip duration and range reflect differences in diet and chick provisioning, through the exploitation of divergent habitats of varying productivity. We tested whether these relationships hold in small procellarriids by equipping chick-rearing Cook’s petrel Pterodroma cookii (200 g) with geolocation-immersion loggers, conducting isotope analysis of blood and measuring chick meal mass following foraging trips of varying duration. Cook’s petrel tracked during chick rearing from Little Barrier Island (LBI) and Codfish Island (CDF), New Zealand had larger maximum ranges during longer foraging trips. Blood nitrogen isotope signatures (δ¹⁵N) of adults were significantly higher after foraging trips of longer duration, but not of greater maximum range. There was no significant relationship between blood carbon isotope signatures (δ¹³C) and foraging trip characteristics. Proportion of time spent on the sea surface and the mass of the meal brought back to chicks were consistently greater for Cook’s petrel with larger maximum ranges, which in the case of birds from CDF coincided with productive subtropical convergence zone habitats. As predicted, trip duration reflected divergent foraging behaviours in Cook’s petrel during breeding. We suggest that the availability of different prey is a key factor governing at-sea distributions and dietary composition of this species.     

Swimming behavior of juvenile anchovies (<Emphasis Type="Italic">Anchoa</Emphasis> spp.) in an episodically hypoxic estuary: implications for individual energetics and trophic dynamics

Diel swimming behaviors of juvenile anchovies (Anchoa spp.) were observed using stationary hydroacoustics and synoptic physicochemical and zooplankton profiles during four unique water quality scenarios in the Neuse River Estuary, NC, USA. Vertical distribution of fish was restricted to waters with DO greater than 2.5 mg O₂ l⁻¹, except when greater than 70% of the water column was hypoxic and a subset of fish were occupying water with 1 mg O₂ l⁻¹. We made the prediction that an individual fish would select a swim speed that would maximize net energy gain given the abundance and availability of prey in the normoxic waters. During the day, fish adopted swim speeds between 7 and 8.8 bl s⁻¹ that were near the theoretical optimum speeds between 7.0 and 8.0 bl s⁻¹. An exception was found during severe hypoxia, when fish were swimming at 60% above the optimum speed (observed speed = 10.6 bl s⁻¹, expected = 6.4 bl s⁻¹). The anchovy is a visual planktivore; therefore, we expected a diel activity pattern characteristic of a diurnal species, with quiescence at night to minimize energetic costs. Under stratified and hypoxic conditions with high fish density coupled with limited prey availability, anchovies sustained high swimming speeds at night. The sustained nighttime activity resulted in estimated daily energy expenditure over 20% greater than fish that adopted a diurnal activity pattern. We provide evidence that the sustained nighttime activity patterns are a result of foraging at night due to a lower ration achieved during the day. During severe hypoxic events, we also observed individual fish making brief forays into the hypoxic hypolimnion. These bottom waters generally contained higher prey (copepod) concentrations than the surface waters. The bay anchovy, a facultative particle forager, adopts a range of behaviors to compensate for the effects of increased conspecific density and reduced prey availability in the presence of stratification-induced hypoxia.