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.     

Feeding ecology and niche segregation in oceanic top predators off eastern Australia

We examined the feeding ecology and niche segregation of the ten most abundant fish species caught by longline operations off eastern Australia between 1992 and 2006. Diets of 3,562 individuals were examined. Hook timer data were collected from a further 328 fish to examine feeding behaviour in relation to depth and time of day. Prey biomass was significantly related to predator species, predator length and year and latitude of capture. Although the fish examined fed on a mix of fish, squid and crustacea, fish dominated the diet of all species except small albacore (Thunnus alalunga) which fed mainly on crustacea and large swordfish (Xiphias gladius) and albacore which fed mainly on squid. Cannibalism was observed in lancetfish (Alepisaurus spp.). Multidimensional scaling identified three species groups based on their diet composition. One group consisted of yellowfin tuna (T. albacares), striped marlin (Tetrapturus audax) and dolphinfish (Coryphaena hippurus); a second group consisted of bigeye tuna (T. obesus), swordfish and albacore; and a third consisted of southern bluefin tuna (T. maccoyii) and blue shark (Prionace glauca). Of note was the separation of mako shark (Isurus oxyrhynchus) and lancetfish from all other predators. Prey length generally increased with increasing predator length although even large predators fed on a wide range of prey lengths including very small prey. Overall, differences in prey type and size, feeding times and depths were noted across the range of species examined to the extent that predators with overlapping prey, either in type or size, fed at different times of the diel period or at different depths. Taken together these data provide evidence for feeding niche segregation across the range of oceanic top predators examined.     

Diet and trophic position of Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) inferred from stable carbon and nitrogen isotope analysis

Stable ¹³C and ¹⁵N isotope analyses of scale, bone, and muscle tissues were used to investigate diet and trophic position of North Atlantic bluefin tuna (Thunnus thynnus Linnaeus) during residency in the northwest Atlantic Ocean off the northeast coast of the United States. Adult bluefin tuna scales collected from fish between June and October 2001 were significantly enriched in ¹³C compared to both muscle and bone across all months, while muscle was significantly enriched in ¹⁵N compared to either bone or scale throughout the same period. In muscle tissue, there was evidence of a shift over the summer from prey with ¹³C values (–17 to –18) that were characteristic of silver hake (Merluccius bilinearis) to species with ¹³C values of –20 to –21 that were similar to Atlantic herring (Clupea harengus) and sandlance (Ammodytes americanus). Depletion of ¹⁵N values in adult scales and bone compared to muscle tissue may be explained by bone and scale samples representing juvenile or life-long feeding habits, isotopic routing, or isotopic differences in amino acid composition of the three tissue types. Adult bluefin tuna were estimated to be feeding at a trophic position similar to pelagic sharks in the northwest Atlantic Ocean, while the trophic positions of yellowfin tuna (Thunnus albacares), albacore tuna (Thunnus alalunga), and juvenile bluefin tuna were indicative of a diet of up to a full trophic position below adult bluefin tuna. The close relationship between the juvenile bluefin ¹⁵N values and those of suspension feeders suggests that nektonic crustaceans or zooplankton may contribute significantly to the diet of bluefin tuna, a food source previously overlooked for this species in the northwest Atlantic Ocean.Communicated by J.P. Grassle, New Brunswick     

Trophic niche overlap among dolphinfish and co-occurring tunas near the northern edge of their range in the western North Atlantic

We present the first quantitative analyses of dolphinfish (Coryphaena hippurus) foraging habits and trophic interactions with co-occurring yellowfin (Thunnus albacares) and albacore (T. alalunga) tunas in the Southern New England region of the western North Atlantic Ocean. Fish caught by recreational anglers in offshore waters of Massachusetts were sampled during the summers of 2007–2010. Diet analysis revealed that shortfin squid (Illex illecebrosus) and small pelagic crustaceans were principal prey to dolphinfish, yellowfin tuna, and albacore tuna. A wide variety of Sargassum-associated fishes were also important to dolphinfish and yellowfin tuna diets. Dietary (Schoener’s index: 0.82–0.86) and isotopic niche (isotopic ellipse overlap: 53.6–64.7 %) overlap was high, and dolphinfish and tunas occupied equivalent trophic positions (TP = 3.4–3.6). Relative prey size in dolphinfish and yellowfin tuna diets exhibited convergence with ontogeny. Overall, dolphinfish had the greatest isotopic niche width, which was twice as large as yellowfin tuna and three times as large as albacore tuna; dolphinfish also consumed the greatest range of prey sizes. Results quantify dolphinfish trophic interactions in the western Atlantic near the northern extent of their geographical range, and are relevant for ecosystem-based management of the offshore pelagic guild in the context of shifting fish populations and fisheries in response to climate and ecological change.     

A rapid ontogenetic shift in the diet of juvenile yellowfin tuna from Hawaii

Within the tropical and subtropical oceans, tuna forage opportunistically on a wide variety of prey. However, little is known about the trophic ecology of the smallest size classes which play an important role in stock assessments and fisheries management. The foraging behavior of yellowfin tuna, Thunnus albacares (23.5–154.0 cm FL), collected from nearshore Fish Aggregating Devices (FADs) around Oahu was studied using stable isotope and stomach contents analyses. Emphasis was placed on small juveniles. Yellowfin tuna changed their diets significantly between 45 and 50 cm forklength (ca. 1.5 kg). Smallest size classes fed on planktonic organisms inhabiting the shallow mixed layer, primarily larval stomatopod and decapod crustaceans, whereas larger tuna fed on teleosts and adult Oplophorus gracilirostris, a vertically migrating mesopelagic species of shrimp. When interpreting the variation in prey δ ¹⁵N values, we considered both their relative trophic position and δ ¹⁵N values of the nitrogen at the base of the food web. Based on the distinct diet shift of the yellowfin tuna, demonstrated by both isotope and stomach content analyses, we propose a critical mass threshold was reached at about 45 cm FL that enabled sufficient endothermic capability to allow tuna to access prey dwelling in deeper, colder water. These ontogenetic changes in foraging range and commensurate shift in diet of small tunas would affect their vulnerability to fishing pressure.     

Variability of albacore (<Emphasis Type="Italic">Thunnus alalunga</Emphasis>) diet in the Northeast Atlantic and Mediterranean Sea

This study aims to describe the variability of albacore (Thunnus alalunga) diet in the Northeast Atlantic and Mediterranean Sea and to identify possible relationships between this variability and the features of different feeding areas, the behavior, and the energetic needs of albacore. Stomach contents from albacore caught in five zones of the Bay of Biscay and surrounding waters (n = 654) and three zones of the Mediterranean Sea (n = 152) were analyzed in terms of diet composition and stomach fullness. Carbon and nitrogen stable isotope and C/N ratios were measured for white muscle and liver from albacore in the Bay of Biscay (n = 41) and Mediterranean Sea (n = 60). Our results showed a spatial, seasonal, inter-annual, and size-related variability in the diet of albacore. Albacore diet varied by location in the Mediterranean Sea, with a particularly high proportion of cephalopods, and low δ¹⁵N values in the Tyrrhenian Sea. In the Northeast Atlantic, albacore consumed a higher proportion of crustaceans and a lower proportion of fishes in the most offshore sampling zone than inshore. The digestion states of the major prey reflected a diurnal feeding activity, indicative of feeding in deeper waters offshore, whereas on the continental slope, feeding probably occurred in surface waters at night. Important seasonal and inter-annual diet variability was observed in the southeast of the Bay of Biscay, where preferred albacore prey appeared to be anchovy (Engraulis encrasicolus). Stomach fullness was inversely related to body size, probably reflecting higher energetic needs for smaller individuals. Albacore from the Bay of Biscay had significantly lower δ¹³C and higher δ¹⁵N values compared with albacore from the Mediterranean Sea, indicative of regional baseline shifts, and trophic position and muscle lipid stores in albacore increased with body size.     

Environmental preferences of yellowfin tuna (Thunnus albacares) at the northern extent of its range

We used acoustic telemetry to examine the small-scale movement patterns of yellowfin tuna (Thunnus albacares) in the California Bight at the northern extent of their range. Oceanographic profiles of temperature, oxygen, currents and fluorometry were used to determine the relationship between movements and environmental features. Three yellowfin tuna (8 to 16 kg) were tracked for 2 to 3 d. All three fish spent the majority of their time above the thermocline (18 to 45 m in depth) in water temperatures >17.5 °C. In the California Bight, yellowfin tuna have a limited vertical distribution due to the restriction imposed by temperature. The three fish made periodic short dives below the thermocline (60 to 80 m), encountering cooler temperatures (>11 °C). When swimming in northern latitudes, the depth of the mixed layer largely defines the spatial distribution of yellowfin tuna within the water column. Yellowfin prefer to spend most of their time just above the top of the thermocline. Oxygen profiles indicated that the tunas encountered oceanic water masses that ranged most often from 6.8 to 8.6 mg O₂ l⁻¹, indicating no limitation due to oxygen concentrations. The yellowfin tuna traveled at speeds ranging from 0.46 to 0.90 m s⁻¹ (0.9 to 1.8 knots h⁻¹) and frequently exhibited an oscillatory diving pattern previously suggested to be a possible strategy for conserving energy during swimming. Received: 14 February 1997 / Accepted: 14 April 1997     

ATP content and adenylate energy charge of the mussel Mytilus edulis during the annual reproductive cycle in Lindåspollene,Western Norway

The oxygen consumption rates ( VO₂) of 6 specimens (6 to 13 kg) of the albacore tuna Thunnus alalunga were measured at sea, using specimens collected 300 km west of San Diego, California (USA) during July and August, 1981. Fish were tested in a closed continuous-flow respirometer, where they swam at about 1.3 body lengths s^-1 velocity in 15° to 19°C water. The albacore tuna is a temperate pelagic species experiencing water temperatures from about 10° to 20°C and attaining a maximum weight of 45 kg. The VO₂ ranged from 1 249 to 3 336 ml h^-1 (the mean VO₂ for the 6 fish was 2 228 ml h^-1); such values approach those of mammals of a similar size and are 3 to 4 times those of most active fishes (e.g. sockeye salmon). Among fishes, the only higher VO₂ values yet recorded were for the skipjack tuna Katsuwonus pelamis, a tropical species. The remarkably high metabolic rates of tunas are presumably correlated with their continuous swimming activity and the maintenance of endothermy. The exponent relating VO₂ to body weight (1.18), although large, is not statistically different from the exponents for most other active vertebrates.     

Horizontal movements and depth distribution of large adult yellowfin tuna (Thunnus albacares) near the Hawaiian Islands, recorded using ultrasonic telemetry: implications for the physiological ecology of pelagic fishes

We measured the horizontal and vertical movements of five adult yellowfin tuna (Thunnus albacares, estimated body mass 64 to 93 kg) near the main Hawaiian Islands, while simultaneously gathering data on oceanographic conditions and currents. Fish movements were recorded by means of ultrasonic depth-sensitive transmitters. Depth–temperature and depth–oxygen profiles were measured with vertical conductivity–temperature–depth (CTD) casts, and the current-velocity field was surveyed using an acoustic Doppler current profiler (ADCP). Large adult yellowfin tuna spent ≃60 to 80% of their time in or immediately below the relatively uniform-temperature surface-layer (i.e. above 100 m), a behavior pattern similar to that previously reported for juvenile yellowfin tuna, blue marlin (Makaira nigricans), and striped marlin (Tetrapturus audax) tracked in the same area. In all three species, maximum swimming depths appear to be limited by water temperatures 8 C° colder than the surface-layer water temperature. Therefore, neither large body mass, nor the ability to maintain elevated swimming-muscle temperatures due to the presence of vascular counter-current heat exchangers in tunas, appears to permit greater vertical mobility or the ability to remain for extended periods below the thermocline. In those areas where the decrease in oxygen with depth is not limiting, the vertical movements of yellowfin tuna, blue marlin and striped marlin all appear to be restricted by the effects of water temperature on cardiac muscle function. Like juvenile yellowfin tuna, but unlike blue marlin and striped marlin, adult yellowfin tuna remained within 18.5 km of the coast and became associated with floating objects, including anchored fish-aggregating devices (FADs) and the tracking vessel. Like juvenile yellowfin tuna, large adult yellowfin repeatedly re-visit the same FAD, and appear able to navigate precisely between FADs that are up to 18 km apart. The median speed over ground ranged from 72 to 154 cm s⁻¹. Neither speed nor direction was strongly influenced by currents. Received: 27 March 1998 / Accepted: 13 November 1998     

Gill and intestinal Na+-K+ ATPase activity, and estimated maximal osmoregulatory costs, in three high-energy-demand teleosts: yellowfin tuna (Thunnus albacares), skipjack tuna (Katsuwonus pelamis), and dolphin fish (Coryphaena hippurus)

We hypothesize that the morpho-physiological adaptations that permit tunas to achieve maximum metabolic rates (MMR) that are more than double those of other active fishes should result in high water and ion flux rates across the gills and concomitant high osmoregulatory costs. The high standard metabolic rates (SMR) of tunas and dolphin fish may, therefore, be due to the elevated rates of energy expenditure for osmoregulation (i.e. teleosts capable of achieving exceptionally high MMR necessarily have SMR). Previous investigators have suggested a link between activity patterns and osmoregulatory costs based on Na⁺-K⁺ ATPase activity in the gills of active epipelagic and sluggish deep-sea fishes. Based on these observations, we conclude that high-energy-demand fishes (i.e. tunas and dolphin fish) should have exceptionally elevated gill and intestinal Na⁺-K⁺ ATPase activity reflecting their elevated rates of salt and water transfer. To test this idea and estimate osmoregulatory costs, we measured Na⁺-K⁺ ATPase activity (V _max) in homogenates of frozen samples taken from the gills and intestines of skipjack and yellowfin tunas, and the gills of dolphin fish. As a check of our procedures, we made similar measurements using tissues from hybrid red tilapia (Oreochromis mossambicus ×O. niloticus). Contrary to our supposition, we found no difference in Na⁺-K⁺ ATPase activity per unit mass of gill or intestine in these four species. We estimate the cost of osmoregulation to be at most 9% and 13% of the SMR in skipjack tuna and yellowfin tuna, respectively. Our results, therefore, do not support either of our original suppositions, and the cause(s) underlying the high SMR of tunas and dolphin fish remain unexplained. Received: 7 September 2000 / Accepted: 4 December 2000     

Self-fertilisation in Celleporella hyalina

The accumulation of parasites in a fish host is modelled as a function of the total amount of prey consumed. The accumulated parasite load is then expressed as a function of fish length so that the asymptotic growth, L , of any population of commonly infected fish can be estimated. Estimates of L are obtained for orange roughy (Hoplostethus atlanticus), albacore tuna (Thunnus alalunga) and the New Zealand southern arrow squid (Nototodarus sloanii).     

Blood oxygen-binding characteristics of bigeye tuna (Thunnus obesus), a high-energy-demand teleost that is tolerant of low ambient oxygen

 We found blood from bigeye tuna (Thunnus obesus) to have a significantly higher O₂ affinity than blood from other tunas. Its P₅₀ (partial pressure of oxygen, P_O2 required to reach 50% saturation) was 1.6 to 2.0 kPa (12 to 15 mmHg) when equilibrated with 0.5% CO₂. Previous studies employing similar methodologies found blood from yellowfin tuna (T. albacares), skipjack tuna (Katsuwonus pelamis), and kawakawa (Euthynnus affinis) to have a P₅₀ of 2.8 to 3.1 kPa (21 to 23 mmHg). These observations suggest that bigeye tuna are more tolerant of low ambient oxygen than other tuna species, and support similar conclusions derived from laboratory whole-animal studies, depth-of-capture data, and directly-recorded vertical movements of fish in the open ocean. We also found the O₂ affinity of bigeye tuna blood to be essentially unaffected by a 10 C° open-system temperature change (as is the blood of all tuna species studied to date). The O₂ affinity of bigeye tuna blood was, however, more affected by a 10 C° closed-system temperature change than the blood of any tuna species yet examined. In other words, bigeye tuna blood displayed a significantly enhanced Bohr effect (change in log P₅₀ per unit change in plasma pH at P₅₀) when subjected to the inevitable changes in partial pressure of carbon dioxide (P_CO2) and plasma pH that accompany closed-system temperature shifts, than when subjected to changes in plasma pH accomplished by changing P_CO2 alone. In vivo, the resultant large decrease in O₂ affinity (i.e. the increase in P₅₀) that occurs as the blood of bigeye tuna is warmed during its passage through the vascular counter-current heat exchangers ensures adequate rates of O₂ off-loading in the swimming muscles of this high-energy-demand teleost. Received: 12 March 1999 / Accepted: 18 December 1999     

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.     

Longitudinal and vertical trends in stable isotope signatures (δ13C and δ15N) of omnivorous and carnivorous copepods across the South Atlantic Ocean

Stable isotope (SI) ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) were measured in omnivorous and carnivorous deep-sea copepods of the families Euchaetidae and Aetideidae across the Atlantic sector of the Southern Ocean to establish their trophic positions. Due to high and variable C/N ratios related to differences in lipid content, δ¹³C was corrected using a lipid-normalisation model. δ¹⁵N signals ranged from 3.0–6.9‰ in mesopelagic species to 7.0–9.5‰ in bathypelagic congeners. Among the carnivorous Paraeuchaeta species, the epi- to mesopelagic species Paraeuchaeta antarctica had lower δ¹⁵N values than the mesopelagic P. rasa and bathypelagic P. barbata. The same trend was observed among omnivorous Aetideidae, but was not significant. In the most abundant species P. antarctica, individuals from the western Atlantic had higher δ¹³C and δ¹⁵N values than specimens at the eastern stations. These longitudinal changes in δ¹³C and δ¹⁵N values were attributed to regional differences in hydrography and sea surface temperature (SST), in particular related to a northward extension of the Antarctic Polar Front (APF) at the easternmost stations. The results indicate that even in a mesopelagic carnivorous species, the changes in surface stable isotope signatures are pronounced.     

Genetic analysis of tuna populations,<Emphasis Type="Italic"> Thunnus thynnus thynnus</Emphasis> and<Emphasis Type="Italic"> T</Emphasis>.<Emphasis Type="Italic"> alalunga</Emphasis>

The genetic population structures of Atlantic northern bluefin tuna ( Thunnus thynnus thynnus) and albacore ( T. alalunga) were examined using allozyme analysis. A total of 822 Atlantic northern bluefin tuna from 18 different samples (16 Mediterranean, 1 East Atlantic, 1 West Atlantic) and 188 albacore from 5 samples (4 Mediterranean, 1 East Atlantic) were surveyed for genetic variation in 37 loci. Polymorphism and heterozygosity reveal a moderate level of genetic variability, with only two highly polymorphic loci in both Atlantic northern bluefin tuna ( FH* and SOD- 1*) and albacore ( GPI- 3* and XDH*). The level of population differentiation found for Atlantic northern bluefin tuna and albacore fits the pattern that has generally been observed in tunas, with genetic differences on a broad rather than a more local scale. For Atlantic northern bluefin tuna, no spatial or temporal genetic heterogeneity was observed within the Mediterranean Sea or between the East Atlantic and Mediterranean, indicating the existence of a single genetic grouping on the eastern side of the Atlantic Ocean. Very limited genetic differentiation was found between West Atlantic and East Atlantic/Mediterranean northern bluefin tuna, mainly due to an inversion of SOD- 1* allele frequencies. Regarding albacore, no genetic heterogeneity was observed within the Mediterranean Sea or between Mediterranean and Azores samples, suggesting the existence of a single gene pool in this area.     

Behavior of yellowfin (<Emphasis Type="Italic">Thunnus albacares</Emphasis>) and bigeye (<Emphasis Type="Italic">T. obesus</Emphasis>) tuna in a network of fish aggregating devices (FADs)

The influence of multiple anchored fish aggregating devices (FADs) on the spatial behavior of yellowfin (Thunnus albacares) and bigeye tuna (T. obesus) was investigated by equipping all thirteen FADs surrounding the island of Oahu (HI, USA) with automated sonic receivers (“listening stations”) and intra-peritoneally implanting individually coded acoustic transmitters in 45 yellowfin and 12 bigeye tuna. Thus, the FAD network became a multi-element passive observatory of the residence and movement characteristics of tuna within the array. Yellowfin tuna were detected within the FAD array for up to 150 days, while bigeye tuna were only observed up to a maximum of 10 days after tagging. Only eight yellowfin tuna (out of 45) and one bigeye tuna (out of 12) visited FADs other than their FAD of release. Those nine fish tended to visit nearest neighboring FADs and, in general, spent more time at their FAD of release than at the others. Fish visiting the same FAD several times or visiting other FADs tended to stay longer in the FAD network. A majority of tagged fish exhibited some synchronicity when departing the FADs but not all tagged fish departed a FAD at the same time: small groups of tagged fish left together while others remained. We hypothesize that tuna (at an individual or collective level) consider local conditions around any given FAD to be representative of the environment on a larger scale (e.g., the entire island) and when those conditions become unfavorable the tuna move to a completely different area. Thus, while the anchored FADs surrounding the island of Oahu might concentrate fish and make them more vulnerable to fishing, at a meso-scale they might not entrain fish longer than if there were no (or very few) FADs in the area. At the existing FAD density, the ‘island effect’ is more likely to be responsible for the general presence of fish around the island than the FADs. We recommend further investigation of this hypothesis.

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Diet of the social groups of long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas</Emphasis>) in the Strait of Gibraltar

The Strait of Gibraltar is inhabited throughout the year by a group of pilot whales (Globicephala melas), but their spatial distribution varies between Summer and Autumn. In this paper, we have used carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) stable isotope signatures to investigate the differences in diet amongst seasons, sex and stable social units. Skin samples were collected from 56 individually photo-identified pilot whales during Autumn 2005 and Summer 2006. These individuals were genetically sexed and their isotopic signature determined. The level of inter-individual association both within and between stable social units were compared to Euclidean distances between individual isotopes signatures. No differences in either δ¹⁵N or δ¹³C were found according to the sex of individuals, but significant seasonal differences were found in δ¹⁵N, although not in the δ¹³C values. This suggests that pilot whales are resident year round in the Strait, a finding supported by independent photo-identification. The variation in δ¹⁵N could reflect a shift in pilot whale diet through the year, with pilot whales feeding at a higher trophic level in Autumn compared to Summer. This could also represent a change in the diet of pilot whale prey species. The δ¹³C values were significantly different amongst the four stable social units sampled and individual δ¹³C values were significantly related to the level of inter-individual association, while no relationship was found for δ¹⁵N. These results suggest that within the same general area (i.e. the Strait of Gibraltar), there is some level of specialisation in habitat or prey choice between pilot whales social units.     

Contrasting C and N isotope ratios from sperm whale skin and squid between the Gulf of Mexico and Gulf of California: effect of habitat

To investigate feeding variation between populations of an apex oceanic predator, stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) have been compared in skin of female and immature male sperm whales (Physeter macrocephalus) from the northern Gulf of Mexico (GoM) and the Gulf of California (GC). Whale sexes were determined genetically. The δ¹³C and δ¹⁵N values from squid muscle were used from the GC, and from inshore and offshore sites in the GoM. We documented contrastingly lower δ¹³C and δ¹⁵N from whales and squid of the GoM compared with those from the GC. While this difference may be associated with variation in trophic position, geographic variation in biochemical cycling influenced significantly the contrasting isotope values between gulfs. Within the northern GoM, the highly distinct δ¹⁵N values of neritic squid versus mesopelagic squid provide further evidence of habitat specificity in δ¹⁵N.     

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.     

Stable isotopes document winter trophic ecology and maternal investment of adult female southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>) breeding at the Kerguelen Islands

Individual specialisation is widespread and can affect a population’s ecological and evolutionary dynamics. Whether intra-specific niche differences can influence reproductive investment was examined in a marine mammal, the southern elephant seal (Mirounga leonina), whose females were known to forage in two different areas during the austral winter. The study was conducted at Kerguelen Islands (49°21′S, 70°18′E), southern Indian Ocean, in late winter–early spring 2006. Pups were used as proxies of their mothers’ biology and combined information on their weaning mass (a proxy of females’ foraging success and short-term fitness) together with their blood δ¹³C value (a proxy of female foraging zone). First, the use of isotopic signature of pups was validated to study the female foraging ecology during their pre-breeding trip by demonstrating that δ¹³C and δ¹⁵N values of pups and their mothers were positively and linearly correlated. Then, blood samples were taken from a large number of newly-weaned pups, which were also weighed, to provide information at the population level. Estimated δ¹³C values of female seals encompassed a large range of values (from −23.7 to −19.1‰) with an unimodal frequency distribution, suggesting no contrasted foraging areas within the population. No significant relationship was found between pup weaning mass and their carbon signature, indicating no link between female foraging areas and maternal foraging success and investment. Finally, blood δ¹³C and δ¹⁵N values gave new insights into southern elephant seal ecology, suggesting that females mainly foraged north of the Polar Front where they preyed upon myctophid fish in late winter.