Using fatty acids as dietary tracers in seabird trophic ecology: theory, application and limitations

Analysis of fatty acids (FAs) is an increasingly utilized tool in studies of trophic ecology in marine ecosystems. This powerful technique has proved useful in delineating spatial and temporal variability in diets, identifying the consumption of key species, and providing quantitative estimates of diet composition. Although consumer FA signatures are undeniably influenced by diet, they can also be affected by other factors including life-history stage, diet quality, and physiological state. Here, we review how FAs are assimilated, deposited, and metabolized in birds, and the implications of these processes on the various tissues commonly sampled for FA analyses. We then examine the assumptions underlying FA signature analysis when used in studies of seabird trophic ecology and propose a direction for future laboratory experiments that are needed to refine the approach. The correct interpretation of FA data relies on accounting for factors that alter predator FA metabolism and controlling for variability in the lipid content and FA composition of prey. Efforts should also be made to incorporate uncertainty associated with predator metabolism into models designed for quantitative diet estimation.     

18S rRNA metabarcoding diet analysis of a predatory fish community across seasonal changes in prey availability

Predator–prey relationships are important ecological interactions, affecting biotic community composition and energy flow through a system, and are of interest to ecologists and managers. Morphological diet analysis has been the primary method used to quantify the diets of predators, but emerging molecular techniques using genetic data can provide more accurate estimates of relative diet composition. This study used sequences from the 18S V9 rRNA barcoding region to identify prey items in the gastrointestinal (GI) tracts of predatory fishes. Predator GI samples were taken from the Black River, Cheboygan Co., MI, USA (n = 367 samples, 12 predator species) during periods of high prey availability, including the larval stage of regionally threatened lake sturgeon (Acipenser fulvescens Rafinesque 1817) in late May/early June of 2015 and of relatively lower prey availability in early July of 2015. DNA was extracted and sequenced from 355 samples (96.7%), and prey DNA was identified in 286 of the 355 samples (80.6%). Prey were grouped into 33 ecologically significant taxonomic groups based on the lowest taxonomic level sequences that could be identified using sequences available on GenBank. Changes in the makeup of diet composition, dietary overlap, and predator preference were analyzed comparing the periods of high and low prey abundance. Some predator species exhibited significant seasonal changes in diet composition. Dietary overlap was slightly but significantly higher during the period of high prey abundance; however, there was little change in predator preference. This suggests that change in prey availability was the driving factor in changing predator diet composition and dietary overlap. This study demonstrates the utility of molecular diet analysis and how temporal variability in community composition adds complexity to predator–prey interactions.     

The effects of diet and caloric restriction on adipose tissue fatty acid signatures of tufted puffin (<Emphasis Type="Italic">Fratercula cirrhata</Emphasis>) nestlings

Fatty acid (FA) signature analysis is a powerful tool to investigate foraging ecology and food web dynamics in marine ecosystems. However, use of FA signatures to qualitatively or quantitatively infer diets is potentially complicated by effects of nutritional state on lipid metabolism. Estimation of diets using the quantitative fatty acid signature analysis (QFASA) model requires the use of calibration coefficients to account for predator metabolism of individual FAs. We conducted a captive feeding experiment to determine the effects of a 50% reduction in food intake on growth rate and adipose tissue FA signatures of tufted puffin (Fratercula cirrhata) nestlings, a species that routinely experiences food restriction during growth. FA signatures of chicks fed low- and high-calorie diets both exhibited a change in composition in response to the dietary shift with the direction of change in the composition of individual FAs matching the direction of change in the dietary FAs. Despite a growth rate in the restricted nestlings that was 38% of those in the well-fed group, rates of FA turnover were not different between high and low-calorie treatments, and turnover was close to, but not entirely complete, after 27 days on both high-calorie and restricted diets. FA signatures of tufted puffin nestlings were significantly affected by caloric restriction, but these effects were much less pronounced than those of dietary turnover, and calibration coefficients of puffins fed low and high-calorie diets were highly correlated. Our results demonstrate that changes in physiological state can affect FA metabolism, but future research is required to better understand whether the size of these effects is sufficient to substantially alter diet estimation using the QFASA model.     

Physiological constraints and the influence of diet on fatty acids in the yolk of gentoo penguins, Pygoscelis papua

Avian yolk fatty acids (FA) composition is influenced by two main factors: maternal diet and genetic factors that regulate FA metabolism. However, due to embryonic developmental requirements, yolk FA are thought to be physiologically constrained and less useful for dietary and trophic studies. We assessed the relative contributions of diet and physiological constraints in determining the yolk FA composition of a marine bird, the gentoo penguin (Pygoscelis papua) by comparing FA signatures of yolks and prey between a captive, controlled- feeding experiment and a wild population. Captive and wild yolk FA signatures differed even though both groups' yolk lipids were composed primarily of three FA (16:0, 18:0 and 18:1n-9). Differences were due to FA occurring in relatively low abundance, but which mirrored differences in the FA composition of diets. However, yolk FA signatures were correlated across three penguin species suggesting that common developmental constraints can be relatively more important than species-specific differences in diet or egg-laying physiology. While yolk FA are constrained, several minor components of yolk FA are reflective of diets and the calibration coefficients resulting from this study have the potential to be incorporated into predictive models and allow for quantitative dietary and trophic studies using FA analysis of penguin egg yolks.     

Trophic variability of Arctic fishes in the Canadian Beaufort Sea: a fatty acids and stable isotopes approach

Trophic ecology of most demersal Arctic fishes remains one of the major knowledge gaps for understanding food web dynamics and connectivity among ecosystems. In this study, fatty acids (FA) and stable isotopes (SI) were used to study the feeding ecology of seven species (n = 106) of the most abundant benthic fishes (eelpouts, sculpins and agonids) in the Canadian Beaufort Sea from shallow (20–75 m), slope (200–350 m) and deep (500–1000 m) habitats. Both FA and SI results revealed among- and within-species variability in diet composition. Correspondence analysis of FA signatures identified high within-species variability in diet, resulting in high overlap among species. Calanus-derived FA were present in all species (Calanus markers up to 13 % of total FA) and were particularly important in Ribbed Sculpin, Adolf’s and Longear Eelpout collected in deep habitats, suggesting a strong contribution of pelagic-derived FA to benthic fish communities. Incorporation of this signal in the benthos may result from either direct consumption of deep overwintering copepods (i.e., off-bottom feeding) or through detrital accumulation in benthic invertebrate prey. Mean SI values differed among species and indicated that a large range of trophic positions (δ¹⁵N varied from 14.09 to 17.71 ‰ for Canadian Eelpout and Adolf’s Eelpout, respectively) and carbon dietary sources are preyed upon (δ¹³C range from ?21.13 to ?23.85 ‰ for Longear Eelpout and Ribbed Sculpin, respectively). SI analyses suggested that most species examined were low- to mid-trophic generalist benthic carnivores, with the exception of Ribbed Sculpin, which was a low-trophic pelagic predator.     

Variation in the fatty acid composition of blubber in Cape fur seals (<Emphasis Type="Italic">Arctocephalus pusillus pusillus</Emphasis>) and the implications for dietary interpretation

Analysis of the fatty acid (FA) composition of blubber is a valuable tool in interpreting the diet of marine mammals. This technique is based on the principle that particular FA present in prey can be incorporated largely untransformed into predator adipose tissue stores, thereby providing biochemical signatures with which to identify prey species. Several studies of phocid seals and cetaceans have documented vertical stratification in the FA composition of blubber such that inferences about diet may vary greatly depending on the layer of the blubber that is analysed. It is not known whether blubber in otariid seals (fur seals and sea lions) also displays vertical stratification in FA composition. Furthermore, it is not known whether the FA composition of blubber is uniform in these species. In the present study, the vertical and regional variation in FA composition of blubber was investigated in seven adult female Cape fur seals (Arctocephalus pusillus pusillus). The proportion of monounsaturated fatty acids (MUFA) was greater in the outer (43.6±1.3%) than inner portion (40.9±1.2%; t₂₀=5.59, P<0.001) whereas the proportions were greater in the inner than outer portions for saturated fatty acids (23.6±0.5% and 21.9±0.6%, respectively, t₂₀ = 5.31, P<0.001) and polyunsaturated fatty acids (PUFA, 35.5±0.7% and 34.5±0.7%, respectively, t₂₀ = 3.81, P < 0.001). There was an inverse relationship between MUFA and PUFA in the blubber, independent of sampling location. In addition, with the exception of the inner portion from non-lactating females, blubber from the mammary area had the highest proportions of 18:19c and total MUFA, followed by blubber from the rump and neck, suggesting that the deposition and mobilisation of blubber lipids may not be uniform around the body in otariid seals. These results support the need for blubber tissue to be sampled from the same site on animals, and to the full depth of the blubber layer, to minimise variation in FA profiles that could occur if different sites and depths were sampled. Such standardisation of sampling will further aid in interpreting diet in otariid seals using the FA Signature Analysis approach.     

Fatty acid signatures of stomach oil and adipose tissue of northern fulmars (<Emphasis Type="Italic">Fulmarus glacialis</Emphasis>) in Alaska: implications for diet analysis of Procellariiform birds

Procellariiforms are unique among seabirds in storing dietary lipids in both adipose tissue and stomach oil. Thus, both lipid sources are potentially useful for trophic studies using fatty acid (FA) signatures. However, little is known about the relationship between FA signatures in stomach oil and adipose tissue of individuals or whether these signatures provide similar information about diet and physiology. We compared the FA composition of stomach oil and adipose tissue biopsies of individual northern fulmars (N = 101) breeding at three major colonies in Alaska. Fatty acid signatures differed significantly between the two lipid sources, reflecting differences in dietary time scales, metabolic processing, or both. However, these signatures exhibited a relatively consistent relationship between individuals, such that the two lipid sources provided a similar ability to distinguish foraging differences among individuals and colonies. Our results, including the exclusive presence of dietary wax esters in stomach oil but not adipose tissue, are consistent with the notion that stomach oil FA signatures represent lipids retained from prey consumed during recent foraging and reflect little metabolic processing, whereas adipose tissue FA signatures represent a longer-term integration of dietary intake. Our study illustrates the potential for elucidating short- versus longer-term diet information in Procellariiform birds using different lipid sources.     

Mechanisms of prey selection by predaceous stoneflies: roles of prey morphology,behavior and predator hunger

Summary Laboratory feeding experiments using Hesperoperla pacifica (Banks), Perlidae, and Megarcys signata (Hagen), Perlodidae, as predators and Baetis tricaudatus Dodds and Ephemerella altana Allen as prey indicate a strong effect of prey morphology and mobility and predator hunger on prey selection by stoneflies. Knowledge of both dietary composition and feeding behavior was necessary to fully understand prey selection by these stoneflies.Fasted stoneflies presented with live prey ate more E. altana while satiated stoneflies ate approximately equal numbers of the two mayfly species. This pattern of dietary composition was the result of a reduction of attack frequency on the slower swimming E. altana with predator satiation and a continued high attack rate on B. tricaudatus regardless of recent feeding history. In contrast, fasted H. pacifica fed fresh frozen mayflies ate more B. tricaudatus indicating the importance of differences in prey mobility in controlling dietary composition.The high degree of similarity in patterns of feeding and mechanisms underlying those patterns for H. pacifica and M. signata suggest that they may be using similar rules for choosing mayfly prey and we suggest that mayfly prey are ranked by stoneflies on the basis of handling times. A general mechanistic model for stoneflies feeding on mayflies is presented and discussed.     

Validating quantitative fatty acid signature analysis to estimate diets of spectacled and Steller’s eiders (Somateria fischeri and Polysticta stelleri)

Fatty acid (FA) signature analysis has been used to study foraging ecology and food webs in marine ecosystems. This powerful method provides information about diets over an extended time period (e.g., 2–4 weeks), rather than just the most recent meal as with most traditional approaches. Using consumer FA signatures, along with a comprehensive database of diet FA signatures, and accounting for consumer FA metabolism, it is possible to estimate the proportions of diet items in the consumer’s diet using quantitative FA signature analysis (QFASA). However, before applying QFASA to free-ranging populations, ideally, controlled feeding studies are performed to determine FA deposition and turnover characteristics. We conducted feeding experiments to validate QFASA in captive spectacled eiders (Somateria fischeri) and Steller’s eiders (Polysticta stelleri) as a minimally invasive method for studying the diets of these threatened species. We determined FA deposition in eider adipose tissue relative to long-term diet, and developed calibration coefficients (CCs) to account for eider lipid metabolism. Using these CCs with subsequent diet trials, QFASA accurately indicated diet and diet switches. QFASA estimates also indicated that turnover of dietary FAs was not complete by 21 or 29 days, and confirmed that diets could be estimated over an extended period of >29 days. Thus, our understanding of diet can be backtracked to more than a month in captive feeding eiders. We conclude that applying QFASA techniques to eiders and other birds in the wild has the potential to provide valuable information about their diets at various life history stages.     

Fatty‐acid profiles of juvenile lake trout reflect experimental diets consisting of natural prey

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Fatty acid composition of sprat (<Emphasis Type="Italic">Sprattus sprattus</Emphasis>) and herring (<Emphasis Type="Italic">Clupea harengus</Emphasis>) in the Baltic Sea as potential prey for salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>)

Sprat (Sprattus sprattus) and small herring (Clupea harengus) are the dominant prey fish of Atlantic salmon (Salmo salar) in the Baltic Sea. If the fatty acid (FA) proportions of sprat and herring differ, the dietary history of ascending salmon could be determined from their FA profiles. Therefore, we investigated the FA composition of several age groups of whole sprat and small herring, caught from the three main feeding areas of salmon in autumn and spring. Oleic acid (18:1n-9) was the most prevalent FA in sprat and characteristic of this species. In herring, palmitic acid (16:0) was the most common FA, but herring lipid was characterized by n-6 polyunsaturated FAs, and moreover, by palmitoleic acid (16:1n-7) and vaccenic acid (18:1n-7). Due to the higher lipid content of sprat, the concentrations of all other FAs, excluding these, were higher in sprat than in herring. The concentration of docosahexaenoic acid (DHA, 22:6n-3) increased with an increase in the lipid content and was consequently highest in the youngest specimens, being in young sprat almost double that of young herring, and 2.6 times higher in the sprat biomass than in that of herring. As a result of a decrease in the DHA concentration with age, the ratio thiamine/DHA increased with respect to age in both species, and was lower in sprat than in herring. It is concluded that an abundance of DHA in the diet of salmon most likely increases oxidative stress because of the susceptibility of DHA to peroxidation, and thus decreases thiamine resources of fasting, prespawning salmon. Because the FA composition of sprat and herring differs, and the relative abundancies of prey fish differ between the feeding areas of salmon, the feeding area of ascending salmon can most probably be derived by comparing their FA profiles.     

Ontogenetic niche shifts and resource partitioning in a subarctic piscivore fish guild

The feeding ecology of three piscivorous fish species (perch (Perca fluviatilis), pike (Esox lucius) and burbot (Lota lota)), was studied in the subarctic Pasvik watercourse (69 °N), northern Norway and Russia. All three species primarily occupied the benthic habitats in the watercourse. Perch and burbot exhibited distinct ontogenetic niche shifts in food resource use, perch changing from a dominance of zooplankton to zoobenthos to fish, and burbot from zoobenthos to fish. Fish prey dominated the diet of all the investigated size-classes of pike, but small-sized pike (<20 cm) were not represented in the sample. Fish prey size was positively related to predator size in all three species. Whitefish (Coregonus lavaretus) was the dominant prey of pike and large-sized burbot and perch. Nine-spined sticklebacks (Pungitius pungitius) was also an important prey and appeared to be a dietary stepping-stone enhancing the transition from invertebrate feeding to consumption of large-sized whitefish prey for all three predators. A cluster analysis separated the different size groups of the three predator species into five functional feeding groups, most of them containing two or all three species. Within these feeding groups, and especially among the piscivorous size-classes, there was a strong and significant interspecific overlap in prey selection, and the dietary similarities between the species were in general much larger than the intraspecific similarities between ontogenetic stages. All three piscivorous species are important top predators in the aquatic food web of the watercourse, and their ontogenetic diet shifts and resource partitioning patterns generate a substantial food web complexity in this subarctic ecosystem.     

Prey composition and ontogenetic shift in coastal populations of longnose gar Lepisosteus osseus

Longnose gar Lepisosteus osseus were collected from May 2012 to July 2013 in the Charleston Harbor and Winyah Bay estuaries (SC, U.S.A.). This study examined trends in stomach fullness, described major prey components and their importance in the diet of L. osseus, compared stomach content‐based trophic level estimates with the stable‐isotope‐based proxy: δ¹⁵N and tested for the occurrence of an ontogenetic diet shift using stomach content analysis and stable C and N isotopes (δ¹³C and δ¹⁵N). Dominant prey families were Clupeidae, Sciaenidae, Penaeidae, Fundulidae and Mugilidae, with the highest consumption rates in autumn. Trophic levels calculated using stomach contents did not correspond to δ¹⁵N (P > 0·05). Stomach contents and stable‐isotope signatures indicate ontogenetic prey composition shifts from low trophic level benthic prey (fundulids) to higher trophic level pelagic prey (clupeids) as the fish grow between 400 and 600 mm in standard length. Due to their biomass, abundance and top predator status, L. osseus play a significant ecological role in the estuarine community composition, although this effect has often been overlooked by past researchers and should be considered in future estuarine community studies.     

Chylomicrons Promote Intestinal Absorption and Systemic Dissemination of Dietary Antigen (Ovalbumin) in Mice

Background

A small fraction of dietary protein survives enzymatic degradation and is absorbed in potentially antigenic form. This can trigger inflammatory responses in patients with celiac disease or food allergies, but typically induces systemic immunological tolerance (oral tolerance). At present it is not clear how dietary antigens are absorbed. Most food staples, including those with common antigens such as peanuts, eggs, and milk, contain long-chain triglycerides (LCT), which stimulate mesenteric lymph flux and postprandial transport of chylomicrons through mesenteric lymph nodes (MLN) and blood. Most dietary antigens, like ovalbumin (OVA), are emulsifiers, predicting affinity for chylomicrons. We hypothesized that chylomicron formation promotes intestinal absorption and systemic dissemination of dietary antigens.

Methodology/Principal Findings

Absorption of OVA into MLN and blood was significantly enhanced when OVA was gavaged into fasted mice together with LCT compared with medium-chain triglycerides (MCT), which do not stimulate chylomicron formation. The effect of LCT was blocked by the addition of an inhibitor of chylomicron secretion, Pluronic L-81. Adoptively transferred OVA-specific DO11.10 T-cells proliferated more extensively in peripheral lymph nodes when OVA was gavaged with LCT than with MCT or LCT plus Pluronic L-81, suggesting that dietary OVA is systemically disseminated. Most dietary OVA in plasma was associated with chylomicrons, suggesting that these particles mediate systemic antigen dissemination. Intestinal-epithelial CaCo-2 cells secreted more cell-associated, exogenous OVA when stimulated with oleic-acid than with butyric acid, and the secreted OVA appeared to be associated with chylomicrons.

Conclusions/Significance

Postprandial chylomicron formation profoundly affects absorption and systemic dissemination of dietary antigens. The fat content of a meal may affect immune responses to dietary antigens by modulating antigen absorption and transport.     

Carboxylesterase1/Esterase-x Regulates Chylomicron Production in Mice

Elevated postprandial plasma triacylglycerol (TG) concentrations are commonly associated with obesity and the risk of cardiovascular disease. Dietary fat contributes to this condition through the production of chylomicrons. Carboxylesterases have been mainly studied for their role in drug metabolism, but recently they have been shown to participate in lipid metabolism; however, their role in intestinal lipid metabolism is unknown. Carboxylesterase1/esterase-x (Ces1/Es-x) deficient mice become obese, hyperlipidemic and develop hepatic steatosis even on standard chow diet. Here, we aimed to explore the role of Ces1/Es-x in intestinal lipid metabolism. Six-month old wild-type and Ces1/Es-x deficient mice were maintained on chow diet and intestinal lipid metabolism and plasma chylomicron clearance were analyzed. Along the intestine Ces1/Es-x protein is expressed only in proximal jejunum. Ablation of Ces1/Es-x expression results in postprandial hyperlipidemia due to increased secretion of chylomicrons. The secreted chylomicrons have aberrant protein composition, which results in their reduced clearance. In conclusion, Ces1/Es-x participates in the regulation of chylomicron assembly and secretion. Ces1/Es-x might act as a lipid sensor in enterocytes regulating chylomicron secretion rate. Ces1/Es-x might represent an attractive pharmacological target for the treatment of lipid abnormalities associated with obesity, insulin resistance and fatty liver disease.     

Fatty acid composition of the blubber in white whales (Delphinapterus leucas)

Fatty acid (FA) composition of the blubber in free-ranging white whales (Delphinapterus leucas) from Svalbard's waters was determined and compared with the fatty acid composition of potential prey species in an attempt to assess diet. This methodology is based on the common assumption that unique arrays of FAs found within groups of organisms are transferred, largely unaltered, up marine food chains and thus may be useful for assessment of diet composition. Complete-column blubber biopsies were sampled from white whales (n=7) during the summers of 1996 and 1997. All captured animals were adult males. FAs were extracted from 2–4 replicates taken from an area about 10 cm in front of the mid-dorsal ridge. FA data from a total of 12 potential prey species from the Svalbard area were compared to the white-whale blubber samples. Twenty-two FAs were consistently found in relative amounts >0.5% of the total FA composition in white whales. These FAs accounted for 94–96% of the total FAs present. The blubber was composed almost entirely of triacylglycerols. The major saturated FAs were 14:0 and 16:0; 16:1(n-7), 18:1(n-9) and 20:1(n-9) were the major monounsaturated FAs and 20:5(n-3) and 22:6(n-3) were the major polyunsaturated FAs. Sixteen of the 22 FAs consistently found in the white-whale blubber were also found in considerable amounts (>0.5% of total FAs) in most of the potential species. Principal Component Analysis run on these 16 FAs suggests that polar cod (Boreogadus saida) had the most similar FA composition to the white-whale blubber, followed by capelin (Mallotus villosus), the copepod Calanus hyperboreus and the shrimp Pandalus borealis. Accepted: 27 November 1999     

A new approach to the solution of the linear mixing model for a single isotope: application to the case of an opportunistic predator

Mixing models are used to determine diets where the number of prey items are greater than one, however, the limitation of the linear mixing method is the lack of a unique solution when the number of potential sources is greater than the number (n) of isotopic signatures +1. Using the IsoSource program all possible combinations of each source contribution (0–100%) in preselected small increments can be examined and a range of values produced for each sample analysed. We propose the use of a Moore Penrose (M-P) pseudoinverse, which involves the inverse of a 2×2 matrix. This is easily generalized to the case of a single isotope with (p) prey sources and produces a specific solution. The Antarctic leopard seal (Hydrurga leptonyx) was used as a model species to test this method. This seal is an opportunistic predator, which preys on a wide range of species including seals, penguins, fish and krill. The M-P method was used to determine the contribution to diet from each of the four prey types based on blood and fur samples collected over three consecutive austral summers. The advantage of the M-P method was the production of a vector of fractions f for each predator isotopic value, allowing us to identify the relative variation in dietary proportions. Comparison of the calculated fractions from this method with means from IsoSource allowed confidence in the new approach for the case of a single isotope, N.     

Resource pulses affect prey selection and reduce dietary diversity of dingoes in arid Australia

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Resource partitioning by sympatric pagophilic seals in Alaska: monitoring effects of climate variation with fatty acids

Four species of pagophilic phocid seals in the Western Arctic—bearded (Erignathus barbatus), ringed (Phoca hispida), ribbon (Phoca fasciata), and spotted (Phoca largha)—are particularly vulnerable to arctic warming trends. Documenting diet composition over time is one way in which the flexibility of a species, in the face of broad ecosystem changes, can be measured. Because the fatty acid (FA) composition of depot lipids has long been known to reflect diet, we analysed the FA composition of blubber samples collected from bearded (n = 30), ringed (n = 15), ribbon (n = 32) and spotted seals (n = 24). All animals were taken near Little Diomede Island in May and June 2003, providing a unique opportunity to study interspecies differences with minimal effects of spatial and temporal variation. Bearded seal FA composition was significantly different from all other seals (P < 0.001 in all cases), ringed seal FA composition was significantly different from that of both ribbon and spotted seals (P < 0.001), but ribbon and spotted seal FA compositions could not be distinguished from each other. Overall, the blubber FA compositions of ribbon, ringed and spotted seals implied diets dominated by pelagic prey. Inferences we made about current diets of bearded and ringed seals suggest that certain prey important to them in this area in the 1970s remain important today, despite notable changes in the ecosystem. We believe that blubber FA analysis will be a useful tool in assessing the response of ice-associated pinnipeds and their ecosystem to changes associated with arctic warming.     

Ontogenetic dietary shifts and feeding ecology of the rasptail skate Raja velezi and the brown smoothhound shark Mustelus henlei along the Pacific coast of Costa Rica,Central America

Stomachs from 511 Raja velezi and 340 Mustelus henlei captured as by‐catch in the commercial trawling fishery (2010–2012) were analysed to examine diet composition, ontogenetic shifts and degree of dietary overlap between species life stages in the Pacific Ocean of Costa Rica. Shrimps were the most important prey categories in the diet of R. velezi, while teleosts and cephalopods dominated the diet of M. henlei. Diet comparisons between different stages of R. velezi and M. henlei revealed clear ontogenetic dietary shifts: crustaceans (mainly shrimps, crabs and stomatopods) dominated the diet of immature individuals, and adults had a higher proportion of teleosts. The results suggest that R. velezi is an epibenthic predator that specializes in shrimps during early life stages, and to a lesser extent, teleosts as it matures, while M. henlei is an opportunistic predator with a highly diverse diet consisting of teleosts, cephalopods, shrimps and stomatopods. This study also found little evidence of dietary overlap between species or life stages and suggests that intra‐ and interspecific competition between R. velezi and M. henlei may be reduced by: (1) diet specialization in immature stages of R. velezi, (2) ontogenetic dietary shifts between immature and mature individuals, (3) prey‐size selectivity in larger individuals of R. velezi and (4) differences in depth utilization in overlapping geographical regions.