Short- and long-term importance of small sharks in the diet of the rare deep-sea shark Dalatias licha

Knowing the trophic ecology of marine predators is essential to develop an understanding of their ecological role in ecosystems. Research conducted on deep-sea and threatened shark species is limited. Here, by combining analyses of individual stomach contents and stable isotope values, we examined the trophic ecology (dietary composition and trophic position) of the kitefin shark Dalatias licha, a deep-sea shark considered as near threatened globally and as data deficient in the Mediterranean Sea. Results revealed the importance of small sharks in the diet of the kitefin shark at short- and long-term scales, although fin-fish, crustaceans and cephalopods were also found. Predation on sharks reveals the high trophic position of the kitefin shark within the food web of the western Mediterranean Sea. Stable isotope values from liver and muscle tissues confirmed our results from stomach content analysis and the high trophic position.     

Long-term changes in the Georges Bank food web: trends in stable isotopic compositions of fish scales

Fish scales from seven species of demersal fish in an archival collection were analyzed for stable isotopic compositions of carbon and nitrogen to study long-term changes in trophic structure of the Georges Bank food web. Nitrogen isotopic compositions are often used to infer trophic level. In the case of haddock, Melanogrammus aeglefinus (Linnaeus), there was a trend towards feeding at 2/3 of one trophic level (2.45% in ¹⁵N) lower in 1987 than in 1929. Values of °¹³C, frequently employed to identify sources of organic carbon to consumers, declined by 1.5 from 1929 to 1960, and then increased again toward the present, suggesting changes in the food web at the level of the primary producers. Superimposed on long-term isotopic trends were short-term variations (1 to 10 yr).To identify potential causes for these isotopic trends, canonical correlation analysis was performed between isotopic data and a suite of environmental and population factors including sea surface temperature, the Greenland Regional Pressure Anomaly (GRPA), the North Atlantic Oscillation (NAO), and the following haddock stock parameters; stock size, fishing mortality, recruitment, and weight-at-age-2 (a measure of growth rate). Isotopic variation was significantly correlated with a combination of environmental and population variables: GRPA, NAO, weight-at-age-2, stock size, and fishing mortality. On the basis of published gut content analyses, the seven fish species were predicted to vary in trophic level (TL) from American plaice, Hippoglossoides platessoides (Fabricius), (TL 2.9) to summer flounder, Paralichthys dentatus (Linnaeus), (TL 4.5), whereas measured °¹⁵N values suggested smaller differences in the trophic levels of these species (less than one TL). Four species showed good agreement between gut-predicted and measured ¹⁵N values, while three species did not. Inclusion of information on ontogenetic dietary shifts in our predictions improved the agreement in some cases but not in other. Differences between stable isotope analysis and gut content analysis in terms of what they measure, i.e., integrated assimilated diet vs short-term ingested diet, respectively, may account for some of the differences in results. Based on our analyses and previous studies, feeding habits of these fish may undergo considerable year-to-year and geographic variation, some of which may have been missed in gut content analyses. To the extent that these fish are representative members of the food web, trophic variation in these fish may indicate more general changes in the food web.     

The role of Greenland sharks (Somniosus microcephalus) in an Arctic ecosystem: assessed via stable isotopes and fatty acids

The Greenland shark (Somniosus microcephalus) is the only shark species known to inhabit ice-covered seas in the North Atlantic, but remains a missing component in most studies of Arctic food webs. In the present study, stable isotopes (SIs) of nitrogen (δ¹⁵N) and carbon (δ¹³C) and fatty acids (FAs) were analyzed to identify the role of Greenland sharks (sampled during June 2008–2009) in Kongsfjorden, a productive fjord on the west coast of Svalbard, Norway (~79ºN, 12–13ºE). The Greenland shark fed at a high trophic position (4.8) based on δ¹⁵N values, and δ¹³C confirmed that most (70 %) of their carbon was derived from phytoplankton-based food chains, which is consistent with a heavy reliance on pelagic teleosts and seals. Greenland sharks from Kongsfjorden had fatty acid profiles in both muscle and plasma (e.g., low 20:1n-9, high 22:5n-3) that suggested a low portion of Greenland halibut (Reinhardtius hippoglossoides) and high proportion of gadoids and seals in their diet compared to Greenland sharks sampled in Cumberland Sound, Canada, during April 2008, which were previously shown to derive much of their energy from Greenland halibut. The high proportions of seal fatty acids in both slow- (muscle) and fast- (plasma) turnover tissues indicate that trophic interactions between Greenland sharks and seals in Kongsfjorden are a common occurrence. Results from the present study suggest that Greenland sharks likely play a unique and significant role in Arctic marine food webs as a top predator of fishes and marine mammals.     

Mercury bioaccumulation and trophic transfer in sympatric snapper species from the Gulf of Mexico.

Consumption of marine fish is a major route of toxic methyl mercury (MeHg) exposure to ocean apex predators and human populations. Here we explore the influence of trophic structure on total mercury (Hg) accumulation in red snapper (RS, Lutjanus campechanus) and gray snapper (GS, Lutjanus griseus) from the coastal Louisiana region of the Gulf of Mexico, west of the Mississippi River. The objectives of this investigation were to: (1) determine the effectiveness of the use of offshore recreational fishing charter boats and marinas as sources of fish samples and (2) compare species differences in Hg bioaccumulation, trophic position, and carbon sources. Our data show that length-normalized Hg concentrations (> or = 97% as MeHg in tissue of both species) were 230% greater in GS in comparison to RS collected from the same general area. Stable C and N isotope signatures (delta15N and delta13C) indicate that GS occupy a slightly higher trophic position (approximately 30% of one trophic position higher) on the Gulf food web in comparison to RS and that GS appear to incorporate higher trophic positioned prey, continually and at smaller sizes. Mercury was strongly correlated with combined delta15N and delta13C in pooled species data, arguing that most of the substantial difference in Hg bioaccumulation between RS and GS can be explained by modest differences in their trophic position and, to a lesser degree, carbon sources, which had low variation and high overlap among species. These observations demonstrate that even minor to moderate differences in trophic position and food habits in sympatric species can create relatively large differences in bioaccumulation regimes and underscores the importance of quantitative characterization of trophic structure in marine MeHg bioaccumulation studies.     

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.     

Identifying trophic variation in a marine suspension feeder: DNA- and stable isotope-based dietary analysis in Mytilus spp.

Accurate field data on trophic interactions for suspension feeders are lacking, and new approaches to dietary analysis are necessary. Polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE) was integrated with stable isotope analysis to examine dietary patterns in suspension-feeding Mytilus spp. from seven spatially discrete locations within a semi-enclosed marine bay (Strangford Lough, Northern Ireland) during June 2009. Results of the two methods were highly correlated, reflecting dietary variation in a similar manner. Variation in PCR-DGGE data was more strongly correlated with the principal environmental gradient (distance from the opening to the Irish Sea), while values of δ ¹³C and δ ¹⁵N became progressively enriched, suggesting a greater dependence on animal tissue and benthic microalgae. Diatoms and crustaceans were the most frequently observed phylotypes identified by sequencing, but specific DNA results provided little support for the trophic trends observed in the stable isotope data. This combined approach offers an increased level of trophic insight for suspension feeders and could be applied to other organisms.     

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.     

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.     

Size-related trophodynamic changes in three target fish species recovering from intensive trawling

The consequences for food-web structure of protecting marine communities from trawling have been little explored. We investigated the Gulf of Castellammare, northwest Sicily, where a trawl ban initiated in 1989 has led to great increases in the biomass of some target species. Using stable isotopes of carbon and nitrogen, our objectives were to see if mean trophic level and omnivory had increased after the ban in three of the most common fishery-target species, namely, the anglerfish Lophius budegassa, Mediterranean hake Merluccius merluccius, and red mullet Mullus barbatus. We compared size data from before and after the fishery closure, but we also compared recently derived data from outside and inside the closed area. In all three species '¹³C and '¹⁵N were found to differ markedly depending on fish size. '¹³C was found to decrease in L. budegassa and Merluccius merluccius but increase in Mullus barbatus with increasing fish length. '¹⁵N increased in all three species, and this was thought to reflect feeding at increasingly higher trophic levels during the animals' lives. Mean length, and therefore '¹⁵N-derived trophic level, increased after the trawl ban only in the anglerfish L. budegassa. Based on '¹³C and '¹⁵N data, none of the species became more or less omnivorous after the ban. After 9 years of no trawling, increases in numerical abundance were not accompanied by substantial size-related trophodynamic shifts in any of the three species of fish studied.     

The tongue-replacing isopod Cymothoa borbonica reduces the growth of largespot pompano Trachinotus botla

Cymothoa borbonica prevalence in the buccal cavity of Trachinotus botla was high, with 45 % of all fish sampled being infected. Smaller fish were more susceptible to infection with no parasites found in fish over 400 mm FL. The detrimental effects of parasite infection on their hosts include basihyal (the bone commonly known as the “tongue”) damage, a loss in buccal cavity volume as a result of female parasite attachment, and a severe impact on host growth. By combining short-term dietary analysis and medium-term stable isotope analysis, there was little evidence to suggest a modification in either the diet or feeding habits of infected fish where infected and uninfected fish occupied the same trophic niche. Inhibited growth in infected fish is hypothesized to be from respiratory distress from long-term oxygen deficiency through buccal obstruction.     

Ecological tracers and at-sea observations document the foraging ecology of southern long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas edwardii</Emphasis>) in Kerguelen waters

The food and feeding ecology of the poorly known southern long-finned pilot whale (Globicephala melas edwardii) was investigated using ecological tracers (muscle and skin δ¹³C and δ¹⁵N, and total mercury, Hg) on individuals from two mass strandings together with at-sea observations of live animals in Kerguelen waters, southern Indian Ocean. Sightings of cetaceans from longliners over 9 years (2003–2012) emphasized the regular occurrence of pilot whales in slope waters surrounding the archipelago. Tissue δ¹³C values (a proxy of consumer foraging habitat) suggest that pilot whales fed in slope waters and in oceanic subantarctic waters over the last months preceding stranding. Tissue δ¹⁵N values and Hg concentration (dietary proxies) indicate a high trophic position (~4.7) for the pilot whales, likely corresponding to a mixed diet of fish and squid (not crustaceans) of undetermined species. Both skin and muscle Hg concentrations were positively and linearly correlated to individual size with no concomitant δ¹⁵N changes, which can be interpreted as a progressive Hg accumulation in tissues of individuals throughout life with no parallel dietary shift. Skin and muscle Hg concentrations were linearly and positively related; hence, Hg skin could be used as a proxy of Hg concentration in muscle (a main Hg reservoir of the body). Kerguelen southern long-finned pilot whales were less Hg contaminated than most pilot whale populations studied so far, thus suggesting that they are not at a high risk to Hg-induced damages in the remote islands of the Southern Ocean.     

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.     

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.     

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.     

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.     

Trophic relationships of hydrothermal vent and non-vent communities in the upper sublittoral and upper bathyal zones off Kueishan Island,Taiwan: a combined morphological,gut content analysis and stable isotope approach

This study used morphological, gut content analysis and carbon- and nitrogen-stable isotope analysis to investigate the trophic structure of upper sublittoral (15–30 m deep) and upper bathyal (200–300 m deep) hydrothermal vents and the adjacent non-vent upper bathyal environment off Kueishan Island. The sublittoral vents host no chemosynthetic fauna, but green and red algae, epibiotic biofilm on crustacean surfaces, and zooplankton form the base of the trophic system. Suspension-feeding sea anemones and the generalist omnivorous vent crab Xenograpsus testudinatus occupy higher trophic levels. The upper bathyal hydrothermal vent is a chemoautotrophic-based system. The vent mussel Bathymodiolus taiwanensis forms a chemosynthetic component of this trophic system. Bacterial biofilm, surface plankton, and algae form the other dietary fractions of the upper bathyal fauna. The vent hermit crab Paragiopagurus ventilatus and the vent crab X. testudinatus are generalist omnivores. The vent-endemic tonguefish Symphurus multimaculatus occupies the top level of the trophic system. The adjacent non-vent upper bathyal region contains decapod crustaceans, which function as either predators or scavengers. The assemblages of X. testudinatus from sublittoral and upper bathyal vents exhibited distinct stable isotope values, suggesting that they feed on different food sources. The upper bathyal Xenograpsus assemblages displayed large variations in their stable isotope values and exhibited an ontogenetic shift in their δ¹³C and δ¹⁵N stable isotope signatures. Some individuals of Xenograpsus exhibited δ¹⁵N values close to those of non-vent species, suggesting that the highly mobile Xenograpsus may transfer energy between the upper bathyal hydrothermal vents and the adjacent non-vent upper bathyal environment.     

Preservation methods alter stable isotope values in gelatinous zooplankton: implications for interpreting trophic ecology

Jellyfish are increasingly topical within studies of marine food webs. Stable isotope analysis represents a valuable technique to unravel the complex trophic role of these long-overlooked species. In other taxa, sample preservation has been shown to alter the isotopic values of species under consideration, potentially leading to misinterpretation of trophic ecology. To identify potential preservation effects in jellyfish, we collected Aurelia aurita from Strangford Lough (54^o22′44.73″N, 5^o32′53.44″W) during May 2009 and processed them using three different methods prior to isotopic analysis (unpreserved, frozen and preserved in ethanol). A distinct preservation effect was found on δ¹⁵N values: furthermore, preservation also influenced the positive allometric relationship between individual size and δ¹⁵N values. Conversely, δ¹³C values remained consistent between the three preservation methods, conflicting with previous findings for other invertebrate, fish and mammalian species. These findings have implications for incorporation of jellyfish into marine food webs and remote sampling regimes where preservation of samples is unavoidable.     

Seasonal variations in the stable carbon and nitrogen isotope ratios (13C/12C and 15N/14N) of primary producers and consumers in a western Mediterranean coastal lagoon

Stable carbon and nitrogen isotope ratios (¹³C/¹²C and ¹⁵N/¹⁴N) of primary producers and consumers were investigated seasonally throughout 1999, in order to describe the food web in a western Mediterranean coastal lagoon (Lake of Sabaudia, central Italy). Particulate organic matter and algal material (seagrass epiphytes and macroalgae) seem to constitute the main food sources for primary consumers (zooplankton and small benthic invertebrates, respectively) throughout the sampling year, while the seagrass Cymodocea nodosa appears to play a negligible trophic role. As regards the ichthyofauna, carbon stable isotopes differentiated between planktivore and benthivore fish species. However, a benthic-pelagic coupling seems to occur, with some fish of higher trophic levels feeding both on benthic and pelagic materials. Analysis of variance showed that the interaction between the three main factors (species2size2season) significantly affects the isotopic composition of fish, suggesting the presence of intra- and inter-specific resource partitioning. Wide seasonal variations in the isotopic composition were observed in organic matter sources, invertebrates and fish, with a general trend towards depleted values in winter and enriched values in summer. The winter depletion of organic matter sources may be due to several environmental factors and seems to be mirrored in the upper trophic levels. Primary producers and invertebrates are known to have shorter time-integrated isotopic signatures than vertebrates, yet fish also exhibited seasonal isotopic differences. We concluded that the examined fish species can assume a new muscle isotopic signature relatively quickly in response to changes in the isotopic composition of their diet and/or diet shifts.     

Sexual segregation in distribution,diet and trophic level of seabirds: insights from stable isotope analysis

Considerable attention has focused on inter- and intraspecific variation in trophic niches of marine predators. Although this has revealed evidence for sexual segregation in distribution in some species, few studies have been able to address sex-related dietary specialisation. Stable isotope analysis of blood cells collected from albatrosses and petrels at South Georgia during chick-rearing indicated a difference in δ¹³C, suggesting that females fed to the north of males, only in two species with male-biased sexual size dimorphism; in no species did sexes differ in trophic level (δ¹⁵N). Based on a wider review, significant differences between sexes in isotope signatures were much more common in seabirds during the pre-laying or breeding than the nonbreeding period, presumably reflecting greater between-sex partitioning of resources when foraging ranges are more constrained and competition is greater. Sex differences, or their absence, were usually consistent across successive stages during the pre-laying and breeding periods, but not necessarily year-round nor between populations. Significant differences in isotope signatures between males and females were extremely rare in monomorphic species, suggesting a link between sexual size dimorphism and segregation in diet or distribution. Among the Southern Ocean albatrosses, sex differences in δ¹³C suggested the underlying mechanism was related to habitat specialisation, whereas in other size-dimorphic taxa (both male- and female-biased), sex differences were more common in δ¹⁵N than δ¹³C and therefore more consistent with size-mediated competitive exclusion or dietary specialisation.     

Matching and mismatching stable isotope (δ13C and δ15N) ratios in fin and muscle tissue among fish species: a critical review

Using non-lethal tissue sampling for stable isotope analysis has become standard in many fields, but not for fishes, despite being desirable when species are rare or protected, when repeated sampling of individuals is required or where removal may bias other analyses. Here, we examine the utility of fish dorsal fin membrane as an alternative to muscle for analyzing δ¹³C and δ¹⁵N ratios in two reef fish species (blue cod Parapercis colias and spotty Notolabrus celidotus) that have differing feeding modes. Both species exhibited evidence of size-based feeding from fin δ¹⁵N values, but not from muscle. Blue cod fin δ¹⁵N increased steadily throughout the sampled size range (213–412 mm fork length), whereas spotty exhibited a distinct ontogenetic diet shift at approximately 120–140 mm fork length after which size-based feeding did not occur. Fin membrane was higher than muscle in δ¹³C in both species and in δ¹⁵N for blue cod, but fin δ¹⁵N was lower than muscle in spotty. The δ¹³C and δ¹⁵N fin–muscle offsets were constant in spotty regardless of size, while in blue cod, δ¹³C was constant with fish size, but δ¹⁵N offsets increased with increasing fish size. Non-lethal sampling utilizing fin tissue can be employed to estimate stable isotope values of muscle in fishes, but it is necessary to assess relationships among tissues and the effects of fish size on isotope values a priori for each species studied. Our data indicated that fin membrane may be a more sensitive tissue than muscle for detecting size-based feeding in some fish species using stable isotopes. A critical literature review revealed inconsistencies in tissue types tested, little understanding of tissue-specific trophic shift or turnover rates, and pseudo-replicated analyses leading to erroneous postulating of 1:1 relationships between tissues.