New insights into the trophic ecology of the scalloped hammerhead shark,Sphyrna lewini,in the eastern tropical Pacific Ocean

Scalloped hammerhead shark, Sphyrna lewini, is considered a generalist predator, showing individual specializations and different trophic levels. This species inhabits both oceanic islands and coastal nursery areas in the eastern tropical Pacific (ETP) Ocean. Existing trophic ecology studies on S. lewini have been focused on stomach contents. This study provides new insights into its foraging preferences, habitat use, and trophic position, using stable isotope analysis (SIA) of muscle tissue. Stable isotope signatures of δ¹³C and δ¹⁵N were determined in S. lewini muscles (n?=?29) from the Ecuadorian Pacific in 2013. Trophic position (TP), isotopic niches, and overlap of S. lewini were estimated by sex, age, and maturity stages to infer their dietary habits throughout life stages. SIA revealed complex movement patterns related to sex and age classes of S. lewini, highlighting high degrees of dietary plasticity and habitat use, with a stronger relation to coastal regions than previously reported. This study provides crucial information regarding essential areas and the related migration behavior of S. lewini, with important implications for their conservation and management in the ETP.

     

Investigations of Δ 14C, δ 13C, and δ 15N in vertebrae of white shark (Carcharodon carcharias) from the eastern North Pacific Ocean

The white shark, Carcharodon carcharias, has a complex life history that is characterized by large scale movements and a highly variable diet. Estimates of age and growth for the white shark from the eastern North Pacific Ocean indicate they have a slow growth rate and a relatively high longevity. Age, growth, and longevity estimates useful for stock assessment and fishery models, however, require some form of validation. By counting vertebral growth band pairs, ages can be estimated, but because not all sharks deposit annual growth bands and many are not easily discernable, it is necessary to validate growth band periodicity with an independent method. Radiocarbon (¹⁴C) age validation uses the discrete ¹⁴C signal produced from thermonuclear testing in the 1950s and 1960s that is retained in skeletal structures as a time-specific marker. Growth band pairs in vertebrae, estimated as annual and spanning the 1930s to 1990s, were analyzed for Δ¹⁴C and stable carbon and nitrogen isotopes (δ¹³C and δ¹⁵N). The aim of this study was to evaluate the utility of ¹⁴C age validation for a wide-ranging species with a complex life history and to use stable isotope measurements in vertebrae as a means of resolving complexity introduced into the ¹⁴C chronology by ontogenetic shifts in diet and habitat. Stable isotopes provided useful trophic position information; however, validation of age estimates was confounded by what may have been some combination of the dietary source of carbon to the vertebrae, large-scale movement patterns, and steep ¹⁴C gradients with depth in the eastern North Pacific Ocean.     

Stable-isotope analysis of a deep-sea benthic-fish assemblage: evidence of an enriched benthic food web

In this study, fishes and invertebrates collected from the continental slope (1000 m) of the eastern North Pacific Ocean were analysed using stable-isotope analysis (SIA). Resulting trophic positions (T(P) ) were compared to known diets and habitats from the literature. Dual isotope plots indicated that most species groups (invertebrates and fishes) sorted as expected along the carbon and nitrogen axes, with less intraspecific variability than interspecific variability. Results also indicated an isotopically distinct benthic and pelagic food web, as the benthic food web was more enriched in both nitrogen and carbon isotopes. Trophic positions from SIA supported this finding, resulting in the assignment of fishes to different trophic positions from those expected based on published dietary information. These differences can be explained largely by the habitat of the prey and the percentage of the diet that was scavenged. A mixing model estimated dietary contributions of prey similar to those of the known diet of Bathyraja trachura from stomach-content analysis (SCA). Linear regressions indicated that trophic positions calculated from SIA and SCA, when plotted against B. trachura total length for 32 individuals, exhibited similar variation and patterns. Only the T(P) from SCA yielded significant results (stomach content: P < 0·05, stable isotope: P > 0·05).     

Evidence for dynamic resource partitioning between two sympatric reef shark species within the British Indian Ocean Territory

Stable-isotope analyses (δ¹³C, δ¹⁵N and δ³⁴S) of multiple tissues (fin, muscle, red blood cells and plasma), revealed ontogenetic shifts in resource use by grey reef sharks Carcharhinus amblyrhynchos and resource partitioning with silvertip sharks Carcharhinus albimarginatus within the British Indian Ocean Territory marine protected area (MPA). Resource partitioning varied temporally, with C. albimarginatus feeding on more pelagic prey during October to January, potentially attributable to an influx of pelagic prey from outside the MPA at that time. Reef sharks may therefore be affected by processes outside an MPA, even if the sharks do not leave the MPA.     

Using stable isotope analysis to understand the migration and trophic ecology of northeastern Pacific white sharks (Carcharodon carcharias)

The white shark (Carcharodon carcharias) is a wide-ranging apex predator in the northeastern Pacific (NEP). Electronic tagging has demonstrated that white sharks exhibit a regular migratory pattern, occurring at coastal sites during the late summer, autumn and early winter and moving offshore to oceanic habitats during the remainder of the year, although the purpose of these migrations remains unclear. The purpose of this study was to use stable isotope analysis (SIA) to provide insight into the trophic ecology and migratory behaviors of white sharks in the NEP. Between 2006 and 2009, 53 white sharks were biopsied in central California to obtain dermal and muscle tissues, which were analyzed for stable isotope values of carbon (δ(13)C) and nitrogen (δ(15)N). We developed a mixing model that directly incorporates movement data and tissue incorporation (turnover) rates to better estimate the relative importance of different focal areas to white shark diet and elucidate their migratory behavior. Mixing model results for muscle showed a relatively equal dietary contribution from coastal and offshore regions, indicating that white sharks forage in both areas. However, model results indicated that sharks foraged at a higher relative rate in coastal habitats. There was a negative relationship between shark length and muscle δ(13)C and δ(15)N values, which may indicate ontogenetic changes in habitat use related to onset of maturity. The isotopic composition of dermal tissue was consistent with a more rapid incorporation rate than muscle and may represent more recent foraging. Low offshore consumption rates suggest that it is unlikely that foraging is the primary purpose of the offshore migrations. These results demonstrate how SIA can provide insight into the trophic ecology and migratory behavior of marine predators, especially when coupled with electronic tagging data.     

New records for Dinemoura ferox (Copepoda: Siphonostomatoida: Pandaridae) from Pacific sleeper sharks captured in waters off eastern Taiwan

Four of the 27 Pacific sleeper sharks (Somniosus pacificus Bigelow and Schroeder) captured in the western North Pacific Ocean off eastern Taiwan between 19 March and 18 May 2002 hosted the parasitic copepod Dinemoura ferox (Kr?yer, 1838) on their body surface including the fins. This report documents a new host record as well as a new ocean record for D. ferox, which until now has only been reported from the benthopelagic sharks, Somniosus microcephalus (Bloch and Schneider) and Centrophorus squamosus (Bonnaterre), occurring in the north Atlantic Ocean off Greenland and Iceland.     

Stable carbon and nitrogen isotopes in multiple tissues of wild and captive harbor seals (Phoca vitulina) off the California coast

Stable carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) of serum, red blood cells (RBC), muscle, and blubber were measured in captive and wild northeast Pacific harbor seals (Phoca vitulina richardii) at three coastal California sites (San Francisco Bay, Tomales Bay, and Channel Islands). Trophic discrimination factors (Δ_{Tissue‐Diet}) were calculated for captive seals and then applied in wild counterparts in each habitat to estimate trophic position and feeding behavior. Trophic discrimination factors for δ¹⁵N of serum (+3.8‰), lipid‐extracted muscle (+1.6‰), and lipid‐blubber (+6.5‰) are proposed to determine trophic position. An offset between RBC and serum of +0.3‰ for δ¹³C and ?0.6‰ for δ¹⁵N was observed, which is consistent with previous research. Specifically, weaner seals (<1 yr) had large offsets, suggesting strong trophic position shifts during this life stage. Isotopic values indicated an average trophic position of 3.6 at both San Francisco Bay and Tomales Bay and 4.2 at Channel Islands. Isotopic means were strongly dependent on age class and also suggested that mean diet composition varies considerably between all locations. Together, these data indicate that isotopic composition of blood fractions can be an effective approach to estimate trophic position and dietary behavior in wild pinnipeds.     

Stable isotope analysis reveals ontogenetic feeding shifts in Pacific blue marlin (Makaira nigricans) off eastern Taiwan

To gain a better understanding of the trophic ecology of Pacific blue marlin Makaira nigricans off eastern Taiwan, nitrogen and carbon stable isotopes (δ¹⁵N and δ¹³C) and Bayesian mixing models were used to explore trophic dynamics and potential ontogenetic feeding shifts across M. nigricans of different size classes. Makaira nigricans samples from east of Taiwan (n = 213) and Palau (n = 37), as well as their prey (n = 70), were collected during 2012 and 2013. Results indicated increases in δ¹⁵N with size, with values of larger size classes (> 200 cm eye-to-fork length; L_EF) significantly higher than those < 200 cm L_EF. Values of δ¹³C were negatively correlated with size. Makaira nigricans > 200 cm L_EF had the highest estimated trophic position (4.44) and also exhibited ontogenetic changes in trophic position. Large M. nigricans fed more on dolphinfish Coryphaena hippurus and hairtail Trichiurus lepturus, while smaller M. nigricans consumed smaller forage fish (e.g., moonfish Mene maculata) and cephalopods. These changes may relate to greater swimming speeds and vertical habitat use in larger M. nigricans, allowing capture and consumption of larger prey items at higher trophic positions. The high trophic level of M. nigricans east of Taiwan confirms its important role as an apex predator in marine food webs and how ecological role changes with size.     

Questioning assumptions of trophic behavior in a broadly ranging marine predator guild

We evaluated whether existing assumptions regarding the trophic ecology of a poorly‐studied predator guild, northwest (NW) Atlantic skates (family: Rajidae), were supported across broad geographic scales. Four hypotheses were tested using carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope values as a proxy for foraging behavior: 1) species exhibit ontogenetic shifts in habitat and thus display a shift in ¹³C with differential use of the continental shelf; 2) species exhibit ontogenetic prey shifts (i.e. from smaller to larger prey items) and become enriched in ¹⁵N; 3) individuals acquire energy from spatially confined local resource pools and exhibit limited displacement; and 4) species exhibit similarly sized and highly overlapping trophic niches. We found some evidence for ontogenetic shifts in habitat‐use (δ¹³C) for thorny and little skate and diet (δ¹⁵N) of thorny and winter skate and hypothesize that individuals exhibit gradual trophic niche transition, especially in δ¹⁵N space, rather than a clear and distinct shift in diet throughout ontogeny. Spatial isoscapes generated for little, thorny, and winter skate highlighted distinct spatial patterns in isotopic composition across the coastal shelf. For little and thorny skate, patterns mimicked expected spatial variability in the isotopic composition of phytoplankton/POM, suggesting limited displacement and utilization of spatially confined resource pools. Winter skate, however, exhibited a much narrower range of δ¹³C and δ¹⁵N values, suggesting individuals may use resources from a more confined latitudinal range. Although high total trophic niche overlap was observed between some species (e.g. little and thorny skate), sympatric species (e.g. little and winter skate) exhibited a degree of trophic niche separation. These findings offer new insight into the trophic dynamics of a poorly‐studied, vulnerable group of predators, and highlight a need to re‐examine assumptions pertaining to aspects of their ecology.     

Variability in isotope discrimination factors in coral reef fishes: implications for diet and food web reconstruction

Interpretation of stable isotope ratios of carbon and nitrogen (δ(13)C and δ(15)N) is generally based on the assumption that with each trophic level there is a constant enrichment in the heavier isotope, leading to diet-tissue discrimination factors of 3.4‰ for (15)N (ΔN) and ～0.5‰ for (13)C (ΔC). Diet-tissue discrimination factors determined from paired tissue and gut samples taken from 152 individuals from 26 fish species at Ningaloo Reef, Western Australia demonstrate a large amount of variability around constant values. While caution is necessary in using gut contents to represent diet due to the potential for high temporal variability, there were significant effects of trophic position and season that may also lead to variability in ΔN under natural conditions. Nitrogen enrichment increased significantly at higher trophic levels (higher tissue δ(15)N), with significantly higher ΔN in carnivorous species. Changes in diet led to significant changes in ΔN, but not tissue δ(15)N, between seasons for several species: Acanthurus triostegus, Chromis viridis, Parupeneus signatus and Pomacentrus moluccensis. These results confirm that the use of meta-analysis averages for ΔN is likely to be inappropriate for accurately determining diets and trophic relationships using tissue stable isotope ratios. Where feasible, discrimination factors should be directly quantified for each species and trophic link in question, acknowledging the potential for significant variation away from meta-analysis averages and, perhaps, controlled laboratory diets and conditions.     

Evidence of two subaggregations of humpback whales on the Kodiak,Alaska, feeding ground revealed from stable isotope analysis

Knowledge of humpback whale (Megaptera novaeangliae) foraging on feeding grounds is becoming increasingly important as the growing North Pacific population recovers from commercial whaling and consumes more prey, including economically important fishes. We explored spatial and temporal (interannual, within‐season) variability in summer foraging by humpback whales along the eastern side of the Kodiak Archipelago as described by stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios of humpback whale skin (n = 118; 2004–2013). The trophic level (TL) of individual whales was calculated using basal food web δ¹⁵N values collected within the study area. We found evidence for the existence of two subaggregations of humpback whales (“North,” “South”) on the feeding ground that fed at different TLs throughout the study period. Linear mixed models suggest that within an average year, Kodiak humpback whales forage at a consistent TL during the feeding season. TL estimates support mixed consumption of fish and zooplankton species in the “North” (mean ± SE; 3.3 ± 0.1) and predominant foraging on zooplankton in the “South” (3.0 ± 0.1). This trend appears to reflect spatial differences in prey availability, and thus, our results suggest North Pacific humpback whales may segregate on feeding aggregations and target discrete prey species.     

Stable isotope tracking of endangered sea turtles: validation with satellite telemetry and δ15N analysis of amino acids

Effective conservation strategies for highly migratory species must incorporate information about long-distance movements and locations of high-use foraging areas. However, the inherent challenges of directly monitoring these factors call for creative research approaches and innovative application of existing tools. Highly migratory marine species, such as marine turtles, regularly travel hundreds or thousands of kilometers between breeding and feeding areas, but identification of migratory routes and habitat use patterns remains elusive. Here we use satellite telemetry in combination with compound-specific isotope analysis of amino acids to confirm that insights from bulk tissue stable isotope analysis can reveal divergent migratory strategies and within-population segregation of foraging groups of critically endangered leatherback sea turtles (Dermochelys coriacea) across the Pacific Ocean. Among the 78 turtles studied, we found a distinct dichotomy in δ(15)N values of bulk skin, with distinct "low δ(15)N" and "high δ(15)N" groups. δ(15)N analysis of amino acids confirmed that this disparity resulted from isotopic differences at the base of the food chain and not from differences in trophic position between the two groups. Satellite tracking of 13 individuals indicated that their bulk skin δ(15)N value was linked to the particular foraging region of each turtle. These findings confirm that prevailing marine isoscapes of foraging areas can be reflected in the isotopic compositions of marine turtle body tissues sampled at nesting beaches. We use a Bayesian mixture model to show that between 82 and 100% of the 78 skin-sampled turtles could be assigned with confidence to either the eastern Pacific or western Pacific, with 33 to 66% of all turtles foraging in the eastern Pacific. Our forensic approach validates the use of stable isotopes to depict leatherback turtle movements over broad spatial ranges and is timely for establishing wise conservation efforts in light of this species' imminent risk of extinction in the Pacific.     

Geographic and ontogenetic variation in the diet of two commonly exploited batoids (Chilean eagle ray and Pacific guitarfish) off Peru: evidence of trophic plasticity

This study provides information about the diet across geographic areas and throughout ontogeny and sex of two coastal and commercial batoid species in Peru (Chilean eagle ray Myliobatis chilensis and Pacific guitarfish Pseudobatos planiceps). Data was collected in the central coast (13°30′S to 14°30′S; Pisco district, Lima) and in the northern coast (13°12′S to 13°49′S; San Jose district, Lambayeque) off Peru during the second semester of the years 2015 and 2016 (i.e., winter and spring) in an El Niño event. A total of 357 stomach contents were analyzed in northern and central Peru with different oceanographic and ecological conditions. In the central coast, M. chilensis showed a high trophic position (tertiary consumer) due to its high consumption of Peruvian anchoveta (Engraulis ringens), while P. planiceps had a lower trophic position (secondary consumer) and a less specialized diet of benthic invertebrates (i.e., crustaceans and mollusks) and pelagic fish (e.g., E. ringens). In the northern coast, both species preyed mainly upon benthic invertebrates and to a lesser degree on fish; therefore, their trophic position was lower. Dietary variation was influenced by species, geographic location, and ontogeny. The diet variability between geographic locations shows insights of these batoids’ trophic plasticity and opportunistic feeding behavior in response to differences in the local prey availability, an effect that may be amplified during the El Niño–Southern Oscillation (ENSO). The spatial variation in this species diet may indicate that they play different ecological roles in distinct environments. This study contributes to the scarce literature about batoids’ ecology in the southeast Pacific Ocean and presents novel information on habitat-specific diet composition.

     

Multi‐decadal and ontogenetic trophic shifts inferred from stable isotope ratios of pinniped teeth

Identifying and characterizing top predators’ use of trophic resources provides important information about animal ecology and their response to changing conditions. Information from sources such as stable isotopes can be used to infer changes in resource use as direct observations in the wild are difficult to obtain, particularly in the marine environment. Stable carbon and nitrogen isotope values were recovered from the canine teeth of grey seals collected from haul outs in the central North Sea in the 1970/1980s (n = 44) and 2000s (n = 25), spanning a period of marked ecosystem changes in the region. Extracting material deposited during juvenile and adult life‐stages, we reconstructed a multi‐decadal record of δ¹⁵N and δ¹³C variation. Using established correlations between stable isotope ratios and sea bottom temperature we created a proxy for baseline isotopic variability to account for this source of temporal change. We found 1) a significant long‐term decline in juvenile grey seal δ¹⁵N values, suggesting trophic position has decreased over time; 2) a decline in adult δ¹⁵N values and contraction in stable isotopic niche space after the North Sea regime shift, signifying both a decline in trophic position and change in foraging habits over the 20th century; and 3) evidence for dietary segregation between juvenile and adult animals, showing juvenile individuals feeding at a lower trophic position and in more nearshore areas than adults. Our results demonstrate the efficacy of mining archived biological samples to address ecological questions and imply important ontogenetic and long‐term shifts in the feeding ecology of a top predator. Long‐term changes in grey seal trophic dynamics may be partly in response to well documented ecosystem changes in the North Sea. Such indirect monitoring of marine predators may have utility when set in the context of ecosystem assessments where paucity of long‐term monitoring data is prevalent.     

Mother-offspring isotope fractionation in two species of placentatrophic sharks

Stable-isotope values of a scalloped hammerhead Sphyrna lewini and blacktip shark Carcharhinus limbatus and their respective embryos were analysed. Embryos of both species were enriched in δ(15) N compared to their mothers (0·82 and 0·88‰, respectively), but fractionation of δ(13) C varied. Embryonic S. lewini were enriched (1·00‰) in δ(13) C while C. limbatus were depleted (0·27‰) relative to their mothers.     

The trophic niche of sculpins Cottus spp. in forage fish assemblages of boreal lakes

We compared the trophic niches of freshwater sculpins Cottus spp. with those of other co-habiting forage fishes in two groups of boreal lakes with distinct habitats and fish communities. Near North Lakes (45° 00′ to 47° 30′ N) were deeper, stratified and contained lake trout Salvelinus namaycush as the apex piscivore, whereas Far North Lakes (51° 10′ to 52° 20′ N) were shallower, did not stratify and contained pike Esox lucius and walleye Sander vitreus as the apex piscivores. Trophic niches of sculpins and other forage fishes were compared based on niche metrics calculated from muscle stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios. In Near North Lakes, sculpins were found almost exclusively in deep, offshore waters and their niche positions reflected a greater reliance on pelagic production (lower δ¹³C) and a higher trophic elevation (higher δ¹⁵N) compared with most other forage fishes. Furthermore, sculpins in Near North Lakes tended to have larger trophic niches (occupied greater area in δ¹³C– δ¹⁵N space), particularly in the food chain (δ¹⁵N) dimension, than other cohabiting forage fishes. In contrast, sculpins in Far North Lakes were commonly found in the nearshore and had trophic niche positions and sizes that were similar to those of the other cohabiting forage fishes. This study illustrates the flexibility in the realised trophic niches of sculpins in relation to habitat availability and fish community composition in boreal lakes.     

Combined gut‐content and stable isotope trophic analysis of the pelagic stingray Pteroplaytrygon violacea (Bonaparte, 1832) diet from the western North Atlantic Ocean

The understanding of trophic relationships is vital for correctly modeling ecosystems and ecosystem effects of fisheries removals. The pelagic stingray is found in epipelagic sub‐tropical and tropical waters worldwide and is a common bycatch in pelagic longline fisheries. Between August 2008 and November 2011, 156 specimens (81 males; 75 females) were collected during pelagic longline fishing operations in the US South Atlantic Bight and Gulf of Mexico. Stomach content analyses found that the major prey items were cephalopod molluscs (59.18%), followed by actinopterygiian fishes (37.75%), and decapod crustaceans (35.71%). These concentrations of prey items found in the stomachs coincide with previous studies done in the Pacific Ocean. In contrast to previous studies that found high percentages of empty stomachs (63%), the current percentage of empty stomachs was much lower (25.6%), likely due to shorter times between collection and inspection. Stable isotope analysis (δ¹³C and δ¹⁵N) was performed on white muscle in order to correlate the trophic position with gut‐content analysis. The δ¹³C values ranged from ‐18.81‰ to ‐16.70‰, while the δ¹⁵N ranged from 6.11‰ to 11.88‰. Modeling of stable isotope data suggest that while squid are occasionally an important part of the pelagic stingray diet, prey usually consist of shrimp and other pelagic crustaceans. Pelagic stingrays fed within two trophic levels, but their prey appeared to feed on different carbon sources than those found in other pelagic elasmobranchs. A deeper understanding of the pelagic stingray diet sources can help fisheries management as it begins to transition into ecosystem‐based management.     

New insights on the trophic ecology of blue (Prionace glauca) and shortfin mako sharks (Isurus oxyrinchus) from the oceanic eastern South Pacific

The blue shark (Prionace glauca) and the shortfin mako shark (Isurus oxyrinchus) are two large and highly migratory sharks distributed in most oceans. Although they are often caught in the south Pacific Ocean long-line fisheries, their trophic ecology is poorly understood. Stable isotopes with Bayesian mixing and dependence concentration models were performed to determine the diet and trophic differences between the two species in the South-eastern Pacific Ocean. According to the mixing models, fishes are the most important prey of these sharks. Dolphin calves and remains were found in the stomachs of both species, which represents a novel finding in trophic ecology of South Pacific sharks. Intra-specific differences were found in P. glauca, but not in specimens of I. oxyrinchus. The two sharks showed a high degree of diet overlap (73%), primarily over mackerel and dolphin carcasses. Our results indicate that blue and shortfin mako sharks have a generalist feeding strategy in the eastern Pacific Ocean, with a strong preference for teleost fishes and also for dolphin carcasses. Therefore, trophic studies are useful to understand energy flow through the food web, and the trophic position of key species.     

Resource use of great hammerhead sharks (Sphyrna mokarran) off eastern Australia

Great hammerhead sharks Sphyrna mokarran are the largest member of Sphyrnidae, yet the roles of these large sharks in the food webs of coastal ecosystems are still poorly understood. Here we obtained samples of muscle, liver and vertebrae from large S. mokarran (234–383 cm total length; L_T) caught as by-catch off eastern Australia and used stable-isotope analyses of δ¹⁵N, δ¹³C and δ³⁴S to infer their resource use and any associated ontogenetic patterns. The results indicated large S. mokarran are apex predators primarily relying on other sharks and rays for their diet, with a preference for benthic resources such as Australian cownose rays Rhinoperon neglecta during the austral summer. Teleosts, cephalopods and crustaceans were not significant components of S. mokarran diets, though some conspecifics appeared to rely on more diverse resources over the austral summer. Ontogenetic shifts in resource use were detected but trajectories of the increases in trophic level varied among individuals. Most S. mokarran had non-linear trajectories in ontogenetic resource-use shifts implying size was not the main explanatory factor. Stable isotope values of δ¹³C and δ³⁴S in muscle suggest S. mokarran span coastal, pelagic and benthic food webs in eastern Australia.     

Spatial,ontogenetic and interspecific variability in stable isotope ratios of nitrogen and carbon of Merluccius capensis and Merluccius paradoxus off South Africa

General linear models (GLMs) were used to determine the relative importance of interspecific, ontogenetic and spatial effects in explaining variability in stable isotope ratios of nitrogen (δ¹⁵N) and carbon (δ¹³C) of the co‐occurring Cape hakes Merluccius capensis and Merluccius paradoxus off South Africa. Significant GLMs were derived for both isotopes, explaining 74 and 56% of observed variance in Merluccius spp. δ¹⁵N and δ¹³C, respectively. Spatial effects (west or south coast) contributed most towards explaining variability in the δ¹⁵N model, with Merluccius spp. off the west coast having higher (by c. 1·4‰) δ¹⁵N levels than Merluccius spp. off the south coast. Fish size and species were also significant in explaining variability in δ¹⁵N, with both species showing significant linear increases in δ¹⁵N with size and M. capensis having higher (by c. 0·7‰) δ¹⁵N values than M. paradoxus. Species and coast explained most and similar amounts of variability in the δ¹³C model, with M. capensis having higher (by c. 0·8‰) δ¹³C values than M. paradoxus, and values being lower (by c. 0·7‰) for fishes off the west coast compared with the south coast. These results not only corroborate the knowledge of Merluccius spp. feeding ecology gained from dietary studies, in particular the ontogenetic change in trophic level corresponding to a changing diet, but also that M. capensis feeds at a slightly higher trophic level than M. paradoxus. The spatial difference in Merluccius spp. δ¹⁵N appears due to a difference in isotopic baseline, and not as a result of Merluccius spp. feeding higher in the food web off the west than the south coast, and provides new evidence that corroborates previous observations of biogeographic differences in isotopic baselines around the South African coast. This study also provides quantitative data on the relative trophic level and trophic width of Cape hakes over a large size range that can be used in ecosystem models of the southern Benguela.