Effects of intrinsic and extrinsic factors on the diet of Bathyraja macloviana,a benthophagous skate

The effects of intrinsic (sex, maturity stage and body size) and extrinsic (depth and region) factors on the diet of Bathyraja macloviana, in the south‐west Atlantic Ocean, were evaluated using a multiple‐hypothesis modelling approach. Bathyraja macloviana fed mainly on polychaetes followed by amphipods, isopods and decapods. Effects of intrinsic and extrinsic factors on diet composition of this species were found. The consumption of polychaetes had a humped relationship with total length (L_T), and isopods and decapods increased with increasing L_T. Immature individuals preyed on amphipods more heavily than mature individuals. Furthermore, region and depth had an important effect on the consumption of isopods, decapods and amphipods. Such ontogenetic changes and spatial patterns may provide insights into understanding the regulatory mechanisms of marine communities.     

Feeding habits of ocellate spot skate,Okamejei kenojei (Müller & Henle, 1841), in coastal waters of Taean,Korea

The feeding habits of Okamejei kenojei were studied using 592 specimens collected in the coastal waters of Taean, Korea from April 2008 to March 2009. O. kenojei is a bottom‐feeding carnivore that consumes mainly shrimp, fishes, and crabs. Its diet also includes small quantities of amphipods, mysids, cephalopods, euphausiids, copepods, isopods, and polychaetes. The total length (TL) of individuals in this study ranged from 8.2 to 49.0 cm. Cluster analysis based on %IRI (index of relative importance) identified three size classes. Group A (< 20 cm TL) ate primarily caridean shrimp and amphipods; group B (20–30 cm TL) ate exclusively shrimp; and group C (> 30 cm TL) ate penaeoidean shrimp, fishes, and crabs. O. kenojei showed ontogenetic changes in feeding habits. Although shrimps were the primary food consumed by all size groups, the proportion of shrimp in the total diet decreased and the consumption of fishes and crabs gradually increased with the body size of O. kenojei. Size of the prey organisms also increased. Smaller individuals fed mainly on small prey, such as amphipods, mysids, and small shrimp, whereas larger individuals preferred larger prey, such as larger shrimp, fishes, and crabs. The size‐related diet breadth and the percentage of empty stomachs were significant; the diet breadth gradually increased with body size, whereas the percentage of empty stomachs decreased. Seasonal changes in the O. kenojei diet were not significant, but shrimp constituted 97.3% of the summer diet by %IRI. Seasonal changes in diet breadth and the percentage of empty stomachs were not significant.     

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.     

The food and feeding habits of blackspot snapper, Lutjanus fulviflamma (Pisces: Lutjanidae) in shallow waters of Mafia Island, Tanzania

Natural diet and feeding habits of blackspot snapper, Lutjanus fulviflamma (Forsskål 1775) were investigated in Mafia Island Marine Park (MIMP) and intensively fished areas (IFA) between May 1999 and April 2001. An index of relative importance (IRI) was used to assess the food preference for L. fulviflamma. Feeding habits were studied with respect to sex and size of the predator, seasons and feeding periodicity. Of 4642 L. fulviflamma specimens sampled, 46% had empty stomachs. Brachyurans were the main prey of L. fulviflamma accounting for 48% IRI. Benthic invertebrates dominated by other crustaceans, gastropods, bivalves, cephalopods and polychaetes and demersal teleosts made up the remainder of the diet. Female and male L. fulviflamma were not effectively segregated by trophic dimension in the study areas. Specimens in the 90–120 mm total length (TL) fed exclusively on crustaceans, with shrimps accounting for 40% IRI. Lutjanus fulviflamma in the 270–300 mm TL preferred fish prey accounting for 50% IRI. The diet of fish species showed clear spatial differences, which were dependent on size distributions. Lutjanus fulviflamma fed mostly but not exclusively at night peaking at dusk and dawn. The results clearly demonstrate that L. fulviflamma is an opportunistic forager, which exhibits both diel and ontogenetic feeding habits.     

Diet of the Antarctic starry skate Amblyraja georgiana (Rajidae, Chondrichthyes) at South Georgia (Southern Ocean)

Stomach contents were identified from 206 Antarctic starry skate (Amblyraja georgiana) that were collected during three groundfish surveys (September 2007, April 2008 and January 2009) at South Georgia, Southern Ocean. The diet of A. georgiana varied with skate size and between years. Preferred prey included fish (particularly for larger individuals) and Antarctic krill, Euphausia superba, as well as amphipods, polychaetes and other benthic fauna. The skate A. georgiana appears to be an opportunistic predator, and the clear presence of Antarctic krill in this demersal predator’s diet may indicate a benthic habit of this euphausiid species, which has hitherto mainly been considered as occupying a purely pelagic niche.     

Natural born killers: an invasive amphipod is predatory throughout its life-history

Introduced predators can have profound impacts on prey populations, with subsequent ramifications throughout entire ecosystems. However, studies of predator–prey interaction strengths in community and food-web analyses focus on adults or use average body sizes. This ignores ontogenetic changes, or lack thereof, in predatory capabilities over the life-histories of predators. Additionally, large individual predators might not be physically capable of consuming very small prey individuals. Both situations are important to resolve, as native prey may or may not therefore experience ontogenetic or size refuges from invasive predators. Here, we find that the freshwater amphipod invader, Gammarus pulex, is predatory throughout its development from juvenile through to adult. All size classes collected in the field had a common prey, nymphs of the mayfly Baetis rhodani, in their guts. In an experiment with predator, prey and experimental arenas scaled for body size, G. pulex juveniles and adults consumed B. rhodani in all size-matched categories. In a second experiment, the largest G. pulex individuals were able to prey on the smallest B. rhodani. Thus, the prey do not benefit from any ontogenetic or size refuge from the predator. This corroborates with the known negative population abundance relationships between this invasive predator and its native prey species. Understanding and predicting invasive predator impacts will be best served when interactions among all life-history stages of predator and prey are considered.     

Food habits of the longnose skate, <Emphasis Type="Italic">Raja rhina</Emphasis> (Jordan and Gilbert, 1880), in central California waters

Feeding studies can provide researchers with important insights towards understanding potential fishery impacts on marine systems. Raja rhina is one of the most common elasmobranch species landed in central and northern California demersal fisheries, yet life history information is extremely limited for this species and aspects of its diet are unknown. Specimens of R. rhina were collected between September, 2002 and August, 2003 from fisheries-independent trawl surveys. Percent Index of Relative Importance values indicated that the five most important prey items in 618 stomachs of R. rhina were unidentified teleosts (31.6% IRI), unidentified shrimps (19.6% IRI), unidentified euphausiids (10.9% IRI), Crangonidae (7.4% IRI), and Neocrangon resima (6.0% IRI). There were significant dietary shifts with increasing skate total length and with increasing depths. Smaller skates ate small crustaceans and larger skates ate larger fishes and cephalopods. With increasing depths, diet included bentho-pelagic teleosts and more cephalopods and euphausiids. The findings of this study are consistent with previous researchers that report similar diet shifts in skate species with size and depth.     

Effects of body size, age and maturity stage on diet in a large shark: ecological and applied implications

Ontogenetic diet shifts are a widespread phenomenon among vertebrates, although their relationships with life history traits are poorly known. We analyzed the relative importance of body size, age and maturity stage as determinants of the diet of a marine top predator, the copper shark, Carcharhinus brachyurus, by examining stomach contents using a multiple-hypothesis modeling approach. Copper sharks shifted their diet as size and age increased and as they became sexually mature, incorporated larger prey as they grew, and had a discrete shift in diet with body size, with only individuals larger than ≈200 cm total length able to prey on chondrichthyans. Body size was the most important trait explaining the consumption of chondrichthyans, while age determined the consumption of pelagic teleosts. Pelagic teleosts were consumed mostly by medium-aged sharks, a result, probably, of a risk-reducing feeding strategy at young ages coupled with either a senescence-related decline in performance or a change in sensory capabilities as sharks age. Copper sharks of all sizes were able to cut prey in pieces, implying that gape limitation (i.e., the impossibility of eating prey larger than a predator’s mouth) did not play a role in producing the diet shift. Our results suggest that, contrary to the current practice of setting minimum but not maximum size limits in catches, any plan to conserve or restore the ecological function of sharks, through their predatory control of large prey, should aim to maintain the largest individuals.     

Feeding habits and predator‐prey size relationships of mandarin fish Siniperca chuatsi (Basilewsky) in a shallow lake,central China

The diet and predator‐prey size relationships of mandarin fish Siniperca chuatsi (Basilewsky) in Lake Xiaosihai along the middle reach of the Yangtze River were studied through stomach content analysis. A total of 401 specimens (91–539 mm total length) were collected in 2007. The diet was dominated by topmouth gudgeon Pseudorasbora parva, sharpbelly Hemiculter leucisculus, redfin culter Cultrichthys erythropterus, and crucian carp Carassius auratus, with significant seasonal and ontogenetic differences. Ontogenetic variation in diet was apparent that larger prey items such as crucian carp and redfin culter became more common, while smaller prey such as topmouth gudgeon, bitterlings and shrimps gradually declined in the larger fish. Mandarin fish total length (TL) was strongly related to mouth gape width (GW) and gape height (GH). Mandarin fish TL and prey fish TL as well as mandarin fish GW and prey fish body depth (BD) were positively and linearly related for sharpbelly, redfin culter and crucian carp. Strong size selectivity for topmouth gudgeon, sharpbelly and redfin culter as prey indicated that the piscivorous mandarin fish can have potential impact on the population size structure of the three prey fish.     

Feeding habits of <Emphasis Type="Italic">Bathydraco marri</Emphasis> (Pisces,Notothenioidei, Bathydraconidae) from the Ross Sea,Antarctica

In comparison with other bathydraconids, all species of the genus Bathydraco are poorly known from an ecological perspective. The diet of juvenile Bathydraco marri Norman, 1938 was studied for the first time in specimens collected in the southwestern Ross Sea during summer 1998. Fish were collected in a single otter trawl catch at 330–340 m depth. The stomach content analysis showed that this species fed exclusively on crustaceans. Overall, 20 prey taxa were identified to genus or species level. Mysids, amphipods and copepods were the most important prey in decreasing order of importance. Other prey, such as Euphausia superba, isopods and tanaids were eaten occasionally and in very small amounts. A multivariate analysis was applied to feeding data to assess ontogenetic or sex-related changes in diet. No difference was detected between sexes, whereas diet of small and large fish differed in some degree. An ontogenetic shift from small and pelagic crustaceans such as copepods to benthic–benthopelagic prey such as amphipods and mysids was observed. Relating present results with published data on physiological characteristics of B. marri, it was possible to infer their feeding behaviour and mode of life. Like other bathydraconids, this species appeared to be an inactive and sluggish fish, which relied on more or less motile benthic or epibenthic prey adopting a “sit and wait” feeding strategy. On the other hand, smaller fish seem to be more active, feeding also on pelagic prey such as copepods that can be seasonally abundant, thus reducing the intraspecific competition for food.     

Ontogenetic dietary shifts and feeding behavior of the tiger shark, Galeocerdo cuvier, in Hawaiian waters

Synopsis Stomach content data from 281 tiger sharks caught during shark control programs in Hawaii between 1967 and 1969, and during 1976 were analyzed to examine feeding habits and ontogenetic shifts in diet. As sharks increased in size, prey diversity and frequency of occurrence of large prey items increased. The percent occurrence of teleosts and cephalopods in stomachs decreased as sharks increased in length, while occurrence of elasmobranchs, turtles, land mammals, crustaceans, and undigestible items increased. Comparisons between the diets of tiger sharks from Hawaii and other locations indicate that ontogenetic shifts are universal in this species and that tiger sharks may be opportunistic feeders that prey heavily on abundant, easy to capture prey. Small tiger sharks may be spatially segregated from medium and large sharks and appear to be primarily nocturnal, bottom feeders. Large tiger sharks feed near the bottom at night, but also feed at the surface during the day. Prey, similar in size to humans, begin to occur in the diet of tiger sharks approximately 230 cm TL, and therefore sharks of this size and larger may pose the greatest threat to humans. Ontogenetic shifts in diet may be attributed to increased size of sharks, expanded range and exploitation of habitats of larger sharks, and/or improved hunting skill of larger sharks.Deceased 1974     

Standardized diet compositions and trophic levels of skates (Chondrichthyes: Rajiformes: Rajoidei)

Skates by virtue of their abundance and widespread occurrence appear to play an influential role in the food webs of demersal marine communities. However, few quantitative dietary studies have been conducted on this elasmobranch group. Therefore, to better understand the ecological role of skates, standardized diet compositions and trophic level (TL) values were calculated from quantitative studies, and compared within and among skate and shark taxa. Prey items were grouped into 11 general categories to facilitate standardized diet composition and TL calculations. Trophic level values were calculated for 60 skate species with TL estimates ranging from 3.48 to 4.22 (mean TL = 3.80 ± 0.02 SE). Standardized diet composition results revealed that decapods and fishes were the main prey taxa of most skate species followed by amphipods and polychaetes. Correspondingly, cluster analysis of diet composition data revealed four major trophic guilds, each dominated by one of these prey groups. Fish and decapod guilds were dominant comprising 39 of 48 species analyzed. Analysis of skate families revealed that the Arhynchobatidae and Rajidae had similar TL values of 3.86 and 3.79 (t-test, P = 0.27), respectively. The Anacanthobatidae were represented by a single species, Cruriraja parcomaculata, with a TL of 3.53. Statistical comparison of TL values calculated for five genera (Bathyraja, Leucoraja, Raja, Rajella, Rhinoraja) revealed a significant difference between Bathyraja and Rajella (t-test, P = 0.03). A positive correlation was observed between TL and total length (L _T) with larger skates (e.g. >100 cm L _T) tending to have a higher calculated TL value (>3.9). Skates were found to occupy TLs similar to those of several co-occurring demersal shark families including the Scyliorhinidae, Squatinidae, and Triakidae. Results from this study support recent assertions that skates utilize similar resources to those of other upper trophic-level marine predators, e.g. seabirds, marine mammals, and sharks. These preliminary findings will hopefully encourage future research into the trophic relationships and ecological impact of these interesting and important demersal predators.     

Trophic ecology of yellownose skate Zearaja chilensis,a top predator in the south‐western Atlantic Ocean

The diet and trophic level (T_L) of the yellownose skate Zearaja chilensis in the south‐western Atlantic Ocean (35°–54° S), and how these varied in relation to body size, sex, maturity stage, depth and region were determined by analysis of stomach contents. From 776 specimens analysed, 671 (86·5%) ranging from 180 to 1190 mm total length (L_T) had prey in their stomachs. The diet was dominated by fishes, mainly the notothenioid Patagonotothen ramsayi and the Argentine hake Merluccius hubbsi. The consumption of fishes and crabs increased with increasing predator size, and these preys were more important in the north than in the south. Isopods and other crustaceans were consumed more in the south and their consumption decreased as the size of Z. chilensis increased. The T_L of Z. chilensis increased with L_T from 4·29 to 4·59 (mean 4·53), confirming their ecological role as a top predator. The small and large size classes exhibited a low diet overlap and the highest spatial segregation, whereas medium and large specimens had higher co‐occurrence and dietary overlap indices. A clear distinction in tooth shape was noted between sexes in adult specimens, with males having longer cusps. This sexual heterodonty may be related to reproductive behaviour, increasing the grasping ability of males during courtship, because there were no differences in diet between the sexes.     

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.     

Diet composition of greater weever,Trachinus draco (Linnaeus, 1758) captured in the eastern‐central Adriatic Sea in relation to fish size,season and sampling area

The diet compositions of greater weever, Trachinus draco, from the eastern‐central Adriatic Sea, were investigated with respect to seasons, fish size and sampling locations. Analyzed were the stomach contents of 592 specimens, total length (TL) of 9.9–31.2 cm, collected by commercial bottom trawls from January to December 2008. The frequency of empty stomachs varied significantly with the season (43.3% maximum during winter; 6.8% minimum during summer). Prey items identified in the stomachs belonged to seven major taxonomic groups: Bivalvia, Cephalopoda, Decapoda, Mysidacea, Isopoda, Amphipoda and Teleostei. Decapods were the most important prey (%IRI = 72.8) followed by teleosts (%IRI = 16.4) and mysids (%IRI = 15.0); other prey groups were only occasionally ingested. Diet composition showed little seasonal variation; decapods were the most important prey in all seasons. Fish size was the most important factor influencing the composition of the diet. Small individuals (<15 cm TL) fed primarily on small crustaceans (mysids), whereas larger fish consumed bigger prey, such as decapods, teleosts and cephalopods. There was high dietary similarity between sampling locations. Based on the diet, greater weever can be a considered a generalistic predator.     

Dietary niche differentiation of five sympatric species of Platycephalidae

The dietary composition and partitioning of food resources between five sympatric species of Platycephalidae inhabiting the coastal waters of New South Wales, Australia was investigated. Samples were collected monthly between March and November 2007 onboard commercial ocean prawn trawlers based in the ports of Yamba and Newcastle. Monthly percentage weight contribution of 12 prey categories was analysed to determine if diet was influenced by the variables: species, location, depth, size and maturity. Of the 959 stomachs from the five species examined, 28–54% contained prey. All Platycephalid species primarily consumed teleosts, however the diversity of prey and the proportion each prey type contributed to the overall diet varied substantially between species. Platycephalus caeruleopunctatus, P. longispinis, P. richardsoni and Ambiserrula jugosa were generalist carnivores and consumed prey from a wide variety of phyla including teleosts, crustaceans, polychaetes, molluscs and echinoderms. In contrast, Ratabulus diversidens were primarily piscivorous. Partitioning of prey resources between species was more evident in waters at Yamba than at Newcastle. Differences in diet between locations were considered a result of differential prey exploitation rather than shifts in the suite of prey consumed. Dietary composition was observed to be influenced by size, maturity status and depth however these differences were not observed for all species.     

Seasonal diet and amphipod size selection of juvenile common bully, <Emphasis Type="Italic">Gobiomorphus cotidianus</Emphasis>, in a coastal New Zealand lake

We examined the seasonal diet of juvenile (≤32 mm total length) common bullies, Gobiomorphus cotidianus, in Lake Waihola, South Island, New Zealand and whether they preyed selectively on the amphipod, Paracalliope fluviatilis, which was a dominant prey item. Generally, ≥75% of all fish sampled had consumed amphipods, which comprised 80%, or more, of the total prey volumes in fish stomachs over most of a year except on one sampling occasion. Copepods, snails, chironomids, Daphnia, mysids and isopods also contributed to the diet. Comparisons of the size distribution of amphipods in fish stomachs and the lake on the date of sampling showed that on four occasions, juvenile bullies selectively consumed small amphipods. No selective feeding was detected on the other occasions. The dominance of primarily benthic amphipods in the diet of juvenile bullies which, in turn, are preyed on by piscivores, suggests a strong benthic-pelagic link in the transfer of energy in Lake Waihola. Handling editor: J. Cambray     

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.