Comparative feeding ecology of four sympatric skate species off central California, USA

The big (Raja binoculata), California (R. inornata), longnose (R. rhina), and sandpaper (Bathyraja kincaidii), skates are commonly found on soft-bottom regions of the central California continental shelf and upper slope. The feeding ecology of this assemblage was compared to evaluate the degree of trophic separation among species, based on the results of previous species-specific diet studies. Specimens were collected from fishery independent trawl surveys conducted during September 2002–March 2003 at depths of 9–536 m. Using single and compound measures, diet composition of small (≤60 cm TL) and large (>60 cm TL) individuals were compared within continental shelf (≤200 m) and slope (>200 m) regions using traditional, multivariate, and novel techniques. Diet compositions within size classes were similar in both regions. Diet compositions between size classes generally differed, however, with fishes more important and crustaceans (especially shrimps and euphausiids) less important in the diets of large individuals. Crabs contributed substantially to skate diet compositions on the shelf, but were uncommon prey items at deeper depths, probably because of their relative scarcity in slope waters. Conversely, cephalopods were common prey items at slope depths, but were rarely ingested at shelf depths. The studied skate assemblage appears to consist primarily of generalist crustacean and fish predators that exhibit high dietary overlap at similar sizes. It is possible that resource competition among skates and groundfish species has been reduced because of considerable recent declines in the biomass of upper trophic level groundfishes. Skates may therefore play important roles in contemporary benthic food web dynamics off central California.     

Dietary comparisons of six skate species (Rajidae) in south-eastern Australian waters

The diet of six skate species caught as bycatch in south-eastern Australian waters was examined over a 2-year period. The skates were segregated into two regions (continental shelf and continental slope) based on prey species and depth of capture. The shelf group consisted of four species, Dipturus sp. A, D. cerva, D. lemprieri and D. whitleyi, while the slope group comprised two species, Dipturus sp. B and D. gudgeri. The two groups varied in feeding strategies with the shelf species generally occupying a broader feeding niche and preying on a larger diversity of prey including a variety of crustaceans (brachyurans, anomurans, achelates, carideans and dendobranchiates), cephalopods, elasmobranchs and teleosts. Within the slope group, Dipturus sp. B and D. gudgeri were more specialised. Dipturus sp. B preyed primarily on anomurans (galatheids) and bachyurans (homolids), whereas D. gudgeri preyed primarily on teleosts. A size related change in diet was evident for all species with the exception of D. gudgeri in which all sizes preyed predominantly on teleosts. Smaller representatives of the four shelf species all preyed on numerous amounts of caridean shrimps, in particular Leptochela sydniensis. In contrast, the continental slope species, Dipturus sp. B consumed anomurans when small, shifting to brachyurans with increasing size. Of the six skate species examined in this study, three were secondary consumers (trophic level <3) and the remaining three tertiary consumers (trophic level >4). Although ANOSIM found significant differences in dietary composition between species within groups, there was some overlap in prey species amongst co-existing skates, which suggests that there is some degree of resource partitioning amongst them.     

Reanalysis of long‐term surveys on the ecology and biology of mud skate (Rhinoraja taranetzi Dolganov, 1985) in the northwestern Pacific (1993–2002)

Based on data sampled in 1992–2002, the occurrence, spatial distribution, bathymetry, bottom temperatures preferences, size composition, feeding, and some features of reproductive biology of mud skate Rhinoraja taranetzi in the Pacific waters off the northern Kuril Islands and southeastern Kamchatka are considered. Throughout the year, the mud skate was most abundant in the central part of the study area, from the coast of Onekotan Island to the southern tip of Kamchatka peninsula. The proportion of this species in bottom trawl catches in different seasons has changed slightly. However, maximum catch occurred in September–December. In April–May mud skates occupied shallower depths (mean 230–270 m), moving deeper in the summer period (mean 340–390 m). In December–March the skate occurred at lower bottom temperatures (mean 0.8–1.6°C) whereas the rest of the year it inhabited warmer waters with mean bottom temperatures of 2.5–3.1°C. Throughout the year decreasing body weight with depth was observed, indicating that adult and juvenile mud skates inhabit different depths. Total length of mud skates in catches ranged from 17 to 70 cm with a mean of 51.71 cm. Relation between total length (TL, cm) and body weight (W, g) was: W = 0.0029TL^3.1614 (r² = 0.978). Males were more abundant among small skates (<30 cm) only, whereas females predominated among larger skates (>30 cm) comprising about 70% in 60–70 cm length class. Female mud skates were longer and heavier than males (mean length 56.9 vs 51.2 cm and mean weight 1206 vs 807 g, respectively). This species is considered to be benthophagous, consuming mostly amphipods (34.9% by weight), polychaete worms (27.6%), decapod crustaceans (12.7%), and fishery discards (13.9%). Small skates (20–40 cm) fed mostly on amphipods (85.4%); medium‐sized (40–60 cm) ate amphipods (40.2%), polychaetes (29.1%) and decapods (19.3%); the largest individuals (>60 cm) consumed fishery offal (27.9%) and less amphipods (26.6%) and polychaetes (27.7%). Preliminary data on the maturation of the species in the western Bering Sea showed that males and females become mature at lengths above 61 cm TL.     

Feeding habits of the Magellan skate: effects of sex, maturity stage, and body size on diet

The aim of this study was to evaluate the effects of sex, maturity stage, and body size on the diet of the Magellan skate, Bathyraja magellanica, in the Southwest Atlantic off Argentina, by examining stomach contents using a multiple hypothesis modeling approach. Relationships between the number of prey and sex, maturity stage, and total length (TL) were assessed by built generalized linear models (GLM). Furthermore, we tested whether there was a threshold size at which B. magellanica started or quit consuming a given prey. The overall diet of B. magellanica was mainly consisted of teleosts, followed by amphipods, isopods, and decapods. Ontogenetic diet shifts were independent of sex and maturity stage. However, discrete shifts in diet with TL were found, with individuals larger than 554 and 623 mm TL ceasing to consume amphipods and isopods, respectively. The consumption of teleosts progressively increased with increasing predator size. Likewise, ontogenetic shifts in foraging behavior were also observed with smaller individuals showing specialization on amphipods with larger specimens consuming teleosts. These results confirm that ontogenetic shifts in diet of B. magellanica are more a function of predator size rather than any other life-history traits. We propose that these food shifts are probably related to morphological limitations and abilities associated with feeding habits of skate, so when specimens of B. magellanica reach an optimum body size, they may have access to higher quality trophic resources. Our results suggest that evaluating the importance of life-history stages on the feeding habits of a species is essential for understanding how that species exploits food resources, which, in turn, is an important factor in developing a suitable plan of marine ecosystem conservation.     

Feeding ecology of the estuarine dolphin (Sotalia guianensis) on the coast of Rio Grande do Norte,Brazil

We analyzed the stomach contents of 40 estuarine dolphins, Sotalia guianensis (van Benédén 1864), beached on the coast of Rio Grande do Norte, Brazil, between February 2000 and February 2007. A total of 223 prey items were identified, including 18 species of teleosts and 5 species of cephalopods. The index of relative importance (IRI) showed that Larimus breviceps, Haemulon plumieri, Lutjanus synagris, Trichiurus lepturus, Mugil curema, and Diapterus rhombeus were the six most important species. The IRI showed that L. breviceps was the main prey for both adults and the young. H. plumieri was the most important for the males and T. lepturus for the females. Seven species of teleosts and two of cephalopods were recorded in the diet of estuarine dolphins for the first time in the country. Our results suggest that the estuarine dolphin can be a feeding specialist and that foraging activity occurs mainly in estuarine areas, where the animals can use passive listening to detect prey.     

Feeding of the Brazilian sharpnose shark Rhizoprionodon lalandii (Müller & Henle, 1839) from southern Brazil

Stomach contents of 175 Brazilian sharpnose sharks, Rhizoprionodon lalandii, were examined to assess their feeding habits. Caught by artisanal fishers between June 2006 and July of 2007 at Ipanema Beach, on the central coast of the state of Paraná, southern Brazil, R. lalandii appears to be piscivorous (89.7% IRI), feeding mainly on families of Clupeidae (5.1% IRI) and Sciaenidae (3.7% IRI) and secondarily on squid Lolliguncula brevis (8.6% IRI). Diets were similar between sexes and seasons. However, there were ontogenetic changes in their diets. The major items for neonates were Penaeidae crustaceans (%IRI = 56.4) and Engraulidae fishes (%IRI = 29.3); for the juveniles, Sciaenidae (%IRI = 62.5) and Clupeidae (%IRI = 19.7), and for adults Clupeidae (%IRI = 45.8) and Sciaenidae (%IRI = 15.9). Only neonate sharks consumed crustaceans, whereas all size classes consumed cephalopods and teleosts. Neonates had the lowest trophic level (TR = 3.8), with the diet of juveniles and adults the highest (TR = 4.2). Given its abundance and diet, R. lalandii may be an important predator of demersal and pelagic prey in coastal waters of Brazil.     

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.     

Spatial Segregation in Eastern North Pacific Skate Assemblages

Skates (Rajiformes: Rajoidei) are common mesopredators in marine benthic communities. The spatial associations of individual species and the structure of assemblages are of considerable importance for effective monitoring and management of exploited skate populations. This study investigated the spatial associations of eastern North Pacific (ENP) skates in continental shelf and upper continental slope waters of two regions: central California and the western Gulf of Alaska. Long-term survey data were analyzed using GIS/spatial analysis techniques and regression models to determine distribution (by depth, temperature, and latitude/longitude) and relative abundance of the dominant species in each region. Submersible video data were incorporated for California to facilitate habitat association analysis. We addressed three main questions: 1) Are there regions of differential importance to skates?, 2) Are ENP skate assemblages spatially segregated?, and 3) When skates co-occur, do they differ in size? Skate populations were highly clustered in both regions, on scales of 10s of kilometers; however, high-density regions (i.e., hot spots) were segregated among species. Skate densities and frequencies of occurrence were substantially lower in Alaska as compared to California. Although skates are generally found on soft sediment habitats, Raja rhina exhibited the strongest association with mixed substrates, and R. stellulata catches were greatest on rocky reefs. Size segregation was evident in regions where species overlapped substantially in geographic and depth distribution (e.g., R. rhina and Bathyraja kincaidii off California; B. aleutica and B. interrupta in the Gulf of Alaska). Spatial niche differentiation in skates appears to be more pronounced than previously reported.     

Neuropeptide Y-like immunoreactivity in the cardiovascular nerve plexus of the elasmobranchs Raja erinacea and Raja radiata

Summary The distribution of nerves showing neuropeptide Y (NPY)-like immunoreactivity in the cardiovascular system of elasmobranchs and teleosts has been investigated. Two species of teleosts, the rainbow trout (Salmo gairdneri) and the Atlantic cod (Gadus morhua), and three species of elasmobranchs, the spiny dogfish (Squalus acanthias), the little skate (Raja erinacea) and the starry ray (Raja radiata), were used in this study. An innervation of the cardiovascular system by an NPY-like substance was found only in the two species of Raja. A rich innervation was encountered in these skates, with the highest density of fibres in the wall of the ventricle, the conus arteriosus, the coeliac artery and smaller mesenterial vessels. In the vessels, the fibres formed a plexus at the adventitio-mediol border. Few fibres were found in the walls of the dorsal aorta, the sinus venosus and the atrium, and no fibres were observed in the walls of the ventral aorta. Falck-Hillarp fluorescence histochemistry showed the presence of a rich innervation of arteries and arterioles of the gut by catecholamine-containing nerve fibres.     

Observations on the electric organ discharge of two skate species (Chondrichthyes: Rajidae) and its relationship to behaviour

Synopsis The electric organ discharge (EOD) of the little skate,Raja erinacea and winter skate,R. ocellata was recorded both from isolated individuals and from small groups using methods that allowed for the identification of individuals producing EODs. Pulse duration, train lengh, frequency, and pulse patterns are characterized and correlated with behaviour. The two species,R. erinacea andR. ocellata, were found to have characteristically different EOD pulse durations of 70 ms and 217 ms respectively. Isolated skates rarely discharged whereas groups of skates were found to discharge regularly. The EOD was evoked by tactile prodding, physical contact with other skates and electrical stimulation. Skates also discharged reflexively in response to an artificially induced head-positive DC stimulus, sine wave and monopolar square pulses. During approach and contact, skates responded to each other with interacting EOD displays. EOD interaction and pulse duration differences between other species suggest a possible intra-specific communication function of the EOD inRaja.     

Age and growth of big skate (Raja binoculata) and longnose skate (R. rhina) in the Gulf of Alaska

In 2003, big skates, Raja binoculata, and longnose skates, Raja rhina, were the target of a commercial fishery around Kodiak Island in the Gulf of Alaska (GOA) for the first time. The sudden development of a fishery for these species prompted the need for improved life history information to better inform fishery managers. Due to the selective nature of the skate fishery, mostly larger individuals were captured. Back-calculation from skate vertebral measurements was used to estimate size-at-age for younger skates. Because back-calculated age-length data within individuals were highly correlated, bootstrap resampling methods were used to test for differences between male and female growth curves. Results from bootstrapping indicated that differences between male and female growth were statistically significant for both species. This investigation indicates that growth of big skates in the GOA (max size 178 cm total length, max age 15 years) is similar to that in California, but different from that in British Columbia. For longnose skates, our GOA results agree with those reported in British Columbia, but were considerably older (max size 130 cm, max age 25 years) than those reported in California, which may not be surprising because longnose skates in the present study were generally larger. This life history information suggests that both big and longnose skates are at risk of unsustainable exploitation by targeted fisheries.     

Diet of two commercially important shark species in the United Arab Emirates: milk shark,Rhizoprionodon acutus (Rüppell, 1837), and slit‐eye shark,Loxodon macrorhinus (Müller & Henle, 1839)

The diets of the milk shark, Rhizoprionodon acutus, and the slit‐eye shark, Loxodon macrorhinus, landed from the artisanal fishery in the Arabian Gulf waters of the United Arab Emirates were investigated to determine their dietary preferences. Stomach contents from 57 milk sharks and 53 slit eye sharks were collected from Abu Dhabi (R. acutus, n = 23), Dubai (R. acutus, n = 5; L. macrorhinus, n = 15) and Ras Al Khaimah (R. acutus, n = 29; L. macrorhinus, n = 38) during fishery surveys from January to May 2012. Prey items were identified to the lowest possible taxonomic level, grouped into five categories including ‘teleost fish’, ‘cephalopods’, ‘crustaceans’, ‘invertebrates’, and ‘other’. The diets of both species were described using the numeric, frequency and weight methods, and the index of relative importance (IRI). The majority of stomachs for both species had food, with 66.6% of milk shark stomachs and 90.5% of slit‐eye shark stomachs containing prey items, both dominated by small teleosts. Rhizoprionodon acutus fed on a wide variety of teleost species, primarily Engraulidae (anchovies) (28%), Gerreidae (mojarras) (5.6%) and Carangidae (jacks) (1.6%) with occasional crustacean and cephalopod prey (8%). On the other hand, L. macrorhinus seemed to have a preference for one species in terms of teleosts (anchovies) (35.1%) and fed on a wider variety of crustaceans and cephalopods (22.6%). There was little overlap in the diets of these two species, suggesting that they may either be using different habitats or that in these waters, the milk shark is a generalist species while the slit‐eye is a specialist feeder.     

Diet of the milk shark,Rhizoprionodon acutus (Rüppel, 1837) (Chondrichthyes: Carcharhinidae), from the Senegalese coast

The diet of the milk shark, Rhizoprionodon acutus, from the Senegalese coast (12°30′N–14°45′N) was investigated in 3600 specimens with total lengths ranging from 44 to 113 cm for females and from 45 to 110 cm for males. Conducted from May 2010 to April 2011, the study revealed that of the 3600 stomachs examined, 577 contained food (16.03%). Cumulative prey diversity curves reached a stable level at 175 stomachs and thus the sample size was large enough to describe the overall milk shark diet. Relevant differences in the diet were found between sexes, maturity stages, sampling seasons and locations. The milk shark diet was composed of teleosts, crustaceans, mollucs, nematodes, annelids, and unidentified invertebrates. R. acutus seemed to have a preference for teleosts (98.75% in terms of IRI). The trophic level of milk shark calculated in this study was 4.2.     

Comparative analysis of feeding habits and dietary niche breadth in skates: the importance of body size,snout length,and depth

Skates (Elasmobranchii, Rajiformes) are a morphologically conservative group of bentophagous chondrichthyan fishes with a high degree of endemism, that occur on marine soft bottoms. Subtle morphological aspects and bathymetric distribution are traits that vary among skate species that could have implications for their feeding ecology. We test how body size, snout length and bathymetric distribution influence the feeding habits and dietary niche breadth in skates using data on 71 species taken from the literature. We hypothesized that snout length has an effect on diet composition. We also hypothesized that dietary niche breadth increases with increasing depth range and decreases with increasing body size of skate species. Generalized additive models for location scale and shape were fitted with taxonomic level (genera nested within family) included as a random effect term in each model. A model selection approach to test the level of support for alternative models was applied. We found that skate species that forage on large prey have the largest body size and skate species with the smallest body size prey on small and medium-sized invertebrates. The results indicated that body size has an effect on feeding habits of skates, whereas an effect of snout length was not supported. Bathymetric variables have an effect on the diet of skates. Our prediction that dietary niche breadth increases with increasing depth range and decreases with increasing body size of skate species was supported in part: in a first phase the relationship between dietary niche breadth and body size is positive, then in a second phase, including species larger than 1000 mm total length, the relationship become negative.     

Biodiversity and systematics of skates (Chondrichthyes: Rajiformes: Rajoidei)

Skates (Rajiformes: Rajoidei) are a highly diverse fish group, comprising more valid species than any other group of cartilaginous fishes. The high degree of endemism exhibited by the skates is somewhat enigmatic given their relatively conserved body morphology and apparent restrictive habitat, e.g. soft bottom substrates. Skates are primarily marine benthic dwellers found from the intertidal down to depths in excess of 3,000 m. They are most diverse at higher latitudes and in deepwater, but are replaced in shallower, warm temperate to tropical waters by stingrays (Myliobatodei). The number of valid skate species has increased exponentially, with more species having been described since 1950 (n = 126) than had been described in the previous 200 years (n = 119). Much of the renaissance in skate systematics has largely been through the efforts of a few individuals who through author–coauthor collaboration have accounted for 78 of the 131 species described since 1948 and for nine of 13 genera named since 1950. Furthermore, detailed regional surveys and accounts of skate biodiversity have also contributed to a better understanding of the diversity of the skates. A checklist of the living valid skate species is presented.     

Food habits of the sandpaper skate, Bathyraja kincaidii (Garman, 1908) off central California: seasonal variation in diet linked to oceanographic conditions

The stomachs of 130 sandpaper skates, Bathyraja kincaidii (Garman, 1908), were sampled from off central California to determine their diet composition. The overall diet was dominated by euphausiids, but shrimps, polychaetes and squids were also important secondary prey. A three-factor MANOVA demonstrated significant differences in the diet by sex, maturity status and oceanographic season using numeric and gravimetric measures of importance for the major prey categories. These three main factors explained more variation in diet than interactions between the factors, and season explained the most variance overall. A detailed analysis of the seasonal variation among the prey categories indicated that abundance changes in the most important prey, euphausiids, were coupled with seasonal changes in the importance of other prey. When upwelling occurred and productivity was great (Upwelling and Oceanic seasons), euphausiids were likely highly abundant in the study area and were the most important prey for B. kincaidii. As productivity declined (Davidson Current season), euphausiids appeared to decrease in abundance and B. kincaidii switched to secondary prey. At that time, gammarid amphipods and shrimps became the most important prey items and polychaetes, mysids and euphausiids were secondary.     

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.     

Southern African skate biodiversity and distribution

The skates (Family Rajidae) have 12 genera and possibly 28 species off southern Africa (southern Angola, Namibia, South Africa and Mozambique). The geographic and bathymetric distribution and the taxonomic composition of the southern African skate fauna are analysed and the distribution mapped. The southern African skate fauna is best known off the temperate west coast of South Africa from the intertidal to approximately 1,200 meters, but poorly known below 1,200 m and sketchily known in warm-temperate and tropical parts of the area. Southern African skates of the temperate continental shelves above 100 m are not diverse and regularly include one species of the genus Dipturus, one species of Leucoraja, two species of Raja (including R. straeleni, the most abundant skate in southern African waters) and the giant skate Rostroraja alba. All of these skates are ‘shelf overlap’ species that range onto the outer shelves and uppermost slopes, and none are confined to inshore environments. Skate diversity increases on the outer shelves and upper slopes. At least half of the skate species are endemic to the southern African region; other species also occur off East or West Africa, a few extend to European waters, and records of one species, Amblyraja taaf, appear to be of strays from nearby sub-Antarctic seas. The genus Bathyraja and softnose skate group (Arhynchobatinae) are surprisingly limited (a single species) in deep-water off southern Africa (unlike other regions including the Antarctic), and almost all of southern African skates are members of the Rajinae. Amongst rajines, the tribes Amblyrajini (Amblyraja, two species, Leucoraja, two species, and Rajella, five species) Rajini (Dipturus, six species, Okamejei, one species, Raja, two species, and Rostroraja, one species), and Anacanthobatini (Anacanthobatis, two species, and Cruriraja, three species) predominate, while Gurgesiellini has a species of Neoraja and possibly two of Malacoraja.     

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.