Two new North Pacific skates (Rajidae) and a revised key toBathyraja in the area

Two new species of the skates,Bathyraja hubbsi andB. pseudoisotrachys are described from the North Pacific. Among the North PacificBathyraja, B. hubbsi most closely resemblesB. violacea Suvorov in possessing a disc without thorns, a median tail thorns irregularly spaced, and minute fine prickles covering the entire dorsal surface. The two species are distinguished from each other by their dorsal color pattern, proportional measurement of tail in relation to total length, clasper components, and number of precaudal vertebrae.Bathyraja pseudoisotrachys has long been confused withB. isotrachys (Günther) by many Japanese authors, but is distinguished from other North PacificBathyraja by dorsal squamation and unique clasper components such as two clefts, terminal bridge and ridge.Bathyraja kincaidii Garman is synonymized with the valid speciesB. interrupta Gill et Townsend.Bathyraja abyssicola Gilbert andB. trachura Gilbert are redescribed in detail. Provisionally, 21 species of the genusBathyraja are recognized in the present study from this region, and a revised key is given to the North Pacific species of the genus.     

Identification of multiple nursery habitats of skates in the eastern Bering Sea

The use of more than a single nursery habitat type is examined for oviparous elasmobranchs using data summarized from studies conducted on the Alaska skate Bathyraja parmifera and the Aleutian skate Bathyraja aleutica in the eastern Bering Sea. The eastern Bering Sea skate species use two discrete areas as nurseries, one for egg deposition and a second for newly emergent juveniles. Egg deposition sites were located along the outer shelf and upper slope near canyons in the eastern Bering Sea. Newly emergent juveniles were found along the outer and middle shelf for B. parmifera and deep‐slope for B. aleutica, suggesting that habitat used by newly emergent juvenile skates is distinct from habitat used for egg deposition and embryo development. In reviewing many studies on oviparous elasmobranchs, similar patterns emerge of habitat use during their early life history. To distinguish these distinct habitats, appropriate terminology is proposed. Egg case nursery is suggested for areas of egg deposition and juvenile nursery is suggested for areas where juveniles aggregate after emergence. Criteria to describe each habitat type are outlined.     

Molecular systematics of the skate subgenus Arctoraja (Bathyraja: Rajidae) and support for an undescribed species, the leopard skate, with comments on the phylogenetics of Bathyraja

Sequence variability in the cytochrome c oxidase I (COI) gene from 226 samples of the species previously considered Bathyraja parmifera (Rajidae) revealed three distinct haplotypes, one of which represents an undescribed species, the leopard skate. Further genetic examination of four closely related North Pacific and Bering Sea skate species, Bathyraja parmifera, B. simoterus, B. smirnovi, and the leopard skate in comparison with 19 related species indicates that together these four species comprise the subgenus Arctoraja. Phylogenetic analysis suggests that Arctoraja is monophyletic, but that the genus Bathyraja may be paraphyletic due to the phylogenetic position of Rhinoraja.     

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.     

Age and growth of the roughtail skate Bathyraja trachura (Gilbert 1892) from the eastern North Pacific

This study provides the first published age estimates for the roughtail skate, Bathyraja trachura. Age and growth characteristics of B. trachura, a poorly-known deepwater species, were determined from samples collected along the continental slope of the contiguous western United States. A new maximum size was established at 91.0 cm TL. Age was determined using a traditional structure (vertebral thin sections) with widespread application on multiple skate species and a non-lethal structure (caudal thorns) recently used for age analysis on skate species. Caudal thorns were determined not to be a useful ageing structure for this species based on poor precision and significantly lower age estimates when compared to age estimates from vertebral thin sections. The best model for describing growth of B. trachura was the two parameter VBGF, assuming annual vertebral band deposition and using length-at-age data. Although females grew slower and reached a larger maximum size than males, their growth was not statistically different (ARSS; P = 0.90); therefore, data were pooled (L_∞ = 99.38, k = 0.09). Annual band deposition was found to be a reasonable assumption for this species, but has yet to be validated. The maximum age estimated for B. trachura was 20 years for males and 17 years for females using vertebral thin sections.     

Age, growth, maturity, and mortality of the Alaska skate, Bathyraja parmifera, in the eastern Bering Sea

The Alaska skate, Bathyraja parmifera, is the most abundant species of skate on the eastern Bering Sea shelf, accounting for over 90% of total skate biomass. However, little is known regarding the life history of this species despite its common occurrence as bycatch in several Bering Sea fisheries. This is the first study to focus on the age and growth of B. parmifera. From 2003 to 2005, more than one thousand specimens were collected by fisheries observers and on scientific groundfish surveys. Annual banding patterns in more than 500 thin sections of vertebral centra were examined for age determination. Caudal thorns were tested as a potentially non-lethal ageing structure. Annual band pair deposition was verified through edge and marginal increment analyses. A three-parameter von Bertalanffy growth function and a Gompertz growth function were fit to observed length-at-age data. Both models provided significant fits, although the Gompertz function best described the overall pattern of growth in both males and females, based upon statistical criteria and parameter estimates. Age and size at 50% maturity were 9 years and 92 cm TL for males and 10 years and 93 cm TL for females. The maximum observed ages for males and females were 15 years and 17 years, respectively. Estimates of natural mortality (M) ranged from 0.14 to 0.28, and were based on published relationships between M and longevity, age at maturity, and the von Bertalanffy growth coefficient. Due to these life history characteristics and a lack of long-term species-specific stock data, a conservative management approach would be appropriate for B. parmifera.     

New data on sexual dimorphism and reproductive biology of Alaska skate <Emphasis Type="Italic">Bathyraja parmifera</Emphasis> from the northwestern Pacific Ocean

New data on sexual dimorphism and features of reproductive biology of Alaska skate Bathyraja parmifera from the western Bering Sea and the northern Sea of Okhotsk are presented. Based on comparative analysis of 11 external morphological characters, the males differ from the females in the length of the disc and snout length (to the eyes, to the mouth, and to the nostrils). In the western Bering Sea, 50% of the females and 50% of the males reach the first sexual maturity at 84.5 cm TL and 85.2–90.2 cm TL, respectively; in the Sea of Okhotsk, 50% of the males reach their first sexual maturity at 80.8–83.8 cm TL (the assessment depends on the features of the method). Among smaller skates (less than 45 cm TL), the males prevail, but the proportion of the females increases in larger size groups; the sex ratio becomes equal in the exemplars 60–80 cm TL, but the males prevail again among the largest fishes (more than 80 cm TL).     

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.     

Trophic niche partitioning of five skate species of genus Bathyraja in northern and central Patagonia,Argentina

Overexploitation of marine communities can lead to modifications in the structure of the food web and can force organisms like elasmobranchs to change their feeding habits. To evaluate the impact that fisheries have on food webs and on the interactions between species, it is necessary to describe and quantify the diet of the species involved and follow it through time. This study compares the diet of five skate species using the data obtained from the by-catch of the Argentine hake (Merluccius hubbsi) fishery in north and central Patagonia, Argentina. Diet composition was assessed by analysing the digestive tract contents and trophic overlapping between species of the genus Bathyraja: Bathyraja albomaculata, Bathyraja brachyurops, Bathyraja macloviana, Bathyraja magellanica and Bathyraja multispinis. A total of 184 stomachs were analysed. The diets of B. albomaculata and B. macloviana mainly comprised annelids, whereas that of B. brachyurops primarily comprised fish, including hake heads discarded by the fishery. The diets of B. magellanica and B. multispinis were largely based on crustaceans. Despite the morphological similarities and their shared preference for benthic habitats, no complete diet overlaps were found between the different species. These results suggest that these skate species have undergone a process of diet specialisation. This is a common feeding strategy that occurs to successfully eliminate competition when resources are limited, which corresponds to the conditions found in an environment being affected by the pressures of overfishing.     

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.     

Distribution and some biological features of skate <Emphasis Type="Italic">Bathyraja parmifera</Emphasis> (Rajidae,Rajiformes) in the northwestern part of the Sea of Japan

In the northwestern part of the Sea of Japan, skate Bathyraja parmifera in catches of the bottom trawl occurs at depths of 40 to 830 m at the near-bottom temperature of 0.4–4.9°C. Throughout the year, the skate performs migrations within the continental slope and, in the feeding period, inhabits mainly the depth range of 300 to 500 m. Maximal average density of aggregations of B. parmifera is observed in waters of northern Primorye; minimal average density is observed in Peter the Great Bay and on the Yamato Bank. In catches, skates with a length of 17–130 cm and a weight of 0.5–18.8 kg are recorded; however, fish with a length of 80–115 cm dominate. Individuals with a length of up to 40 cm feed mainly on amphipods, and larger immature and adult individuals feed on decapods, cephalopods, and fish. The value of the daily ration of B. parmifera with its growth decreases from 2.4 to 0.9% of the body weight. Specific ecological features of B. parmifera are a cause of its considerable underfishing by the bottom trawl; therefore, the total resource of this species in the considered region may be at the level of maximal estimates obtained during surveys and comprise no less than 20000 t.     

Size and age estimates at sexual maturity for the little skate Leucoraja erinacea from the western Gulf of Maine,U.S.A.

Size and age estimates at sexual maturity were determined for 162 male and 273 female little skates Leucoraja erinacea collected from the western Gulf of Maine. Maturity ogives suggest that 50% maturity in females occurs at age 9·5 years and 480 mm total length (L_T), whereas 50% maturity in males occurs at a slightly younger age of 7·7 years and smaller size of 460 mm L_T. Age estimates were made from 389 L. erinacea ranging in size from 93 to 570 mm L_T. The index of average per cent error and age‐bias plots indicated that the ageing methods were precise and non‐biased. Additionally, annual periodicity of band formation was validated with oxytetracycline in eight individuals (three males and five females) ranging in age from 3 to 12 years. In conclusion, results from this study indicate that L. erinacea exhibits characteristics that make other elasmobranch populations highly susceptible to overexploitation.     

Reproductive biology of the brown smoothhound shark Mustelus henlei,in the northern Gulf of California,México

Female brown smoothhound sharks Mustelus henlei were found to reproduce annually. A mature female carried both developing oocytes in the ovary and developing embryos in the uteri concurrently for c. 1 year. A great variability in the size of embryos was recorded each month, and the maximum embryo sizes were found from late January to mid‐March. The largest oocytes in mature females were observed in mid‐March. Gestation lasted c. 10 months. A linear relationship between maternal total length (L_T) and the number of pups per litter (litter size one to 21) was estimated. Birth L_T was reached in c. 280 mm. Females and males matured at 570–660 and 550–560 mm L_T, respectively. Difference in the litter size among Californian coast (one to 10) and northern Gulf of California (one to 21) populations existed for this smoothhound shark.     

Reproductive biology of the southern thorny skate Amblyraja doellojuradoi (Chondrichthyes,Rajidae)

The total lengths (L_T) of 193 males (209–556 mm) and 130 females (275–515 mm) of Amblyraja doellojuradoi, a commercial by‐catch species on the Argentinean continental shelf, which are increasingly retained, were analysed. No sexual dimorphism was observed in the L_T at which 50% of individuals were sexually mature; males matured at 448 mm and females at 411 mm, c. 80 and 82% of maximum L_T. The hepato‐somatic index was similar among sexes, but significantly different between maturity stages, being lower in mature than immature specimens. Males had no seasonal difference in the hepato‐somatic index and females had the lowest index in autumn. The gonado‐somatic index was lower in males than in females and significantly higher in mature than immature specimens of both sexes. Males had the highest index in autumn and females had no seasonal difference. Collectively, these results would indicate that A. doellojuradoi breeds in autumn.     

Age and size at sexual maturity of the smooth skate Malacoraja senta from the western Gulf of Maine

Age and size at sexual maturity was determined for 185 male and 96 female smooth skates Malacoraja senta (ranging in size from 370 to 680 mm total length L_T), collected from the western Gulf of Maine. Maturity ogives for males, based on clasper length, testis mass and the proportion of mature spermatocysts in the testes, suggest that 50% maturity occurs between 9 and 10 years and 560 mm L_T. Maturity ogives for females, based on ovary mass, shell‐gland mass and maximum follicle size, suggest that 50% maturity occurs at age 9 years and 540 mm L_T.     

Size at maturity and egg capsules of the softnose skates Bathyraja brachyurops (Fowler, 1910) and Bathyraja macloviana (Norman, 1937) (Elasmobranchii: Rajidae) in the SW Atlantic (37°00'–39°30'S)

The softnose skates Bathyraja brachyurops and Bathyraja macloviana represent an important portion of the skate catches of the Uruguayan trawling fleet in the southwestern Atlantic. From March to October 2004, specimens of these species were collected at 75–200 m depth range in the area situated between latitudes 37°00'–39°30'S. For B. brachyurops , total length at which 50% of the specimens were retained by the gear was 68.0 cm for both sexes; T _L50 was estimated at 65.4 cm for males and 67.0 cm for females. For B. macloviana, total length at which 50% of the specimens were retained was 56.0–57.0 cm for both sexes; T _L50 was estimated at 53.5 cm for males and 52.0 cm for females. Egg capsule length varied from 79–91 mm in B. brachyurops and 69–75.5 mm in B. macloviana . In both species, capsules displayed striated surfaces and similar gross morphology, although egg capsules of B. macloviana had more robust anterior horns and a smaller size than those of B. brachyurops . Egg capsules of the latter also exhibited microscopical prickles. Capsule edges were laterally keeled with a groove along the keel, and a straight and transverse velum was present in the egg capsules of both species.     

Life history of the river goby Glossogobius callidus (Teleostei: Gobiidae)

The river goby Glossogobius callidus is native to freshwater and estuarine habitats in South Africa. Individuals [21.1–144.4 mm total length (L_T)] were sampled from impoundments in the Sundays River Valley, Eastern Cape, from February 2014 to March 2015. The largest female was 137.2 mm L_T, and the largest male was 144.4 mm L_T. Length-at-50% maturity was 75.2 ± 2.1 mm L_T for males and 76.2 ± 2.0 mm L_T for females. Absolute fecundity was 1028.2 ± 131.7 oocytes per fish, and relative fecundity was 50.1 ± 18.1 oocytes per gram. The spawning season extended from October to December. Fish were aged using sectioned sagittal otoliths. The growth zone periodicity was validated using edge analysis. Longevity was more than 7 years for females and more than 6 years for males. Length-at-age was similar for the two sexes and was best described using the von Bertalanffy growth model as L_t = 74.7(1 − e^{–1.0(t + 0.1)}) mm L_T for the entire population. Using the population age structure, the mortality rate was estimated at 1.3 per year.     

Arctic skate Amblyraja hyperborea preys on remarkably large glacial eelpouts Lycodes frigidus

During scientific surveys on the continental slopes north‐west of Spitsbergen and off north‐east Greenland (c. 600 and 1000 m depths), two female Arctic skates Amblyraja hyperborea were caught while swallowing extraordinary large individuals of glacial eelpout Lycodes frigidus. The total length (L_T) of the prey constituted 50 and 80% of the L_T of the skates, which reveal that A. hyperborea are capable predators of fishes of surprisingly large relative size.     

Exploring nursery sites for oviparous chondrichthyans in the Southwest Atlantic (36°S–41°S)

The aim of this paper was to explore egg-laying areas of oviparous chondrichthyans occurring in the northern part of the Southwest Atlantic Ocean (36°S–41°S) at between 50 and 200 metres depth and the Mar del Plata Canyon (from 200 to 3447?m). A total of 515 capsules were collected corresponding to 10 species. Four species accounted for 87.6% of the total catch: Psammobatis normani, Bathyraja macloviana, Amblyraja doellojuradoi and Bathyraja brachyurops. The remaining corresponded to Schroederichthys bivius, Psammobatis rudis, Zearaja chilensis, Bathyraja albomaculata, Psammobatis lentiginosa and Bathyraja sp. Most hauls were monospecific and relatively few hauls contained four or five species (southern part of the area, in sites located at 75 and 94–105?m depth). No egg capsules were recorded between 1712?m and 3447?m. Highest densities (>3000 capsules/km²) were mainly found between 39°46.2′ and 40°29.9′S from 85 to 105?m and they were also recorded near the Mar del Plata Canyon at 37°59.7′S and at 852?m. The highest value recorded was 12,326 capsules/km² (located at 40°18.9′S and 85.4?m). The highest densities for each of the most abundant species are discussed. The finding of high densities of egg cases near the Argentine shelf-break front indicates that many skate species use this area as a nursery site. The egg-laying areas explored here are the first reported sites of this kind in the Southwest Atlantic Ocean. Recognizing and protecting egg-laying habitats may be important steps to a long-term conservation of oviparous chondrichthyan populations.