Skate Species Research Articles

Interannual variability in the skate assemblage on the South Patagonian shelf and slope.

Observer data from the commercial fishery on the Patagonian shelf and slope around the Falkland Islands (home to an assemblage of >16 skate species (Rajiformes), for which commercial catches have been recorded since 1987), as well as survey data from an area closed to skate target fishing after exploitation, were summarized by species to examine changes in the population status of individual skate species. Total skate catch per unit effort increased significantly in the target fishery since 1994, and four species have made up >85% of all skate catch. Bathyraja brachyurops and Zearaja chilensis increased significantly in catch proportions and abundance from 1994 to 2013. Bathyraja albomaculata and Bathyraja griseocauda decreased significantly before rebounding with trends of increasing abundance. Concurrently, B. brachyurops and Z. chilensis showed decreasing trends in size at 50% maturity in areas where skates continue to be targeted commercially. The increasing abundances and concomitant reductions in size at maturity of B. brachyurops and Z. chilensis suggest either plasticity in life-history traits or a density-dependent growth response to fishing pressure. Bathyraja griseocauda decreased in size at 50% maturity in the area that was closed to skate target fishing, where it was initially larger, but only decreased to the same average size as in the commercially targeted areas. Bathyraja albomaculata and Z. chilensis are IUCN-listed as vulnerable and B. griseocauda is listed as endangered, but their abundance trends since 1994 indicate that these populations are not declining in Falkland waters.

The diversity, distribution and status of deep-water elasmobranchs in the Rockall Trough, north-east Atlantic Ocean.

Data from a scientific deep-water trawl fisheries survey in the north-east Atlantic were analysed to determine the spatial and bathymetric distribution of elasmobranch species and assess the change in relative abundance over the period 1998-2013. During this period, commercial fisheries for deep-water sharks went from being entirely unregulated, to being briefly managed, to being completely prohibited. A total of 22 species of shark and 10 species of skate were recorded between depths of 300 and 2030 m. All showed strong species-specific depth-related trends in abundance. Out of the 11 more common species, five showed no change in relative abundance over time, two (Centrophorus squamosus and Centroselachus crepidater) declined significantly and four increased in relative abundance (Apristurus aphyodes, Apristurus microps, Galeus melastomus and Deania calcea). Assuming these populations were depleted by fisheries in the past, the current data do not suggest there has been an overall recovery. Positive signs for some species in the most recent years suggest movement or recruitment back into the area; however, it is of concern that two species continued to decline. There is a continued need to have precautionary management of these elasmobranch species, and the current ban on landing these species in European waters remains appropriate.

How is the morphology of the oviducal gland and of the resulting egg capsule associated with the egg laying habitats of Rajidae species?

Egg capsules of Rajidae species vary in size, shape and adornments. These differences can be used as a taxonomic tool to differentiate species. Furthermore, since egg capsule external morphology is adapted to the environment where they are laid, it can be used make inferences on egg-laying grounds. In the present study, oviducal glands and egg capsules of seven species of skates (Leucoraja naevus, Raja brachyura, Raja montagui, Raja microocellata, Raja miraletus, Raja clavata and Raja undulata) are described and compared, and a taxonomic key developed. The resulting taxonomic key constitute a useful tool to identify skate species inhabiting Portuguese continental waters and, therefore, to improve our knowledge of egg laying habitatsproviding useful information for conservation and management.

In situ observations of a possible skate nursery off the western Antarctic Peninsula.

A dense aggregation of skate egg cases was imaged during a photographic survey of the sea floor along the western Antarctic Peninsula in November 2013. Egg cases were noted in a narrow band between 394 and 443 m depth. Although some skate species in other oceans are known to utilize restricted areas to deposit eggs in great numbers, such nurseries have not been described in the Southern Ocean.

Nutritional content, mercury, and trace element analyses of two skate (Rajidae) species in the Gulf of Alaska

Seafood is recognized as an important source of proteins and long chain omega-3 fatty acids. However, one of the primary concerns with seafood consumption is the level of heavy metals, particularly mercury, present in fish tissues, which may influence the demand of certain fishery products. We sampled muscle and liver tissues from 20 big (Beringraja binoculata) and 20 longnose (Raja rhina) skates collected near Kodiak and Cordova, Alaska, and analyzed their nutritional content (protein, moisture and lipid content and fatty acid profiles), heavy metal (mercury, arsenic, cadmium, lead) and trace element (selenium) load. Big and longnose skate muscle was composed of lean protein (14.7%±0.7% SD) with 1.2% (±0.4%) lipids and 83% (±0.8%) moisture. Skate livers were very high in lipids, between 52.5 and 57.5% and had high percentages of omega-3 fatty acids (30.2%). Mercury in these skates had mean levels of 0.21mg/kg, lower than average levels found in Pacific halibut (Hippoglossus stenolepis). Overall, the risk/benefit ratio of consuming skate muscle was slightly positive (3.62%) based on the balance of mercury toxicity and omega-3 fatty acid benefits. Big skates were overall more beneficial to consume, and only longnose skates from Cordova had a negative risk/benefit ratio. These data can be used by the fishing industry to understand current and future market demands for skate products, and to be aware of any health concerns of consuming Gulf of Alaska skates.

Using biological variables and reproductive strategy of the undulate ray Raja undulata to evaluate productivity and susceptibility to exploitation.

The present work provides a detailed analysis of the reproductive strategy of the undulate ray Raja undulata in Portuguese mainland waters. The species was found mostly between 30 and 40 m deep on sandy bottoms. Egg-laying sites were observed in the north, centre and south-west regions, mainly at depths below 30 m. The peak of the reproductive season occurred from December to May. Asynchrony between reproductively active females and males appeared to occur, although most adult males were capable of reproducing throughout the year. The estimated length at 50% maturity was 86·2 cm (8·7 years) and 76·8 cm (7·6 years) total length for females and males, respectively. The maximum potential fecundity was estimated to be 69·8 follicles per female per reproductive season, which are released in 4·7 batches of 15 follicles. The life-history and demographic parameters of R. undulata are similar to those of other skate species, while the potential rate of population increase (0·49) is above the published values for other elasmobranch species. With these new findings, this study makes an important contribution to the understanding of the life history of R. undulata, and provides a first evaluation of the productivity and susceptibility of the species to exploitation.

Having confidence in productivity susceptibility analyses: A method for underpinning scientific advice on skate stocks?

National and European shark conservation plans aim to manage elasmobranch stocks sustainably. However uncertainties and deficiencies with available data hamper traditional, quantitative assessment methods of stock status to inform those plans, and thus effective management. The International Council for the Exploration of the Sea (ICES) Expert Groups have explored a range of data deficient assessment methods that may be used to support management advice, including Productivity Susceptibility Analysis (PSA). This method was applied to the demersal elasmobranch fauna (21 species) of the Celtic Sea to explore how such approaches could inform the management of skates (Rajidae). This species complex is an important catch component for demersal trawl and gillnet fisheries and is currently managed under a mixed species total allowable catch (TAC). PSAs were conducted on both of these fisheries, by four experts from three countries to introduce independence, and to quantify the range in perceptions of each stock. Confidence scoring of attributes was incorporated and probability distributions generated to model uncertainty in the expert responses to susceptibility attributes. Results showed that three shark species (tope, Galeorhinus galeus; angel shark, Squatina squatina and spurdog, Squalus acanthias) were the most vulnerable species in both fisheries (a consequence of their life history strategy and large size), followed by two skates (otter trawl) and three skates (gillnet). All of these species have some form of restrictive management and, apart from tope, are either currently listed as prohibited species or have a zero TAC in the area. Blonde ray, Raja brachyura was ranked as the next most vulnerable member of the commercially exploited skate complex. This adaptation of the PSA approach enabled skate species of higher and lower risk to be ranked and thus inform where management efforts should be focussed, whilst giving a novel consideration to uncertainty through canvassing expert opinion.

New insights to discriminate between Sympterygia acuta Garman 1877 and Sympterygia bonapartii Müller & Henle, 1841 (Rajidae) of the Southwest Atlantic Ocean: on the use of geometric morphometrics and spinulation patterns

Fil: Orlando, P.. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales; Argentina

DNA barcoding of the northern Northeast Atlantic skates (Chondrichthyes, Rajiformes), with remarks on the widely distributed starry ray

The skate fauna in the northern Northeast (NE) Atlantic is poorly investigated, and misidentifications are common. Here, ‘DNA barcoding’ was used to analyse 105 specimens of 15 species previously reported from the area to investigate the occurrence of species. Of these 15 species, three were new to the region and confirmed with voucher specimens. Three previously reported taxa were not obtained from the study area, providing a total number of 12 skate species for the northern NE Atlantic. Only one specimen of the critically endangered Dipturus batis complex was found. It occurs frequently in the literature and commercial fisheries catch records, and we argue that the vast majority of these are misidentifications. Due to striking differences in Amblyraja radiata life history parameters across the North Atlantic, cryptic species diversity has previously been suspected. A total of 80 A. radiata cytochrome c oxidase subunit I sequences from across the North Atlantic were sampled, and the highest fixation index (FST) was found when maximising geographical distance (FST = 0.133). A lower index was found when grouped according to life history (FST = 0.067). These results are not strongly supportive for the occurrence of cryptic species.

A comparison of age and growth of the Bering skate, Bathyraja interrupta ( ), from two Alaskan large marine ecosystems

The Bering skate, Bathyraja interrupta, is commonly found in both the Bering Sea and Gulf of Alaska, where skates are a regular bycatch in commercial long line and bottom trawl fisheries. This is the first study to examine the age and growth of B. interrupta, and one of the few studies to directly compare growth parameters of a skate species between management regions. Ages were estimated based on band counts using the standard gross-sectioning method of vertebral centra preparation. Additionally, a preliminary investigation was conducted into the utility of the histological method. Several growth models were compared and the best-fit model for both regions was identified as the logistic model using the Akaike's Information Criterion approach. No significant difference was detected between sexes for any model in either region; however, a slight but significant difference was found between regions. The parameters of the logistic growth model based on the gross-sectioned preparation were L∞=82.80cm TL and k−1yr=0.23 for the eastern Bering Sea and L∞=86.94cm TL and k−1yr=0.20 for the Gulf of Alaska. Age estimates from the gross sectioned vertebral centra provided a minimum longevity of 19 years and 21 years, respectively. Estimates based on the histologically prepared samples provide overall slightly higher age estimates, but the same maximum age of 21 years.

Identification of Potential Essential Fish Habitats for Skates Based on Fishers’ Knowledge

Understanding of spatio-temporal patterns of sensitive fish species such as skates (Rajidae) is essential for implementation of conservation measures. With insufficient survey data available for these species in Portuguese Continental waters, this study shows that fishery-dependent data associated with fishers' knowledge can be used to identify potential Essential Fish Habitats (EFH) for seven skate species. Sites with similar geomorphology were associated with the occurrence of juveniles and/or adults of the same group of species. For example, sites deeper than 100m with soft sediment include predominantly adults of Raja clavata, and are the habitat for egg deposition of this species. Raja undulata and R. microocellata are the more coastal species, preferring sand or gravel habitats, while coastal areas with rocks and sand seabed are potential nursery areas for R. brachyura, R. montagui and R. clavata. The main output of this study is the identification of preferential fishing sites enclosing potential EFH for some species, associated with egg-laying and nursery grounds. The location of these areas will be considered for future seasonal closures, and studies will be conducted to evaluate the biological and socio-economic impacts of such measures. As in the past, fishermen will collaborate in the process of evaluating those impacts, since they have practical and applied knowledge that is extremely valuable for evaluating the advantages and disadvantages of such closures. In conclusion, this study is a first contribution to the understanding and identification of EFH for skate species, associated with nursery and egg deposition sites, with direct application to management.

Morphology Meets Molecules: A New Genus and Two New Species of Diphyllidean Cestodes From the Yellowspotted Skate,Leucoraja wallacei, from South Africa

Two morphologically disparate undescribed species of diphyllidean cestodes from the Yellowspotted skate, Leucoraja wallacei , from South Africa were included in a recent molecular phylogenetic study aimed at revising diphyllidean classification. From a molecular standpoint, these species were determined to be only distantly related to one another. One (originally referred to as Echinobothrium n. sp. 2) showed affinities with species of the batoid-parasitizing Echinobothrium sensu stricto and is described here as Echinobothrium marquesi n. sp. This species most closely resembles Echinobothrium joshuai, an affinity supported by the previous molecular study, but differs in the form of its "B" hooks and degree of overlap between bothria and cephalic peduncle. The other species (originally referred to as New genus n. sp. 1), although exhibiting the full complement of scolex armature, grouped among primarily shark-hosted genera, most of which lack or exhibit reduced scolex armatures. That unexpected result suggested that erection of a novel genus might be warranted, but morphological grounds supporting the action were not apparent at that time. The present study aimed to explore the morphology of this taxon in more detail. Light and scanning electron microscopy revealed this taxon to be exceptional in that its lateral hooklets, which are arranged in 2 clusters like those of Echinobothrium and Coronocestus, are unique in being arranged in anterior and posterior rows, rather than in a single row. Andocadoncum n. gen. is erected, with Andocadoncum meganae n. sp. as its type, to accommodate this taxon. A minor adjustment to the existing hook formula by presenting counts for the anterior hooklets (a) separately from the posterior hooklets (b) readily accommodates this new form. Leucoraja wallacei is distinctive among skates in hosting 2 different genera of diphyllideans. In the context of the previous molecular phylogenetic analyses, it appears that the presence of Andocadoncum n. gen. in this skate species may be the result of a host-switching event involving a weakly or unarmed shark-hosted taxon moving to a batoid-a shift that was likely accompanied by a return to the fully armed condition seen, for example, in other batoid-parasitizing members of the order.

Life history of Bathyraja trachura from the eastern Bering Sea, with evidence of latitudinal variation in a deep-sea skate species

Although many skates possess life history characteristics that may make them vulnerable to exploitation, the detailed biological information needed to enact effective management schemes is lacking for most species. The objectives of this study were to provide age, growth, and maturity estimates for the roughtail skate, Bathyraja trachura, from the eastern Bering Sea. Maximum age was estimated at 36 yr based on band pair counts in vertebral centra. Of the four growth models applied, the logistic model provided the best description of growth (asymptotic total length = 911 mm; growth coefficient = 0.131 yr−1). There was no evidence of difference in growth between males and females. Females attained maturity at larger sizes and older ages than males. The median size-at-maturity was estimated at 741 mm total length (TL) for males and 796 mm TL for females; median age-at-maturity was estimated at 21.1 yr and 24.7 yr for males and females, respectively. The results of this study may indicate a latitudinal pattern in size and growth, with individuals from the eastern Bering Sea growing more slowly and reaching higher maximum ages than previously reported for specimens collected off the western coast of the continental United States.

Length–Length and Width–Length Conversion of Longnose Skate and Big Skate Off the Pacific Coast: Implications for the Choice of Alternative Measurement Units in Fisheries Stock Assessment

AbstractMost traditional fisheries models have size or stage–age relationships as their foundation. Two long‐established metrics used are FL and TL. Body length measurements are linked to age, maturity, and fecundity. Obtaining FL or TL measurements are not always convenient, especially for larger fish (>100 cm TL). Two skate species, Longnose Skate Raja rhina and Big Skate R. binoculata, were collected from commercial fishery landings along the coast of Washington State and length–length or width–length conversions were investigated for four metrics of size: TL, tail only length (TOL), interspiracular width (ISW), and interorbital width (IOW). Relationships between TL and alternative measures were examined using model II regression analyses and confirmed a strong linear relationship in all cases (R 2 > 0.94). It is likely that both TL and TOL measures have errors due to worn‐off tail portions; conversely, the ISW is nonlaborious and convenient to obtain. The ISW metric has a smaller coefficient of variation compared with IOW if we assume that both metrics have the same distribution of measurement errors. We recommend ISW as the preferred alternative metric in the assessment of large skate. The ISW will probably yield measurements that are equivalent, or potentially superior, to traditional metrics because the measurement is taken from body parts devoid of worn‐off portions.Received December 30, 2011; accepted June 13, 2013

DNA barcoding unveils skate (Chondrichthyes: Rajidae) species diversity in ‘ray’ products sold across Ireland and the UK

Skates are widely consumed across the globe, but many large species are subject to considerable concern regarding their conservation and management. Within Europe such issues have recently driven policy changes so that, for the first time, reports of skate landings now have to be made under species-specific names. Total allowable catches have also been established for many groups, which have been set to zero for a number of the most vulnerable species (e.g., Dipturus batis, Raja undulata and Rostoraja alba). Whilst accurate species identification has become an important issue for landings, the sale of skates is still usually made under a blanket term of “skate” or “ray”. The matter of identifying species of skate is further complicated by their morphologically conservative nature and the fact that they are commercially valued for their wings. Thus, before sale their bodies are usually discarded (i.e., “winged”) and often skinned, making morphological identification impossible. For the first time, DNA barcoding (of the mitochondrial COI gene) was applied to samples of skate wings from retail outlets across the British Isles, providing insight into which species are sold for consumption. A total of 98 wing samples were analysed, revealing that six species were sold; blonde ray (Raja brachyura), spotted ray (Raja montagui), thornback ray (Raja clavata), cuckoo ray (Leucoraja naevus) small-eyed ray (Raja microocellata) and shagreen ray (Leucoraja fullonica). Statistical testing demonstrated that there were significant differences in the species sold in the distinct retail groups which suggests complex drivers behind the patterns of sale in skates. The results also indicate that endangered species are not commonly being passed on to consumers. In addition, the practice of selling skate wings under ambiguous labels is highlighted as it makes it extremely difficult for consumers to exercise a right to avoid species of conservation concern. Interestingly, a single retailer chain labelled their wings as originating from three smaller-growing species (generally to be considered of lower conservation concern); of the six samples analysed from this company a third were mislabelled and originated from the thornback ray (a larger species that is currently undergoing population declines).

New African Species ofEchinobothrium(Cestoda: Diphyllidea) and Implications for the Identities of their Skate Hosts

Two new species of diphyllidean cestodes of the genus Echinobothrium, each hosted by a different skate species in the Raja miraletus complex, are described. Echinobothrium mercedesae n. sp. is described from R. cf. miraletus 2 off Senegal. Echinobothrium yiae n. sp. is described from R. cf. miraletus 1 off South Africa. Both species are small worms that differ from their 29 described congeners in the combination of number of cephalic peduncle spines per column, hook formula, number and arrangement of testes, and arrangement of vitelline follicles. They are easily distinguished from one another in that whereas the vitelline follicles of E. yiae n. sp. are circumcortical, they are lateral in E. mercedesae n. sp., and also in number of cephalic peduncle spines per column (14-17 vs. 10-12). Echinobothrium yiae n. sp. is also unusual in that the cephalic peduncle spines stop short of the anterior margin of the peduncle. In addition, although the paucity of available material precluded their formal description, evidence of 2 additional new species parasitizing R. miraletus also from Senegal is presented. In combination these worms provide support for the interpretation that what is currently recognized as Raja miraletus actually consists of a complex of geographically restricted species, rather than a polymorphic species of multiple parapatric or allopatrically distributed populations. This interpretation is not only supported by previously published molecular data, but also by newly collected morphological data involving differences in the color patterns of disc ocelli among host specimens of the 3 forms available as a result of digital efforts to ensure the accuracy of host identifications, which are also presented here.

Analysis of body shape for differentiating among species of rajids

This study sought to differentiate the species of skates encountered in Gulf St Vincent (GSV), South Australia using normalized elliptical Fourier analysis of body shape. Significant intraspecific variation was observed among whole body shapes. This was overcome by limiting subsequent analyses to the anterior snout region, where significant differences in shape were detected among the species examined and provided a high degree of classification success for the skates of GSV. More generally, this approach has the potential to provide a cost- and time-efficient means of discrimination among species of skates. Further research is required to investigate the potentially confounding effects of sexual dimorphism and ontogenetic variation in growth to improve the efficacy of the body shape analysis of the skates and batoids species in general. In addition, this approach requires considerable development to facilitate implementation in a fishery setting.

Phylogenetic analysis and reconfiguration of genera in the cestode order Diphyllidea

The generic boundaries of the Diphyllidea are reassessed based on parsimony and likelihood phylogenetic analyses of 28S rDNA (ribonucleic acid large subunit), 18S rDNA (ribonucleic acid small subunit), and COI (cytochrome oxidase subunit I) sequence data for 31 species representing morphological variation across the order. Trees resulting from these analyses yielded a number of well-supported clades that are congruent with unique morphological features mandating generic revision of the order and erection of at least two new genera. Species originally assigned to Echinobothriumvan Beneden, 1849 but bearing a corona of spines on the region of the scolex anterior to the bothria and posterior to the apical organ armature are transferred to Coronocestus n. gen.; members of this genus typically parasitize triakid sharks, although one report from a hemiscylliid shark exists. Species with lateral hooklets arranged in continuous bands, rather than in two distinct clusters, are transferred to Halysioncum n. gen.; all species parasitize batoids, mostly myliobatids and rhinopterids, but a few records also exist from arhynchobatids, rhinobatids, platyrhinids and urotrygonids. Our analyses support transfer of the five species originally assigned to MacrobothridiumKhalil and Abdul-Salam, 1989 owing to their lack of cephalic peduncle spines to Echinobothrium. As a consequence, Echinobothrium sensu stricto includes species both with and without spines on the cephalic peduncle, but all members of the genus possess lateral hooklets arranged in clusters on either side of the dorsal and ventral apical hooks. With respect to diphyllideans parasitizing catsharks, AhamulinaMarques, Jensen and Caira, 2012 is unique in possessing apical hooks but lacking lateral hooklets and DitrachybothridiumRees, 1959 is unique in entirely lacking scolex armature. By far the majority of species of Echinobothrium sensu stricto parasitize skates of the family Rajidae, guitarfish of the family Rhinobatidae, and stingrays of the dasyatid genera Taeniura Müller and Henle, Dasyatis Rafinesque, and Himantura Müller and Henle, although a single species each has been reported from Anacanthobatidae, Rhynchobatidae, Platyrhinidae and Myliobatidae. It now seems clear that while by far the majority of diphyllideans parasitize batoids, the diphyllideans parasitizing sharks, and catsharks in particular, remain problematic. Additional collections from these carcharhiniform hosts are likely to be particularly illuminating.

Fishery Management Complexes: An Impediment or Aid to Sustainable Harvest? A Discussion Based on the Northeast Skate Complex

AbstractThere are seven species of skates (family Rajidae) found along the East Coast of the USA. All seven species are currently managed by the New England Fisheries Management Council as a single management complex extending from the Gulf of Maine to Cape Hatteras. The objective of the management plan is to ensure the long‐term sustainability of fishing for each species via a trip limit approach. Two species are harvested in two distinct commercial fisheries. Northeast Fisheries Science Center trawl survey data and published literature were examined to investigate differences between the individual species in the skate complex. Each species exhibited a unique thermal and geographic range in addition to vital life history traits (e.g., age at maturity, longevity, and maximum size). Thorny Skate Amblyraja radiata and Smooth Skate Malacoraja senta have narrow thermal ranges and maintain a more northern distribution. Barndoor Skate Dipturus laevis have a moderate thermal habitat. Little Skate Leucoraja erinacea and Winter Skate L. ocellata have broad thermal ranges and are distributed throughout the management area. Limited inferences can be made about the thermal preferences of Clearnose Skate Raja eglanteria without data from south of Cape Hatteras, but they appear to have a broad thermal range within the management area. Rosette Skate L. garmani have a narrow thermal range and tend to be found in the deep offshore mid‐Atlantic region. The validity of managing multiple distinct species in a complex is questioned. This example shows that a mixed‐stock management strategy may be inadequate to meet the sustainability needs of each species and the associated fisheries. A management strategy focused on individual species may lead to a more efficient harvest of targeted species while allowing for the rebuilding of overfished species.Received November 8, 2011; accepted December 27, 2012

An empirical model of seasonal depth-dependent fish assemblage structure to predict the species composition of mixed catches

Reliable catch statistics are essential for assessing fishing impacts on individual stocks. However, fisheries that capture a mixture of stocks or species for which catch statistics are not disaggregated pose a challenge. Nonetheless, catch composition can be inferred given information on fishing date and location and a prevalent role of season and habitat in structuring fish assemblage composition. Here, a harmonic regression model for multinomial data, intended to predict the species composition of catches based on season and depth, is developed using bottom-trawl survey data. Model development was motivated by the need to quantify catches of individual skate (Rajidae) species in fisheries for which landing and discard data are only reliable at the family level. The model was validated by applying it to flatfishes (Pleuronectidae), whose catches are generally reliably and consistently disaggregated by species. The predicted species composition of flatfish matched the composition observed in fishery catches well. The present approach should be applicable to other well-surveyed ecosystems where assemblage composition is structured by one or more key environmental variables of known spatial distribution.