All their eggs in one basket: a rocky reef nursery for the longnose skate (Raja rhina Jordan & Gilbert, 1880) in the southern California Bight

Skates (family Rajidae) are oviparous and lay tough, thick-walled eggs. At least some skate species lay their eggs in spatially restricted nursery grounds where embryos develop and hatch (Hitz, 1964; Hoff, 2007). After hatching, neonates may quickly leave the nursery grounds (Hoff, 2007). Egg densities in these small areas may be quite high. As an example, in the eastern Bering Sea, a site <2 km2 harbored eggs of Alaska skate (Bathyraja parmifera) exceeding 500,000/km2. All skate nursery grounds have been identified over soft sea floors (Lucifora and García, 2004; Hoff, 2007).

NOVEL TACHYKININS FROM THE BRAIN OF THE SEA LAMPREY, PETROMYZON-MARINUS, AND THE SKATE, RAJA-RHINA

Using radioimmunoassay for mammalian tachykinins, peptides with substance P-like immunoreactivity and neurokinin A-like immunoreactivity were identified in an extract of the brain of the longnose skate, Raja rhina (elasmobranch) but only a peptide with neurokinin A-like immunoreactivity was identified in the brain of the sea lamprey, Petromyzon marinus (agnathan). The primary structure of the skate peptide with substance P-like immunoreactivity (Ala-Lys-His-Asp-Lys-Phe-Tyr-Gly-Leu-Met-NH2) shows one amino acid substitution (Phe(3) --> His) compared with scyliorhinin I, previously isolated from dogfish brain and gut. The skate neurokinin A-related peptide (His-Lys-Leu-Gly-Ser-Phe-Val-Gly-Leu-Met-NH2) shows tow substitutions (Thr(3) --> Leu and Asp(4) --> Gly) compared with mammalian neurokinin A. Although the COOH-terminus of the lamprey tackhykinin (Arg-Lys-Pro-His-Pro-Lys-Gly-phe-Val-Gly-Leu-Met-NH2) resembles neurokinin A, the presence of the strongly conserved Lys/Arg-Pro-Xaa-Pro motif at the NH2-terminus of the peptide indicates greater structural similarity with substance P. The additional arginine residue at position 1 in the peptide suggests that the lamprey is utilizing a site of postranslational processing in the tachykinin precursor that is different from the equivalent site in mammalian and other lower vertebrate preprotachykinin(s).

Resolving taxonomic uncertainty in vulnerable elasmobranchs : are the Madeira skate (Raja maderensis) and the thornback ray (Raja clavata) distinct species?

We are grateful to all those who helped with sample collection. This includes the MEDITS survey programme (IEO Mallorca) for Mediterranean samples. Portugal mainland samples were collected under the EU Data Collection Framework (DCF, PNAB). Azores specimens from the Department of Oceanography and Fisheries (DOP) of the University of the Azores (UAc) were collected under the project DEMERSAIS “Monitorização das espécies demersais dos Açores” financed by the Azorean government, and the project DEECON “Unravelling population connectivity for sustainable fisheries in the Deep Sea” project approved by the European Science Foundation (ESF) under the EUROCORES programme (proposal No 06-EuroDEEP-FP-008 & SFRH-EuroDEEP/0002/2007). This study was funded by the UK Natural Environment Research Council (NERC) Oceans 2025 Strategic Research Programme Theme 6 (Science for Sustainable Marine Resources). REB was supported by the Fisheries Society of the British Isles (FSBI), BSP was funded by the Fundação para a Ciência e a Tecnologia, SFRH/BPD/72351/2010 and JL was supported by The Alasdair Downes Marine Conservation Fund.

Resolving taxonomic uncertainty in vulnerable elasmobranchs: are the Madeira skate (Raja maderensis) and the thornback ray (Raja clavata) distinct species?

Skates and rays constitute the most speciose group of chondrichthyan fishes, yet are characterised by remarkable levels of morphological and ecological conservatism. They can be challenging to identify, which makes monitoring species compositions for fisheries management purposes problematic. Owing to their slow growth and low fecundity, skates are vulnerable to exploitation and species exhibiting endemism or limited ranges are considered to be the most at risk. The Madeira skate Raja maderensis is endemic and classified as ‘Data Deficient’ by the IUCN, yet its taxonomic distinctiveness from the morphologically similar and more wide-ranging thornback ray Raja clavata is unresolved. This study evaluated the sequence divergence of both the variable control region and cytochrome oxidase I ‘DNA barcode’ gene of the mitochondrial genome to elucidate the genetic differentiation of specimens identified as R. maderensis and R. clavata collected across much of their geographic ranges. Genetic evidence was insufficient to support the different species designations. However regardless of putative species identification, individuals occupying waters around the Azores and North African Seamounts represent an evolutionarily significant unit worthy of special consideration for conservation management.

Resolving taxonomic uncertainty in vulnerable elasmobranchs: are the Madeira skate (Raja maderensis) and the thornback ray (Raja clavata) distinct species?

Skates and rays constitute the most speciose group of chondrichthyan fishes, yet are characterised by remarkable levels of morphological and ecological conservatism. They can be challenging to identify, which makes monitoring species compositions for fisheries management purposes problematic. Owing to their slow growth and low fecundity, skates are vulnerable to exploitation and species exhibiting endemism or limited ranges are considered to be the most at risk. The Madeira skate Raja maderensis is endemic and classified as ‘Data Deficient’ by the IUCN, yet its taxonomic distinctiveness from the morphologically similar and more wide-ranging thornback ray Raja clavata is unresolved. This study evaluated the sequence divergence of both the variable control region and cytochrome oxidase I ‘DNA barcode’ gene of the mitochondrial genome to elucidate the genetic differentiation of specimens identified as R. maderensis and R. clavata collected across much of their geographic ranges. Genetic evidence was insufficient to support the different species designations. However regardless of putative species identification, individuals occupying waters around the Azores and North African Seamounts represent an evolutionarily significant unit worthy of special consideration for conservation management.

Opportunistic feeding of longhorn sculpin (Myoxocephalus octodecemspinosus): Are scallop fishery discards an important food subsidy for scavengers on Georges Bank?

There has been much recent interest in the effects of fishing on habitat and non-target species, as well as in protecting certain areas of the seabed from these effects (e.g. Jennings and Kaiser, 1998; Benaka, 1999; Langton and Auster, 1999; Kaiser and de Groot, 2000). As part of an effort to determine the effectiveness of marine closed areas in promoting recovery of commercial species (e.g. haddock, Melanogrammus aegelfinus; sea scallops, Placopecten magellanicus; yellowtail flounder, Limanda ferruginea; cod, Gadus morhua), nontarget species, and habitat, a multidisciplinary research cruise was conducted by the Northeast Fisheries Science Center (NEFSC), National Marine Fisheries Service. The cruise was conducted in closed area II (CA-II) of the eastern portion of Georges Bank during 19–29 June 2000 (Fig. 1). The area has historically produced high landings of scallops but was closed in 1994 principally for groundfish recovery (Fogarty and Murawski, 1998). The southern portion of the area was reopened to scallop fishing from 15 June to 12 November 1999, and again from 15 June to 15 August 2000. While conducting our planned sampling, we observed scallop viscera (the noncalcareous remains from scallops that have been shucked by commercial fishermen at sea) in the stomachs of several fish species at some of these locations, namely little skate (Raja erinacea), winter skate (R. ocellata), red hake (Urophycis chuss), and longhorn sculpin (Myoxocephalus octodecemspinosus). We examined the stomach contents of a known scavenger, the longhorn sculpin, to evaluate and document the extent of this phenomenon.

Urea-based osmoregulation in the developing embryo of oviparous cartilaginous fish (Callorhinchus milii) : contribution of the extraembryonic yolk sac during the early developmental period

Marine cartilaginous fish retain a high concentration of urea to maintain the plasma slightly hyperosmotic to the surrounding seawater. In adult fish, urea is produced by hepatic and extrahepatic ornithine urea cycles (OUCs). However, little is known about the urea retention mechanism in developing cartilaginous fish embryos. In order to address the question as to the mechanism of urea-based osmoregulation in developing embryos, the present study examined the gene expression profiles of OUC enzymes in oviparous holocephalan elephant fish (Callorhinchus milii) embryos. We found that the yolk sac membrane (YSM) makes an important contribution to the ureosmotic strategy of the early embryonic period. The expression of OUC enzyme genes was detectable in the embryonic body from at least stage 28, and increased markedly during development to hatching, which is most probably due to growth of the liver. During the early developmental period, however, the expression of OUC enzyme genes was not prominent in the embryonic body. Meanwhile, we found that the mRNA expression of OUC enzymes was detected in the extra-embryonic YSM; the mRNA expression of cmcpsIII in the YSM was much higher than that in the embryonic body during stages 28-31. Significant levels of enzyme activity and the existence of mitochondrial-type cmgs1 transcripts in the YSM supported the mRNA findings. We also found that the cmcpsIII transcript is localized in the vascularized inner layer of the YSM. Taken together, our findings demonstrate for the first time that the YSM is involved in urea-based osmoregulation during the early to mid phase of development in oviparous cartilaginous fish.

Morphological and molecular investigations of the holocephalan elephant fish nephron: the existence of a countercurrent-like configuration and two separate diluting segments in the distal tubule

In marine cartilaginous fish, reabsorption of filtered urea by the kidney is essential for retaining a large amount of urea in their body. However, the mechanism for urea reabsorption is poorly understood due to the complexity of the kidney. To address this problem, we focused on elephant fish (Callorhinchus milii) for which a genome database is available, and conducted molecular mapping of membrane transporters along the different segments of the nephron. Basically, the nephron architecture of elephant fish was similar to that described for elasmobranch nephrons, but some unique features were observed. The late distal tubule (LDT), which corresponded to the fourth loop of the nephron, ran straight near the renal corpuscle, while it was convoluted around the tip of the loop. The ascending and descending limbs of the straight portion were closely apposed to each other and were arranged in a countercurrent fashion. The convoluted portion of LDT was tightly packed and enveloped by the larger convolution of the second loop that originated from the same renal corpuscle. In situ hybridization analysis demonstrated that co-localization of Na(+),K(+),2Cl(-) cotransporter 2 and Na(+)/K(+)-ATPase α1 subunit was observed in the early distal tubule and the posterior part of LDT, indicating the existence of two separate diluting segments. The diluting segments most likely facilitate NaCl absorption and thereby water reabsorption to elevate urea concentration in the filtrate, and subsequently contribute to efficient urea reabsorption in the final segment of the nephron, the collecting tubule, where urea transporter-1 was intensely localized.

Molecular evolution of two vertebrate aryl hydrocarbon (dioxin) receptors (AHR1 and AHR2) and the PAS family

The aryl hydrocarbon receptor (AHR) is a ligand-activated transcription factor through which halogenated aromatic hydrocarbons such as 2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD) cause altered gene expression and toxicity. The AHR belongs to the basic helix–loop–helix/Per-ARNT-Sim (bHLH-PAS) family of transcriptional regulatory proteins, whose members play key roles in development, circadian rhythmicity, and environmental homeostasis; however, the normal cellular function of the AHR is not yet known. As part of a phylogenetic approach to understanding the function and evolutionary origin of the AHR, we sequenced the PAS homology domain of AHRs from several species of early vertebrates and performed phylogenetic analyses of these AHR amino acid sequences in relation to mammalian AHRs and 24 other members of the PAS family. AHR sequences were identified in a teleost (the killifish Fundulus heteroclitus), two elasmobranch species (the skate Raja erinacea and the dogfish Mustelus canis), and a jawless fish (the lamprey Petromyzon marinus). Two putative AHR genes, designated AHR1 and AHR2, were found both in Fundulus and Mustelus. Phylogenetic analyses indicate that the AHR2 genes in these two species are orthologous, suggesting that an AHR gene duplication occurred early in vertebrate evolution and that multiple AHR genes may be present in other vertebrates. Database searches and phylogenetic analyses identified four putative PAS proteins in the nematode Caenorhabditis elegans, including possible AHR and ARNT homologs. Phylogenetic analysis of the PAS gene family reveals distinct clades containing both invertebrate and vertebrate PAS family members; the latter include paralogous sequences that we propose have arisen by gene duplication early in vertebrate evolution. Overall, our analyses indicate that the AHR is a phylogenetically ancient protein present in all living vertebrate groups (with a possible invertebrate homolog), thus providing an evolutionary perspective to the study of dioxin toxicity and AHR function.

Urea based osmoregulation and endocrine control in elasmobranch fish with special reference to euryhalinity

Since the landmark contributions of Homer Smith and co-workers in the 1930s there has been a considerable advance in our knowledge regarding the osmoregulatory strategy of elasmobranch fish. Smith recognised that urea was retained in the body fluids as part of the 'osmoregulatory ballast' of elasmobranch fish so that body fluid osmolality is raised to a level that is iso- or slightly hyper-osmotic to that of the surrounding medium. From studies at that time he also postulated that many marine dwelling elasmobranchs were not capable of adaptation to dilute environments. However, more recent investigations have demonstrated that, at least in some species, this may not be the case. Gradual acclimation of marine dwelling elasmobranchs to varying environmental salinities under laboratory conditions has demonstrated that these fish do have the capacity to acclimate to changes in salinity through independent regulation of Na+, Cl- and urea levels. This suggests that many of the presumed stenohaline marine elasmobranchs could in fact be described as partially euryhaline. The contributions of Thomas Thorson in the 1970s demonstrated the osmoregulatory strategy of a fully euryhaline elasmobranch, the bull shark, Carcharhinus leucas, and more recent investigations have examined the mechanisms behind this strategy in the euryhaline elasmobranch, Dasyatis sabina. Both partially euryhaline and fully euryhaline species utilise the same physiological processes to control urea, Na+ and Cl- levels within the body fluids. The role of the gills, kidney, liver, rectal gland and drinking process is discussed in relation to the endocrine control of urea, Na+ and Cl- levels as elasmobranchs acclimate to different environmental salinities. (C) 2003 Elsevier Inc. All rights reserved.

Observer-reported skate bycatch in the commercial groundfish fisheries of Alaska

We analyzed skate catch data collected by observers in the North Pacific Groundfish Observer Program (NPGOP) from 1998 through 2008 to document recent changes in the identification of skates by observers and to examine the species composition of observed skate catch in Alaska’s groundfish fisheries as well as recent trends in skate retention by commercial fishermen. Historically, almost all skate bycatch has been reported by NPGOP observers as “skate unidentified.” However, since 2004 observers have been trained to identify skates to the genus and species level. In 2008 over 95% of all skates were identified at least to the genus level, and over 50% were identified to species. The most common species of skates identified by observers in groundfish fisheries are Bathyraja parmifera (Alaska skate), Raja binoculata (big skate), and Bathyraja aleutica (Aleutian skate). Species composition of reported skate catch generally reflects recent survey-derived biomass estimates, with B. parmifera dominating the catches in the Bering Sea and, to a lesser extent, in the Aleutian Islands region, and species of the genus Raja dominating catches in the Gulf of Alaska. A relatively high percentage of the skate catch on longline vessels is still reported at the family or genus level because of difficulties in the identification of skates not brought onboard the vessel. For the larger skate species, the proportion retained for processing has increased in recent years as the market price for skate product has increased. Although observed skate catch does not give a complete account of skate bycatch in the fisheries of the region, observer data provide critical information for the appropriate management of skate populations in Alaska.

Analysis of permanent magnets as elasmobranch bycatch reduction devices in hook-and-line and longline trials

Previous studies indicate that elasmobranch fishes (sharks, skates and rays) detect the Earth’s geomagnetic field by indirect magnetoreception through electromagnetic induction, using their ampullae of Lorenzini. Applying this concept, we evaluated the capture of elasmobranchs in the presence of permanent magnets in hook-and-line and inshore longline fishing experiments. Hooks with neodymium-iron-boron magnets significantly reduced the capture of elasmobranchs overall in comparison with control and procedural control hooks in the hook-and-line experiment. Catches of Atlantic sharpnose shark (Rhizoprionodon terraenovae) and smooth dogfish (Mustelus canis) were signif icantly reduced with magnetic hook-and-line treatments, whereas catches of spiny dogfish (Squalus acanthias) and clearnose skate (Raja eglanteria) were not. Longline hooks with barium-ferrite magnets significantly reduced total elasmobranch capture when compared with control hooks. In the longline study, capture of blacktip sharks (Carcharhinus limbatus) and southern stingrays (Dasyatis americana) was reduced on magnetic hooks, whereas capture of sandbar shark (Carcharhinus plumbeus) was not affected. Teleosts, such as red drum (Sciaenops ocellatus), Atlantic croaker (Micropogonias undulatus), oyster toadfish (Opsanus tau), black sea bass (Centropristis striata), and the bluefish (Pomatomas saltatrix), showed no hook preference in either hook-and-line or longline studies. These results indicate that permanent magnets, although eliciting species-specific capture trends, warrant further investigation in commercial longline and recreational fisheries, where bycatch mortality is a leading contributor to declines in elasmobranch populations.

Hepatic urea biosynthesis in the euryhaline elasmobranch Carcharhinus leucas

Plasma urea levels and hepatic urea production in the euryhaline bull shark, Carcharhinus leucas, acclimated to freshwater and seawater environments were measured. It was found that plasma urea concentration increased with salinity and that this increase was, in part, the result of a significant increase in hepatic production of urea. This study provides direct evidence that hepatic production of urea plays an important role in the osmoregulatory strategy of C. leucas.

Änderung wissenschaftlicher Gattungsnamen der Echten Rochen (Raja spp.), Familie Rajidae

Based on a comprehensive compilation and evaluation of hitherto published revisions by international rajid systematists, the U.S. authors McEachran and Dunn (1998) published a summarizing revision on phylogenetic interrelationships within the family Rajidae. The family has been subdivided into the two subfamilies Rajinae and Arhynchobatinae comprising 15 and 11 rajid genera, respectively. Within the Rajinae, seven former subgenera of Raja and Gurgesiella have been newly elevated to generic rank, likewise two former subgenera of Raja within the Arhynchobatinae. As a consequence, generic names have changed for quite a number of mainly former Raja species not only in European and North Atlantic waters. Table 1 presents for comparison a selection of European skate species on the shelf with the former and new generic assignment, as well as their German names. As Fishery Conventions and Societies, as well as regional (e.g., ICES) and international organizations (e.g., FAO) are increasingly using the new scientific name combinations for fishery statistics and in relevant publications, all involved in fisheries should be aware of these nomenclatorial alterations.