Global population structure of the spiny dogfish Squalus acanthias,a temperate shark with an antitropical distribution

The spiny dogfish (Squalus acanthias) is a temperate, coastal squaloid shark with an antitropical distribution in the Atlantic and Pacific oceans. The global population structure of this species is poorly understood, although individuals are known to undergo extensive migrations within coastal waters and across ocean basins. In this study, an analysis of the global population structure of the spiny dogfish was conducted using eight polymorphic nuclear microsatellite markers and a 566‐bp fragment of the mitochondrial ND2 gene region. A low level of genetic divergence was found among collections from the Atlantic and South Pacific basins, whereas a high level of genetic divergence was found among Pacific Ocean collections. Two genetically distinct groups were recovered by both marker classes: one exclusive to North Pacific collections, and one including collections from the South Pacific and Atlantic locations. The strong genetic break across the equatorial Pacific coincides with major regional differences in the life‐history characters of spiny dogfish, suggesting that spiny dogfish in areas on either side of the Pacific equator have been evolving independently for a considerable time. Phylogeographic analyses indicate that spiny dogfish populations had a Pacific origin, and that the North Atlantic was colonized as a result of a recent range expansion from the South American coast. Finally, the available data strongly argue for the taxonomic separation of the North Pacific spiny dogfish from S. acanthias and a re‐evaluation of the specific status of S. acanthias is warranted.     

Lipid correction model of carbon stable isotopes for a cosmopolitan predator, spiny dogfish Squalus acanthias

Three lipid correction models were evaluated for liver and white dorsal muscle from Squalus acanthias. For muscle, all three models performed well, based on the Akaike Information Criterion value corrected for small sample sizes (AIC(c) ), and predicted similar lipid corrections to δ(13) C that were up to 2.8 ‰ higher than those predicted using previously published models based on multispecies data. For liver, which possessed higher bulk C:N values compared to that of white muscle, all three models performed poorly and lipid-corrected δ(13) C values were best approximated by simply adding 5.74 ‰ to bulk δ(13) C values.     

Organ-related distribution of phospholemman in the spiny dogfish Squalus acanthias

The distribution of phospholemman among nine different organs of the spiny dogfish (Squalus acanthias) has been determined on the basis of Western blotting of microsomal material. Only rectal gland (100%), brain (43%), heart (18%), and kidney (19%) (abundancies as percent of the concentration in rectal gland) contained the protein, but not gill and colon. The relative abundance in the brain makes this organ a preferential test system for phospholemman in fishes that lack a rectal gland like teleosts.     

Reproductive Biology of the Smooth Dogfish, Mustelus canis, in the Northwest Atlantic Ocean

Shark populations tend to be highly vulnerable to overexploitation due to low fecundity and slow growth rates. Recent fishing pressure on the smooth dogfish in the northwest Atlantic has created a necessity for more information about their basic biology. Length and age at maturity, fecundity, and reproductive season were estimated. Total length at 50% maturity was 102cm for females, and 86cm for males. The majority of females were mature at age 4 or 5, and all males were mature at 2 or 3 years of age. Females had an 11–12 month gestation with parturition occurring in May, ovulation occurring between May and July likely in late May and early June, and mating occurring between May and September. Fecundity ranged between 3 and 18 pups per litter, and was positively related to length and age, with a mean of 9.53 pups per litter. Sperm was found in the terminal zone of the oviducal gland of females throughout the year.     

Sertoli cell cytoplasts in the semen of the spiny dogfish Squalus acanthias

The fine structure of Squalus acanthias (Elasmobranchii) semen was investigated to determine the cellular component responsible for the steroidogenic activity previously demonstrated in the seminal plasma of this species. Semen was found to consist of bundles of spermatozoa, many of which were encased in a sleeve of cytoplasm restricted to the tail region; large, dense bodies lacking a limiting membrane and numerous anuclear cytoplasmic remnants containing lipid droplets, mitochondria, areas of agranular reticulum, and possibly unreleased spermatozoa. These remnants, which we have termed cytoplasts, morphologically resemble in appearance Sertoli cells of S. acanthias at the time of spermiation. This structural similarity, plus the fact that many elasmobranch species slough the apical regions of Sertoli cells during the release of spermatozoa, indicates that the cytoplasts present in the semen of S. acanthias originate from Sertoli cells. Furthermore, the occurrence in these cytoplasts of organelles normally associated with steroid synthesis strongly suggests these structures are the source of steroidogenic enzymes in the semen of S. acanthias. The steroidal contribution to the semen by Sertoli cell cytoplasts may be necessary for either maturation or maintenance of spermatozoa in the excurrent reproductive ducts of S. acanthias.     

Vulnerability to longline fisheries of three hammerhead shark Sphyrna species in the south‐western and equatorial Atlantic Ocean

Catch and effort data from 29 418 longline sets from Brazilian tuna longline vessels operating in the south‐western and equatorial Atlantic Ocean between 2004 and 2011 were analysed to investigate the distribution, catch rate and size of three species of hammerhead sharks (Sphyrna lewini, Sphyrna mokarran and Sphyrna zygaena). During that period, 6172 hammerhead sharks were caught. Among the elasmobranchs, the highest percentage of hammerhead sharks were caught in 2007, when they accounted for 3·90% of the group, while the lowest value of 0·40% was recorded in 2010. In general, the spatial distribution of the mean catch per unit effort (CPUE) by years and quarters showed a trend of higher catches near the equatorial region and in southern Brazil. The nominal mean CPUE was 0·12 Sphyrna spp. 1000⁻¹ hooks, with the highest value being recorded in 2007 (0·30 Sphyrna spp. 1000⁻¹ hooks). The standardized yearly CPUE estimated by a generalized linear model assuming a zero inflated negative binomial (ZINB) distribution were not much different from nominal values. Of the 205 sexed specimens, 117 were females and 88 were males, resulting in a sex ratio with a predominance of females (1·30:1·00), although not statistically significant. The total length of females ranged from 1200 to 2800 mm and of males from 1100 to 3100 mm. Juvenile hammerhead sharks represented 82 and 54% of the sexed female and male specimens, respectively.     

Characterization of large basking shark Cetorhinus maximus aggregations in the western North Atlantic Ocean

Cetorhinus maximus aggregations recorded during extensive aerial survey efforts off the north‐eastern United States between 1980 and 2013 included aggregations centring on sightings with group sizes of at least 30 individuals. These aggregations occurred in summer and autumn months and included aggregation sizes of up to 1398 individuals, the largest aggregation ever reported for this species. The aggregations were associated with sea surface temperatures of 13–24° C and chlorophyll‐a concentrations of 0·4–2·6 mg m^?3 and during one aggregation, a high abundance of zooplankton prey was present. Photogrammetric tools allowed for the estimation of total body lengths ranging between 4 and 8 m. Characterization of these events provides new insight into the potential biological function of large aggregations in this species.     

Reproductive biology of the lesser guitarfish Zapteryx brevirostris from the south-western Atlantic Ocean

This study provides the first detailed information on the reproductive biology of the lesser guitarfish Zapteryx brevirostris in the south-western Atlantic Ocean between latitudes 34° and 42° S. A total of 320 males (196-647 mm total length, L(T)) and 336 females (200-640 mm L(T)) were analysed. No significant differences were observed in size at 50% maturity between males (499 mm L(T)) and females (506 mm L(T)). Three seasonal patterns were related to the testes mass, the development of spermatocysts and the presence of spermatozoa in genital ducts. Females showed a 3 year reproductive cycle, with 2 years of ovarian maturation and 1 year of gestation, including a mean annual fecundity of one to two embryos. Synchronicity of both ovulating females and males with sperm in genital ducts indicated that ovulation and hence mating occurs during winter. Birth was assumed to occur during winter, but neither ovulation nor mating is simultaneous with this reproductive event. Despite its low economic value, a decline in abundance in the region, together with a low reproductive efficiency, indicates that the Z. brevirostris population should be monitored closely.     

Comparative analysis of methods for determining bite force in the spiny dogfish Squalus acanthias

Many studies have identified relationships between the forces generated by the cranial musculature during feeding and cranial design. Particularly important to understanding the diversity of cranial form amongst vertebrates is knowledge of the generated magnitudes of bite force because of its use as a measure of ecological performance. In order to determine an accurate morphological proxy for bite force in elasmobranchs, theoretical force generation by the quadratomandibularis muscle of the spiny dogfish Squalus acanthias was modeled using a variety of morphological techniques, and lever-ratio analyses were used to determine resultant bite forces. These measures were compared to in vivo bite force measurements obtained with a pressure transducer during tetanic stimulation experiments of the quadratomandibularis. Although no differences were found between the theoretical and in vivo bite forces measured, modeling analyses indicate that the quadratomandibularis muscle should be divided into its constituent divisions and digital images of the cross-sections of these divisions should be used to estimate cross-sectional area when calculating theoretical force production. From all analyses the maximum bite force measured was 19.57 N. This relatively low magnitude of bite force is discussed with respect to the ecomorphology of the feeding mechanism of S. acanthias to demonstrate the interdependence of morphology, ecology, and behavior in organismal design.     

Characterization and isolation of DNA microsatellite primers in the spiny dogfish (Squalus acanthias)

The North Atlantic spiny dogfish (Squalus acanthias) population has been declining since the 1980s. Proper management of this population is essential as dogfish are prone to rapid collapse based on the unique biology of this species. We characterized eight microsatellite loci for spiny dogfish to use in determining the genetic structure of this species along the Atlantic coast of the USA.     

Gastrin/CCK-like peptides in the spiny dogfish, Squalus acanthias; concentrations and actions in the gut

1. The presence of gastrin/CCK-like immunoreactive material in both muscle and mucosal layers of the whole gut, except the cardiac stomach, in Squalus acanthias has been confirmed by radioimmunoassay. The highest levels were measured in rectum and spiral intestine respectively. 2. Fractionation of the spiral intestine mucosal extract on DEAE 52 columns indicated the presence of multiple forms of gastrin/CCK in the elasmobranch gut. 3. Synthetic mammalian gastrin- and CCK-like peptides, when effective, increased the rhythmic activity and sometimes the basal tonus in smooth muscle preparations from the intestine or rectum. 4. The irregular effects of mammalian peptides on the motility indicated an essential difference from the mammalian counterpart in the structure of the elasmobranch peptide present in the gastrointestinal nerves.     

Cardiovascular control via angiotensin II and circulating catecholamines in the spiny dogfish, Squalus acanthias

The contributions of circulating angiotensin II (Ang II) and catecholamines to cardiovascular control in the spiny dogfish were investigated by monitoring the effects of exogenous and endogenous dogfish [Asn¹, Pro³, Ile⁵]-Ang II (dfAng II) on plasma catecholamine levels and blood pressure regulation. Bolus intravenous injections of dfAng II (30–1200 pmol kg⁻¹) elicited dose-dependent increases in plasma adrenaline and noradrenaline concentrations, caudal artery pressure (P _CA), and systemic vascular resistance (R _S), and a decrease in cardiac output (Q). Similar injections of Ang II in dogfish pre-treated with the α-adrenoceptor antagonist yohimbine (4 mg kg⁻¹) also elicited dose-dependent increases in plasma catecholamine levels yet the cardiovascular effects were abolished. Dogfish treated with yohimbine were hypotensive and had elevated levels of plasma Ang II and catecholamines. Intravenous injection of the smooth muscle relaxant papaverine (10 mg kg⁻¹) elicited a transient decrease in P _CA and R _S, and increases in plasma Ang II and catecholamine levels. In dogfish first treated with lisinopril (10⁻⁴ mol kg⁻¹), an angiotensin converting enzyme inhibitor, papaverine treatment caused a more prolonged and greater decrease in P _CA and R _S, an attenuated increase in plasma catecholamines, and no change in plasma Ang II. By itself, lisinopril treatment had little effect on P _CA, and no effect on R _S, plasma Ang II or catecholamines. In yohimbine-treated dogfish, papaverine treatment elicited marked decreases in P _CA, R _S, and Q, and increases in plasma Ang II and catecholamines. Among the three papaverine treatments, there was a positive linear relationship between plasma Ang II and catecholamine concentrations, and the cardiovascular and hormonal changes were most pronounced in the yohimbine + papaverine treatment. Therefore, under resting normotensive conditions, while Ang II does not appear to be involved in cardiovascular control, catecholamines play an important role. However, during a hypotensive stress elicited by vascular smooth muscle relaxation, Ang II indirectly contributes to cardiovascular control by dose-dependently stimulating catecholamine release. Accepted: 24 February 1999     

Bomb dating and age validation using the spines of spiny dogfish (Squalus acanthias)

Bomb radiocarbon has previously been used to validate the age of large pelagic sharks based on incorporation into vertebrae. However, not all sharks produce interpretable vertebral growth bands. Here we report the first application of bomb radiocarbon as an age validation method based on date-specific incorporation into spine enamel. Our results indicate that the dorsal spines of spiny dogfish, Squalus acanthias, recorded and preserved a bomb radiocarbon pulse in growth bands formed during the 1960s with a timing which was very similar to that of marine carbonates. Using radiocarbon assays of spine growth bands known to have formed in the 1960s and 1970s as a dated marker, we confirm the validity of spine enamel growth band counts as accurate annual age indicators to an age of at least 45 year. Radiocarbon incorporation into northeast Atlantic dogfish spines occurred in similar years as those in the northwest Atlantic and northeast Pacific, although the amount of radiocarbon differed in keeping with the radiocarbon content of the different water masses. Published reports suggesting that Pacific dogfish are longer lived and slower growing than their Atlantic counterparts appear to be correct, and are not due to errors in interpreting the spine growth bands. Radiocarbon assays of fin spine enamel appears to be well suited to the age validation of sharks with fin spines which inhabit the upper 200 m of the ocean.     

Reproductive biology of the greeneye spurdog Squalus chloroculus (Squaliformes,Squalidae)

The reproduction of the greeneye spurdog Squalus chloroculus was studied based on animals caught in the multispecies and multi‐gear southern and eastern scalefish and shark fishery on the upper continental slope off southern Australia. One hundred and ninety‐nine females (502–990 mm, total length, L_T) and 189 males (515–810 mm L_T) were examined. The female reproductive cycle, based on 41 breeding animals, is continuous and triennial, with the pregnancy period estimated to be 31–34 months, seasonal and synchronous with the ovarian cycle; a third of the breeding female population is estimated to give birth between September and December each year. The estimated L_T at which 50% of females are mature is 799 mm (95% c.i. : 794, 804), whereas the L_T at which 50% are maternal is 825 mm (95% c.i. : 817–833), but these estimates are probably biased by the phenomenon of apparent change of L_T at maternity and L_T at maturity following severe length‐selective fishing mortality. Litters ranged from four to 15 embryos with a 1:1 sex ratio, and litter size increased with maternal length. The breeding cycle of males is neither seasonal nor synchronous with the female cycle. The estimated L_T of males where 50% are mature was 629 mm (95% c.i. : 603, 645).     

Reproductive biology of the spotback skate Atlantoraja castelnaui in the south-west Atlantic Ocean

This study provides information on the reproduction of spotback skate Atlantoraja castelnaui. A total of 232 individuals (119 females and 113 males) were obtained from surveys carried out between 2003 and 2006, from the south-west Atlantic Ocean, between 34 and 42° S and <50 m deep; another 514 specimens (241 females and 273 males) were obtained between 2005 and 2007 from commercial fishery operations carried out in the same area and landings in the port of Mar del Plata, Argentina. Males ranged from 185 to 1250 mm total length (L(T) ) and females from 243 to 1368 mm L(T) . Length at maturity was estimated to be 980 mm for males and 1089 mm L(T) for females. Lack of variation of testis mass together with the continuous production of mature spermatocyst and spermatozoa in deferent ducts suggested that males can reproduce throughout the year. Females reproduced year-round with peaks of reproductive activity an integral part of a continuous cycle. This conclusion is corroborated by the seasonal variation of ovaries, oviducal gland and the occurrence of females with eggs in the uterus throughout the year. Results from this study indicate that A. castelnaui is very susceptible to fishery pressure.     

Type IV carbonic anhydrase is present in the gills of spiny dogfish (Squalus acanthias)

Physiological and biochemical studies have provided indirect evidence for a membrane-associated carbonic anhydrase (CA) isoform, similar to mammalian type IV CA, in the gills of dogfish (Squalus acanthias). This CA isoform is linked to the plasma membrane of gill epithelial cells by a glycosylphosphatidylinositol anchor and oriented toward the plasma, such that it can catalyze the dehydration of plasma HCO(3)(-) ions. The present study directly tested the hypothesis that CA IV is present in dogfish gills in a location amenable to catalyzing plasma HCO(3)(-) dehydration. Homology cloning techniques were used to assemble a 1,127 base pair cDNA that coded for a deduced protein of 306 amino acids. Phylogenetic analysis suggested that this protein was a type IV CA. For purposes of comparison, a second cDNA (1,107 base pairs) was cloned from dogfish blood; it encoded a deduced protein of 260 amino acids that was identified as a cytosolic CA through phylogenetic analysis. Using real-time PCR and in situ hybridization, mRNA expression for the dogfish type IV CA was detected in gill tissue and specifically localized to pillar cells and branchial epithelial cells that flanked the pillar cells. Immunohistochemistry using a polyclonal antibody raised against rainbow trout type IV CA revealed a similar pattern of CA IV immunoreactivity and demonstrated a limited degree of colocalization with Na(+)-K(+)-ATPase immunoreactivity. The presence and localization of a type IV CA isoform in the gills of dogfish is consistent with the hypothesis that branchial membrane-bound CA with an extracellular orientation contributes to CO(2) excretion in dogfish by catalyzing the dehydration of plasma HCO(3)(-) ions.     

Age and growth of the blacknose shark, <Emphasis Type="Italic">Carcharhinus acronotus</Emphasis>, in the western North Atlantic Ocean with comments on regional variation in growth rates

We examined the age and growth of the blacknose shark, Carcharhinus acronotus, in the western North Atlantic Ocean by obtaining direct age estimates using vertebral centra. We verified annual deposition of growth increments with marginal increment analysis and validated it by analyzing vertebrae marked with oxytetracycline from a female blacknose shark held in captivity. Von Bertalanffy growth parameters indicated that female blacknose sharks have a lower growth constant (k), a larger theortical maximum size (L), and are longer lived than males. We compared these growth parameters for blacknose sharks in the western North Atlantic Ocean to growth parameters for blacknose sharks collected in the eastern Gulf of Mexico to test for differences between regions. Females in the western North Atlantic Ocean have a significantly lower L, lower k, and a higher theoretical longevity than females in the Gulf of Mexico. Males in the western North Atlantic Ocean have a higher L<>, lower k, and higher theoretical longevity than males in the Gulf of Mexico. The significant differences between these life history parameters for blacknose sharks suggest that, when possible, future management initiatives concerning blacknose sharks should consider managing the populations in the western North Atlantic and the Gulf of Mexico as separate stocks.