Field guide to requiem sharks (Elasmobranchiomorphi: Carcharhinidae) of the Western North Atlantic

Identification problems are common for many sharks due to a general lack of meristic characteristics that are typically useful for separating species. Other than number of vertebrae and number and shape of teeth, identifications are frequently based on external features that are often shared among species. Identification problems in the field are most prevalent when live specimens are captured and releasing them with a minimum of stress is a priority (e.g., shark tagging programs). Identifications must be accurate and conducted quickly but this can be challenging, especially if specimens are very active or too large to be landed without physical damage. This field guide was designed primarily for use during field studies and presents a simplified method for identifying the 21 species of western North Atlantic Ocean sharks belonging to the family Carcharhinidae (carcharhinids). To assist with identifications a dichotomous key to Carcharhinidae was developed, and for the more problematic Carcharhinus species (12 species), separation sheets based on important distinguishing features were constructed. Descriptive text and illustrations provided in the species accounts were developed from field observations, photographs, and published references. (PDF file contains 36 pages.)

Proceedings of the International Workshop on age determination of oceanic pelagic fishes: Tunas, billfishes, and sharks, Miami, Florida, February 15-18,1982

Accurate and precise estimates of age and growth rates are essential parameters in understanding the population dynamics of fishes. Some of the more sophisticated stock assessment models, such as virtual population analysis, require age and growth information to partition catch data by age. Stock assessment efforts by regulatory agencies are usually directed at specific fisheries which are being heavily exploited and are suspected of being overfished. Interest in stock assessment of some of the oceanic pelagic fishes (tunas, billfishes, and sharks) has developed only over the last decade, during which exploitation has increased steadily in response to increases in worldwide demand for these resources. Traditionally, estimating the age of fishes has been done by enumerating growth bands on skeletal hardparts, through length frequency analysis, tag and recapture studies, and raising fish in enclosures. However, problems related to determining the age of some of the oceanic pelagic fishes are unique compared with other species. For example, sampling is difficult for these large, highly mobile fishes because of their size, extensive distributions throughout the world's oceans, and for some, such as the marlins, infrequent catches. In addition, movements of oceanic pelagic fishes often transect temperate as well as tropical oceans, making interpretation of growth bands on skeletal hardparts more difficult than with more sedentary temperate species. Many oceanic pelagics are also long-lived, attaining ages in excess of 30 yr, and more often than not, their life cycles do not lend themselves easily to artificial propagation and culture. These factors contribute to the difficulty of determining ages and are generally characteristic of this group-the tunas, billfishes, and sharks. Accordingly, the rapidly growing international concern in managing oceanic pelagic fishes, as well as unique difficulties in ageing these species, prompted us to hold this workshop. Our two major objectives for this workshop are to: I) Encourage the interchange of ideas on this subject, and 2) establish the "state of the art." A total of 65 scientists from 10 states in the continental United States and Hawaii, three provinces in Canada, France, Republic of Senegal, Spain, Mexico, Ivory Coast, and New South Wales (Australia) attended the workshop held at the Southeast Fisheries Center, Miami, Fla., 15-18 February 1982. Our first objective, encouraging the interchange of ideas, is well illustrated in the summaries of the Round Table Discussions and in the Glossary, which defines terms used in this volume. The majority of the workshop participants agreed that the lack of validation of age estimates and the means to accomplish the same are serious problems preventing advancements in assessing the age and growth of fishes, particularly oceanic pelagics. The alternatives relating to the validation problem were exhaustively reviewed during the Round Table Discussions and are a major highlight of this workshop. How well we accomplished our second objective, to establish the "state of the art" on age determination of oceanic pelagic fishes, will probably best be judged on the basis of these proceedings and whether future research efforts are directed at the problem areas we have identified. In order to produce high-quality papers, workshop participants served as referees for the manuscripts published in this volume. Several papers given orally at the workshop, and included in these proceedings, were summarized from full-length manuscripts, which have been submitted to or published in other scientific outlets-these papers are designated as SUMMARY PAPERS. In addition, the SUMMARY PAPER designation was also assigned to workshop papers that represented very preliminary or initial stages of research, cursory progress reports, papers that were data shy, or provide only brief reviews on general topics. Bilingual abstracts were included for all papers that required translation. We gratefully acknowledge the support of everyone involved in this workshop. Funding was provided by the Southeast Fisheries Center, and Jack C. Javech did the scientific illustrations appearing on the cover, between major sections, and in the Glossary. (PDF file contains 228 pages.)

Bycatches of sharks in the tuna purse-seine fishery of the eastern Pacific Ocean reported by observers of the Inter-American Tropical Tuna Commission, 1993-2004

Information on bycatches of sharks collected by observers of the Inter-American Tropical Tuna Commission (IATTC) between 1993 and 2004 is presented in this data report. This report contains two sections. The first section summarizes information used by the staff of the IATTC to review and revise IATTC observers’ at-sea species identifications of Carcharhinus falciformis, C. limbatus, and C. longimanus. The revisions were based on 1) data collected on species-specific diagnostic characteristics as part of a special sampling program conducted between March 2000, and March 2001 and 2) a review of observers’ archival field notes for the 1993-2004 period. The second section summarizes the shark bycatches reported by IATTC observers between 1993 and 2004, incorporating the revisions of observers’ at-sea identifications. The IATTC-observed shark bycatch data are summarized as tables with annual tallies of observed bycatches and maps of the spatial distributions of the average bycatches per set and size compositions of the bycatches.

Gillnet selectivity for juvenile blacktip sharks (Carcharhinus limbatus)

Gillnet mesh selectivity parameters were estimated for juvenile blacktip sharks (Carcharhinus limbatus) by using length data from an experimental fishery-independent gillnet survey in the northeastern Gulf of Mexico. Length data for 1720 blacktip sharks were collected over 17 years (1994–2010) with seven mesh sizes ranging from 7.6 to 20.3 cm. Four selectivity models, a normal model assuming fixed spread, a normal model assuming that spread is proportional to mesh size, a lognormal model, and a gamma model were fitted to the data by using the SELECT (share each length’s catch total) method. Each model was run twice under separate assumptions of 1) equal fishing intensity; and 2) fishing intensity proportional to mesh size. The normal, fixed-spread selectivity curve where fishing intensity is assumed to be proportional to mesh size provided the best fit to the data according to model deviance estimates and was chosen as the best model. Results indicate that juvenile blacktip sharks are susceptible as bycatch in some commercial gillnet fisheries.

Postrelease survival, vertical and horizontal movements, and thermal habitats of five species of pelagic sharks in the central Pacific Ocean

From 2001 to 2006, 71 pop-up satellite archival tags (PSATs) were deployed on five species of pelagic shark (blue shark [Prionace glauca]; shortfin mako [Isurus oxyrinchus]; silky shark [Carcharhinus falciformis]; oceanic whitetip shark [C. longimanus]; and bigeye thresher [Alopias superciliosus]) in the central Pacific Ocean to determine species-specific movement patterns and survival rates after release from longline fishing gear. Only a single postrelease mortality could be unequivocally documented: a male blue shark which succumbed seven days after release. Meta-analysis of published reports and the current study (n=78 reporting PSATs) indicated that the summary effect of postrelease mortality for blue sharks was 15% (95% CI, 8.5–25.1%) and suggested that catch-and-release in longline fisheries can be a viable management tool to protect parental biomass in shark populations. Pelagic sharks displayed species-specific depth and temperature ranges, although with significant individual temporal and spatial variability in vertical movement patterns, which were also punctuated by stochastic events (e.g., El Niño-Southern Oscillation). Pelagic species can be separated into three broad groups based on daytime temperature preferences by using the unweighted pair-group method with arithmetic averaging clustering on a Kolmogorov-Smirnov Dmax distance matrix: 1) epipelagic species (silky and oceanic whitetip sharks), which spent >95% of their time at temperatures within 2°C of sea surface temperature; 2) mesopelagic-I species (blue sharks and shortfin makos, which spent 95% of their time at temperatures from 9.7° to 26.9°C and from 9.4° to 25.0°C, respectively; and 3) mesopelagic-II species (bigeye threshers), which spent 95% of their time at temperatures from 6.7° to 21.2°C. Distinct thermal niche partitioning based on body size and latitude was also evident within epipelagic species.

The repulsive and feeding-deterrent effects of electropositive metals on juvenile sandbar sharks (Carcharhinus plumbeus)

Reducing shark bycatch and depredation (i.e., damage caused by sharks to gear, bait, and desired fish species) in pelagic longline fisheries targeting tunas and swordfish is a priority. Electropositive metals (i.e., a mixture of the lanthanide elements lanthanum, cerium, neodymium, and praseodymium) have been shown to deter spiny dogfish (Squalus acanthias, primarily a coastal species) from attacking bait, presumably because of interactions with the electroreceptive system of this shark. We undertook to determine the possible effectiveness of electropositive metals for reducing the interactions of pelagic sharks with longline gear, using sandbar sharks (Carcharhinus plumbeus, family Carcharhinidae) as a model species. The presence of electropositive metal deterred feeding in groups of juvenile sandbar sharks and altered the swimming patterns of individuals in the absence of food motivation (these individuals generally avoided approaching electropositive metal closer than ~100 cm). The former effect was relatively short-lived however; primarily (we assume) because competition with other individuals increased feeding motivation. In field trials with bottom longline gear, electropositive metal placed within ~10 cm of the hooks reduced the catch of sandbar sharks by approximately two thirds, compared to the catch on hooks in the proximity of plastic pieces of similar dimensions. Electropositive metals therefore appear to have the potential to reduce shark interactions in pelagic longline fisheries, although the optimal mass, shape, composition, and distance to baited hooks remain to be determined.

Standard and routine metabolic rates of juvenile sandbar sharks (Carcharhinus plumbeus), including the effects of body mass and acute temperature change*

Standard and routine metabolic rates (SMRs and RMRs, respectively) of juvenile sandbar sharks (Carcharhinus plumbeus) were measured over a range of body sizes (n=34) and temperatures normally associated with western Atlantic coastal nursery areas. The mean SMR Q10 (increase in metabolic rate with temperature) was 2.9 ±0.2. Heart rate decreased with increasing body mass but increased with temperature at a Q10 of 1.8−2.2. Self-paired measures of SMR and RMR were obtained for 15 individuals. Routine metabolic rate averaged 1.8 ±0.1 times the SMR and was not correlated with body mass. Assuming the maximum metabolic rate of sandbar sharks is 1.8−2.75 times the SMR (as is observed in other elasmobranch species), sandbar sharks are using between 34% and 100% of their metabolic scope just to sustain their routine continuous activity. This limitation may help to explain their slow individual and population growth rates, as well as the slow recoveries from overfishing of many shark stocks worl

Estimating consumption rates of juvenile sandbar sharks (Carcharhinus plumbeus) in Chesapeake Bay, Virginia, using a bioenergetics model*

Using a bioenergetics model, we estimated daily ration and seasonal prey consumption rates for six age classes of juvenile sandbar sharks (Carcharhinus plumbeus) in the lower Chesapeake Bay summer nursery area. The model, incorporating habitat and species-specific data on growth rates, metabolic rate, diet composition, water temperature (range 16.8−27.9°C), and population structure, predicted mean daily rations between 2.17 ±0.03 (age-0) and 1.30 ±0.02 (age-5) % body mass/day. These daily rations are higher than earlier predictions for sandbar sharks but are comparable to those for ecologically similar shark species. The total nursery population of sandbar sharks was predicted to consume ~124,000 kg of prey during their 4.5 month stay in the Chesapeake Bay nursery. The predicted consumption rates support the conclusion that juvenile sandbar sharks exert a lesser top-down effect on the Chesapeake Bay ecosystem than do teleost piscivores and hu

Growth and maturity of salmon sharks (Lamna ditropis) in the eastern and western North Pacific, and comments on back-calculation methods

Age and growth estimates for salmon sharks (Lamna ditropis) in the eastern North Pacific were derived from 182 vertebral centra collected from sharks ranging in length from 62.2 to 213.4 cm pre-caudal length (PCL) and compared to previously published age and growth data for salmon sharks in the western North Pacific. Eastern North Pacific female and male salmon sharks were aged up to 20 and 17 years, respectively. Relative marginal increment (RMI) analysis showed that postnatal rings form annually between January and March. Von Bertalanffy growth parameters derived from vertebral length-at-age data are L∞ =207.4 cm PCL, k=0.17/yr, and t0=−2.3 years for females (n=166), and L∞ =182.8 cm PCL, k=0.23/yr , and t0=−1.9 years for males (n=16). Age at maturity was estimated to range from six to nine years for females (median pre-caudal length of 164.7 cm PCL) and from three to five years old for males (median precaudal length of 124.0 cm PCL). Weight-length relationships for females and males in the eastern North Pacific are W=8.2 × 10_05 × L2.759 –06 × L3.383 (r2 =0.99) and W=3.2 × 10 (r2 =0.99), respectively. Our results show that female and male salmon sharks in the eastern North Pacific possess a faster growth rate, reach sexual maturity earlier, and attain greater weight-at-length than their same-sex counterparts living in the western North Pacific.

Exploring intraspecific life history patterns in sharks

Marine ecosystems compose the major source (85%) of world fisheries production (Garcia and Newton, 1997). Although only a few fish species tend to dominate fishery catches (Jennings et al., 2001), a large diversity of fishes representing varied taxonomic levels, ecological guilds, and life histories is commonly taken. Recently, 66% of global marine resources were determined to be either fully, heavily, or over-exploited (Botsford et al., 1997). Considering the current state of many fisheries, the large diversity of species taken globally, and the general lack of resources to adequately assess many stocks, it has become important to develop shortcuts that may provide methods fisheries scientists can use to determine which stocks are in danger of overexploitation and which recovery plans are appropriate when biological data are limited (Stobutzki et al., 2001).

Incidental catch and estimated discards of pelagic sharks from the swordfish and tuna fisheries in the Mediterranean Sea

Large pelagic sharks are caught incidentally in the swordfish and tuna fisheries of the Mediterranean Sea. In our study, twelve shark species were documented as bycatch over three years from 1998 to 2000. Blue shark (Prionace glauca) was the predominant species in all gears and areas examined. Shortfin mako (Isurus oxyrinchus), common thresher shark (Alopias vulpinus), and tope shark (Galeorhinus galeus) were the next most abundant shark species—found in more than half of the areas sampled. Catch composition varied both in the areas and gears investigated. Sharks represented 34.3% in weight of total catches sampled in the Alboran Sea and 0.9% in the Straits of Sicily. Higher shark catches were observed in the swordfish longline fishery, where a nominal CPUE value reached 3.8 sharks/1000 hooks in the Alboran Sea. Size distribution by fishing gear varied significantly. Albacore longline catches consisted mainly of juveniles, whereas subadult and adult specimens were more frequent in the swordfish longline and driftnet fishery. The percentage of sharks brought onboard alive was exceptionally high; only 5.1% of the specimens died. Few discards (seven blue shark) were recorded in the Greek longline fleet during onboard sampling in the Aegean Se

Length at maturity in three pelagic sharks (Lamna nasus, Isurus oxyrinchus, and Prionace glauca) from New Zealand

Reproductive data collected from porbeagle, shortfin mako, and blue sharks caught around New Zealand were used to estimate the median length at maturity. Data on clasper development, presence or absence of spermatophores or spermatozeugmata, uterus width, and pregnancy were collected by observers aboard tuna longline vessels. Direct maturity estimates were made for smaller numbers of sharks sampled at recreational fishing competitions. Some data sets were sparse, particularly over the vital maturation length range, but the availability of multiple indicators of maturity made it possible to develop estimates for both sexes of all three species. Porbeagle shark males matured at 140–150 cm fork length and females at about 170–180 cm. New Zealand porbeagles therefore mature at shorter lengths than they do in the North Atlantic Ocean. Shortfin mako males matured at 180–185 cm and females at 275 –285 cm. Blue shark males matured at about 190 –195 cm and females at 170–190 cm; however these estimates were hampered by small sample sizes, difficulty obtaining representative samples from a population segregated by sex and maturity stage, and maturation that occurred over a wide length range. It is not yet clear whether regional differences in median maturity exist for shortfin mako and

Mitochondrial gene sequences useful for species identification of western North Atlantic Ocean sharks

Molecular-based approaches for shark species identification have been driven largely by issues specific to the fishery. In an effort to establish a more comprehensive identification data set, we investigated DNA sequence variation of a 1.4-kb region from the mitochondrial genome covering partial sequences from the 12S rDNA, 16S rDNA, and the complete valine tRNA from 35 shark species from the Atlantic fishery. Generally, within-species variability was low in relation to interspecific divergence because species haloptypes formed monophyletic groups. Phylogenetic analyses resolved ordinal relationships among Carcharhiniformes and Lamniformes, and revealed support for the families Sphyrnidae and Triakidae (within Carcharhiniformes) and Lamnidae and Alopidae (within Lamniformes). The combination of limited intraspecific variability and sufficient between-species divergence indicates that this locus is suitable for species identification.

Northerly Distribution of White Sharks, Carcharodon carcharias, in the Eastern Pacific and Relation to ENSO Events

Twenty-nine verified records of white sharks, Carcharodon carcharias, from British Columbia and Alaska waters (1961–2004) are presented. Record locations ranged from lat. 48°48ʹN to lat. 60°17ʹN, including the northernmost occurrence of a white shark and the first report of this species from the central Bering Sea. White sharks recorded from the study area were generally large, with 95% falling between 3.8 and 5.4 m in length. Mature white sharks of both sexes occur in British Columbia and Alaska waters, although they do not necessarily reproduce there. White sharks actively feed in the study area; their diet is similar to that reported for this species from Washington and northern California waters. Sea surface temperature (SST) concurrent with white shark records from the study area ranged from 16°C to between 6.4°C and 5.0°C, extending the lower extreme of the range of SST from which this species has been previously reported. White shark strandings are rarely reported, yet 16 (55%) of the records in this study are of beached animals; strandings generally occurred later in the year and at lower latitudes than nonstrandings. No significant correlation was found between white shark records in the study area and El Niño events and no records occurred during La Niña events. The data presented here indicate that white sharks are more abundant in the cold waters of British Columbia and Alaska than previous records suggest.