Additions to a revision of the shark genus Carcharhinus: Synonymy of Aprionodon and Hypoprion, and description of a new species of Carcharhinus (Carcharhinidae)

Features of the valid nominal species of Aprionodon Gill (isodon Valenciennes) and Hypoprion Muller and Henle (hemiodon Valenciennes, macloti Muller and Henle, and signatus Poey), plus those of a previously unrecognized species here described as Carcharhinus leiodon n.sp., are examined and compared with those of Carcharhinus Blainville. Features studied include morphometrics, vertebral numbers and other vertebral characteristics, tooth numbers, color pattern, and some other aspects of external morphology. It is concluded that on these features C. leiodon n.sp. is entirely encompassed within the parameters of Carcharhinus, and that, although A. isodon, H. hemiodon, H. macloti, and H. signatus each extend the range of diversity of Carcharhinus in one or more features, A. isodon is not uniquely different from Carcharhinus, and there is no common pattern of difference between the three species of Hypoprion and Carcharhinus. Accordingly, and because the nature of the teeth of Aprionodon and Hypoprion has been found insufficient to warrant generic distinction from Carcharhinus, the genera Aprionodon and Hypoprion are synonymised with Carcharhinus. A diagnosis and description are given for each of the above species. The descriptions include measurements, counts, and line illustrations that show the whole shark in lateral view, underside of head, nostril, and teeth. The geographic distribution is summarized, as are also the meager biological data available on number of embryos, size at birth, size at sexual maturity, and maximum size. (PDF file contains 32 pages.)

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.

Bomb radiocarbon and tag-recapture dating of sandbar shark (Carcharhinus plumbeus)

The sandbar shark (Carcharhinus plumbeus) was the cornerstone species of western North Atlantic and Gulf of Mexico large coastal shark fisheries until 2008 when they were allocated to a research-only fishery. Despite decades of fishing on this species, important life history parameters, such as age and growth, have not been well known. Some validated age and growth information exists for sandbar shark, but more comprehensive life history information is needed. The complementary application of bomb radiocarbon and tag-recapture dating was used in this study to determine valid age-estimation criteria and longevity estimates for this species. These two methods indicated that current age interpretations based on counts of growth bands in vertebrae are accurate to 10 or 12 years. Beyond these years, we could not determine with certainty when such an underestimation of age begins; however, bomb radiocarbon and tag-recapture data indicated that large adult sharks were considerably older than the estimates derived from counts of growth bands. Three adult sandbar sharks were 20 to 26 years old based on bomb radiocarbon results and were a 5- to 11-year increase over the previous age estimates for these sharks. In support of these findings, the tag-recapture data provided results that were consistent with bomb radiocarbon dating and further supported a longevity that exceeds 30 years for this 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 dynamics of the spinner shark (Carcharhinus brevipinna) off the United States southeast and Gulf of Mexico coasts: a comparison of methods

The age and growth dynamics of the spinner shark (Carcharhinus brevipinna) in the northwest Atlantic Ocean off the southeast United States and in the Gulf of Mexico were examined and four growth models were used to examine variation in the ability to fit size-at-age data. The von Bertalanffy growth model, an alternative equation of the von Bertalanffy growth model with a size-at-birth intercept, the Gompertz growth model, and a logistic model were fitted to sex-specific observed size-at-age data. Considering the statistical criteria (e.g., lowest mean square error [MSE], high coefficient-of-determination, and greatest level of significance) we desired for this study, the logistic model provided the best overall fit to the size-at-age data, whereas the von Bertalanffy growth model gave the worst. For “biological validity,” the von Bertalanffy model for female sharks provided estimates similar to those reported in other studies. However, the von Bertalanffy model was deemed inappropriate for describing the growth of male spinner sharks because estimates of theoretical maximum size (L∞) indicated a size much larger than that observed in the field. However, the growth coefficient (k= 0.14/yr) from the Gompertz model provided an estimate most similar to that reported for other large coastal species. The analysis of growth for spinner shark in the present study demonstrates the importance of fitting alternative models when standard models fit the data poorly or when growth estimates do not appear to be realistic.

The Status of the United States Population of Night Shark, Carcharhinus signatus

Night sharks, Carcharhinus signatus, are an oceanic species generally occurring in outer continental shelf waters in the western North Atlantic Ocean including the Caribbean Sea and Gulf of Mexico. Although not targeted, night sharks make up a segment of the shark bycatch in the pelagic longline fishery. Historically, night sharks comprised a significant proportion of the artisanal Cuban shark fishery but today they are rarely caught. Although information from some fisheries has shown a decline in catches of night sharks, it is unclear whether this decline is due to changes in fishing tactics, market, or species identification. Despite the uncertainty in the decline, the night shark is currently listed as a species of concern due to alleged declines in abundance resulting from fishing effort, i.e. overutilization. To assess their relevance to the species of concern list, we collated available information on the night shark to provide an analysis of its status. Night shark landings were likely both over- and under-reported and thus probably did not reflect all commercial and recreational catches, and overall they have limited relevance to the current status of the species. Average size information has not changed considerably since the 1980’s based on information from the pelagic longline fishery when corrected for gear bias. Analysis of biological information indicates night sharks have intrinsic rates of increase (r) about 10% yr–1 and have moderate rebound potential and an intermediate generation time compared to other sharks. An analysis of trends in relative abundance from four data sources gave conflicting results, with one series in decline, two series increasing, and one series relatively flat. Based on the analysis of all currently available information, we believe the night shark does not qualify as a species of concern but should be retained on the prohibited species list as a precautionary approach to management until a more comprehensive stock assessment can be conducted.

Cook Inlet Belugas, Delphinapterus leucas: Status and Overview

Of the five populations of beluga, Delphinapterus leucas, in Alaska, the most isolated is the one in Cook Inlet (Hazard, 1988; Hill and DeMaster, 1998) (Fig. 1). The geographic and genetic segregation of this stock (O’Corry-Crowe et al., 1997), combined with the beluga’s tendency toward site fidelity in summer, makes this population especially vulnerable to impacts from large or persistent harvests.

Distribution of Belugas, Delphinapterus leucas, in Cook Inlet, Alaska, During June/July 1993–2000

Aerial surveys of belugas, Delphinapterus leucas, in Cook Inlet wre flown each year during June and/or July from 1993 to 2000. This project was designed to delineate distribution and collect aerial counts, elements critical to the managment of this small, isolated stock that was subjected to a persistent harvest by Native hunters. The surveys provided a thorough, annual coverage of the coastal areas of the inlet (1,300 km of shoreline) and included roughly 1,000 km of offshore transects annually. Coastal transects were flown 1.4 km from the waterline, thus surveying most of the area within 3 km of shore. These, along with offshore transects, provided annual systematic searches of 13-33% of the entire inlet. The largest concentration of belugas (151-288 whales by aerial count) was in the northern portion of upper Cook Inlet in the Susitna River Delta and/or in Knik Arm. Another concentration (17-49 whales) was consistently found between Chickaloon River and Point Possession. Smaller groups (generally <20 whales) were occasionally found in Turn-again Arm, Kachemak Bay, Redoubt Bay (Big River), and Trading Bay (McArthur River) prior to 1995 but not thereafter. Over the past three decades, summer distribution has shrunk such that sightings now only rarely occur in lower Cook Inlet and in offshore areas. In the 1990's, most (96-100%) of the sightings were concentrated in a few dense groups in shallow areas near river mouths in upper Cook Inlet.

Historic and Current Use of Lower Cook Inlet, Alaska, by Belugas, Delphinapterus leucas

Dedicated at-sea surveys for marine birds and mammals conducted in lower Cook Inlet in late July and early August from 1995–99 failed to locate any belugas, Delphinapterus leucas. Surveys covered a total of 6,249 linear km and were conducted in both nearshore and offshore habitats. Sightings included 791 individual marine mammals of 10 species. Both historical data and local knowledge indicate that belugas were regularly seen in summer in nearshore and offshore areas of lower Cook Inlet up until the early 1990’s. Diminished presence of belugas in lower Cook Inlet may be a direct function of reduced numbers but may also indicate changes in habitat quality that may inhibit recovery.

Beluga, Delphinapterus leucas, Distribution and Survey Effort in the Gulf of Alaska

Beluga, Delphinapterus leucas, distribution in the Gulf of Alaska and adjacent inside waters was examined through a review of surveys conducted as far back as 1936. Although beluga sightings have occurred on almost every marine mammal survey in northern Cook Inlet (over 20 surveys reported here), beluga sightings have been rare outside the inlet in the Gulf of Alaska. More than 150,000 km of dedicated survey effort in the Gulf of Alaska resulted in sightings of over 23,000 individual cetaceans, of which only 4 beluga sightings (5 individuals) occurred. In addition, nearly 100,000 individual cetaceans were reported in the Platforms of Opportunity database; yet, of these, only 5 sightings (39 individuals) were belugas. Furthermore, approximately 19 beluga sightings (>260 individuals), possibly including resightings, have been reported without information on effort or other cetacean sightings. Of the 28 sightings of belugas outside of Cook Inlet, 9 were near Kodiak Island, 10 were in or near Prince William Sound, 8 were in Yakutat Bay, and 1 anomalous sighting was well south of the Gulf. These sightings support archaeological and commercial harvest evidence indicating the only persistent group of belugas in the Gulf of Alaska occurs in Cook Inlet.

Abundance of Belugas, Delphinapterus leucas, in Cook Inlet, Alaska, 1994–2000

Annual abundance estimates of belugas, Delphinapterus leucas, in Cook Inlet were calculated from counts made by aerial observers and aerial video recordings. Whale group-size estimates were corrected for subsurface whales (availability bias) and whales that were at the surface but were missed (detection bias). Logistic regression was used to estimate the probability that entire groups were missed during the systematic surveys, and the results were used to calculate a correction to account for the whales in these missed groups (1.015, CV = 0.03 in 1994–98; 1.021, CV = 0.01 in 1999– 2000). Calculated abundances were 653 (CV = 0.43) in 1994, 491 (CV = 0.44) in 1995, 594 (CV = 0.28) in 1996, 440 (CV = 0.14) in 1997, 347 (CV = 0.29) in 1998, 367 (CV = 0.14) in 1999, and 435 (CV = 0.23, 95% CI=279–679) in 2000. For management purposes the current Nbest = 435 and Nmin = 360. These estimates replace preliminary estimates of 749 for 1994 and 357 for 1999. Monte Carlo simulations indicate a 47% probability that from June 1994 to June 1998 abundance of the Cook Inlet stock of belugas was depleted by 50%. The decline appears to have stopped in 1998.

Beluga, Delphinapterus leucas, Group Sizes in Cook Inlet, Alaska, Based on Observer Counts and Aerial Video

Belugas, Delphinapterus leucas, groups were videotaped concurrent to observer counts during annual NMFS aerial surveys of Cook Inlet, Alaska, from 1994 to 2000. The videotapes provided permanent records of whale groups that could be examined and compared to group size estimates ade by aerial observers.Examination of the video recordings resulted in 275 counts of 79 whale groups. The McLaren formula was used to account for whales missed while they were underwater (average correction factor 2.03; SD=0.64). A correction for whales missed due to video resolution was developed by using a second, paired video camera that magnified images relative to the standard video. This analysis showed that some whales were missed either because their image size fell below the resolution of hte standard video recording or because two whales surfaced so close to each other that their images appeared to be one large whale. The correction method that resulted depended on knowing the average whale image size in the videotapes. Image sizes were measured for 2,775 whales from 275 different passes over whale groups. Corrected group sizes were calcualted as the product of the original count from video, the correction factor for whales missed underwater, and the correction factor for whales missed due to video resolution (averaged 1.17; SD=0.06). A regression formula was developed to estimate group sizes from aerial observer counts; independent variables were the aerial counts and an interaction term relative to encounter rate (whales per second during the counting of a group), which were regressed against the respective group sizes as calculated from the videotapes. Significant effects of encounter rate, either positive or negative, were found for several observers. This formula was used to estimate group size when video was not available. The estimated group sizes were used in the annual abundance estimates.

Beluga, Delphinapterus leucas, Habitat Associations in Cook Inlet, Alaska

A review of available information describing habitat associations for belugas, Delphinapterus leucas, in Cook Inlet was undertaken to complement population assessment surveys from 1993-2000. Available data for physical, biological, and anthropogenic factors in Cook Inlet are summarized followed by a provisional description of seasonal habitat associations. To summarize habitat preferences, the beluga summer distribution pattern was used to partition Cook Inlet into three regions. In general, belugas congregate in shallow, relatively warm, low-salinity water near major river outflows in upper Cook Inlet during summer (defined as their primary habitat), where prey availability is comparatively high and predator occurrence relatively low. In winter, belugas are seen in the central inlet, but sightings are fewer in number, and whales more dispersed compared to summer. Belugas are associated with a range of ice conditions in winter, from ice-free to 60% ice-covered water. Natural catastrophic events, such as fires, earthquakes, and volcanic eruptions, have had no reported effect on beluga habitat, although such events likely affect water quality and, potentially, prey availability. Similarly, although sewage effluent and discharges from industrial and military activities along Cook Inlet negatively affect water quality, analyses of organochlorines and heavy metal burdens indicate that Cook Inlet belugas are not assimilating contaminant loads greater than any other Alaska beluga stocks. Offshore oil and gas activities and vessel traffic are high in the central inlet compared with other Alaska waters, although belugas in Cook Inlet seem habituated to these anthropogenic factors. Anthropogenic factors that have the highest potential negative impacts on belugas include subsistence hunts (not discussed in this report), noise from transportation and offshore oil and gas extraction (ship transits and aircraft overflights), and water quality degradation (from urban runoff and sewage treatment facilities). Although significant impacts from anthropogenic factors other than hunting are not yet apparent, assessment of potential impacts from human activities, especially those that may effect prey availability, are needed.