Validation and interpretation of annual skeletal marks in loggerhead (Caretta caretta) and Kemp’s ridley (Lepidochelys kempii) sea turtles

Numerous studies have applied skeletochronology to sea turtle species. Because many of the studies have lacked validation, the application of this technique to sea turtle age estimation has been called into question. To address this concern, we obtained humeri from 13 known-age Kemp’s ridley (Lepidochelys kempii) and two loggerhead (Caretta caretta) sea turtles for the purposes of examining the growth marks and comparing growth mark counts to actual age. We found evidence for annual deposition of growth marks in both these species. Corroborative results were found in Kemp’s ridley sea turtles from a comparison of death date and amount of bone growth following the completion of the last growth mark (n=76). Formation of the lines of arrested growth in Kemp’s ridley sea turtles consistently occurred in the spring for animals that strand dead along the mid- and south U.S. Atlantic coast. For both Kemp’s ridley and loggerhead sea turtles, we also found a proportional allometry between bone growth (humerus dimensions) and somatic growth (straight carapace length), indicating that size-at-age and growth rates can be estimated from dimensions of early growth marks. These results validate skeletochronology as a method for estimating age in Kemp’s ridley and loggerhead sea turtles from the southeast United States.

Prey consumption of Steller sea lions (Eumetopias jubatus) off Alaska: How much prey do they require?

The effects of seasonal and regional differences in diet composition on the food requirements of Steller sea lions (Eumetopias jubatus) were estimated by using a bioenergetic model. The model considered differences in the energy density of the prey, and differences in digestive efficiency and the heat increment of feeding of different diets. The model predicted that Steller sea lions in southeast Alaska required 45–60% more food per day in early spring (March) than after the breeding season in late summer (August) because of seasonal changes in the energy density of the diets (along with seasonal changes in energy requirements). The southeast Alaska population, at 23,000 (±1660 SD) animals (all ages), consumed an estimated 140,000 (±27,800) t of prey in 1998. In contrast, we estimated that the 51,000 (±3680) animals making up the western Alaska population in the Gulf of Alaska and Aleutian Islands consumed just over twice this amount (303,000 [±57,500] t). In terms of biomass removed in 1998 from Alaskan waters, we estimated that Steller sea lions accounted for about 5% of the natural mortality of gadids (pollock and cod) and up to 75% of the natural mortality of hexagrammids (adult Atka mackerel). These two groups of species were consumed in higher amounts than any other. The predicted average daily food requirement per individual ranged from 16 (±2.8) to 20 (±3.6) kg (all ages combined). Per capita food requirements differed by as much as 24% between regions of Alaska depending on the relative amounts of low–energy-density prey (e.g. gadids) versus high–energy-density prey (e.g. forage fish and salmon) consumed. Estimated requirements were highest in regions where Steller sea lions consumed higher proportions of low–energy-density prey and experienced the highest rates of population decline

Dynamics of bigeye (Thunnus obesus) and yellowfin (T. albacares) tuna in Hawaii’s pelagic fisheries: analysis of tagging data with a bulk transfer model incorporating size-specific attrition

Tag release and recapture data of bigeye (Thunnus obesus) and yellowfin tuna (T. albacares) from the Hawaii Tuna Tagging Project (HTTP) were analyzed with a bulk transfer model incorporating size-specific attrition to infer population dynamics and transfer rates between various fishery components. For both species, the transfer rate estimates from the offshore handline fishery areas to the longline fishery area were higher than the estimates of transfer from those same areas into the inshore fishery areas. Natural and fishing mortality rates were estimated over three size classes: yellowfin 20–45, 46–55, and ≥56 cm and bigeye 29–55, 56–70, and ≥71 cm. For both species, the estimates of natural mortality were highest in the smallest size class. For bigeye tuna, the estimates decreased with increasing size and for yellowfin tuna there was a slight increase in the largest size class. In the Cross Seamount fishery, the fishing mortality rate of bigeye tuna was similar for all three size classes and represented roughly 12% of the gross attrition rate (includes fishing and natural mortality and emigration rates). For yellowfin tuna, fishing mortality ranged between 7% and 30%, the highest being in the medium size class. For both species, the overall attrition rate from the entire fishery area was nearly the same. However, in the specific case of the Cross Seamount fishery, the attrition rate for yellowfin tuna was roughly twice that for bigeye. This result indicates that bigeye tuna are more resident at the Seamount than yellowfin tuna, and larger bigeye tunas tend to reside longer than smaller individuals. This may result in larger fish being more vulnerable to capture in the Seamount fishery. The relatively low level of exchange between the Sea-mount and the inshore and longline fisheries suggests that the fishing activity at the Seamount need not be of great management concern for either species. However, given that the current exploitation rates are considered moderate (10–30%), and that Seamount aggregations of yellowfin and bigeye tuna are highly vulnerable to low-cost gear types, it is recommended that further increases in fishing effort for these species be monitored at Cross Seamount.

Larval development of the southern sea garfish (Hyporhamphus melanochir) and the river garfish (H. regularis) (Beloniformes: Hemiramphidae) from South Australian waters

Larval development of the southern sea garfish (Hyporhamphus melanochir) and the river garfish (H. regularis) is described from specimens from South Australian waters. Larvae of H. melanochir and H. regularis have completed notochord flexion at hatching and are characterized by an elongate body with distinct rows of melanophores along the dorsal, lateral, and ventral surfaces; a small to moderate head; a heavily pigmented and long straight gut; a persistent pre-anal finfold; and an extended lower jaw. Fin formation occurs in the following sequence: caudal, dorsal and anal (almost simultaneously), pectoral, and pelvic. Despite the similarities between both species and among hemiramphid larvae in general, H. melanochir larvae are distinguishable from H. regularis by 1) having 58–61 vertebrae (vs. 51–54 for H. regularis); 2) having 12–15 melanophore pairs in longitudinal rows along the dorsal margin between the head and origin of the dorsal fin (vs. 19–22 for H. regularis); and 3) the absence of a large ventral pigment blotch anteriorly on the gut and isthmus (present in H. regularis). Both species can be distinguished from similar larvae of southern Australia (other hemiramphids and a scomberosocid) by differences in meristic counts and pigmentation.

Age and growth estimates of the winter skate (Leucoraja ocellata) in the western Gulf of Maine

Age and growth estimates for the winter skate (Leucoraja ocellata) were estimated from vertebral band counts on 209 fish ranging in size from 145 to 940 mm total length (TL). An index of average percent error (IAPE) of 5.8% suggests that our aging method represents a precise approach to the age assessment of L. ocellata. Marginal increments were significantly different between months (Kruskal-Wallis P<0.001) and a distinct trend of increasing monthly increment growth began in July. Estimates of von Bertalanffy growth parameters suggest that females attain a slightly larger asymptotic TL (L∞=1374 mm) than males (L∞=1218 mm) and grow more slowly (k=0.059 and 0.074, respectively). The oldest ages obtained for the winter skate were 19 years for males and 18 years for females, which corresponded to total lengths of 932 mm and 940 mm, respectively. The results indicate that the winter skate exhibits the characteristics that have made other elasmobranch populations highly susceptible to exploitation by commercial fisheries.

Estimation of bycatch in shrimp trawl fisheries: a comparison of estimation methods using field data and simulated data

Bycatch, or the incidental catch of nontarget organisms during fi shing operations, is a major issue in U.S. shrimp trawl fisheries. Because bycatch is typically discarded at sea, total bycatch is usually estimated by extrapolating from an observed bycatch sample to the entire fleet with either mean-per-unit or ratio estimators. Using both field observations of commercial shrimp trawlers and computer simulations, I compared five methods for generating bycatch estimates that were used in past studies, a mean-per-unit estimator and four forms of the ratio estimator, respectively: 1) the mean fish catch per unit of effort, where unit effort was a proxy for sample size, 2) the mean of the individual fish to shrimp ratios, 3) the ratio of mean fish catch to mean shrimp catch, 4) the mean of the ratios of fish catch per time fished (a variable measure of effort), and 5) the ratio of mean fish catch per mean time fished. For field data, different methods used to estimate bycatch of Atlantic croaker, spot, and weakfish yielded extremely different results, with no discernible pattern in the estimates by method, geographic region, or species. Simulated fishing fleets were used to compare bycatch estimated by the fi ve methods with “actual” (simulated) bycatch. Simulations were conducted by using both normal and delta lognormal distributions of fish and shrimp and employed a range of values for several parameters, including mean catches of fish and shrimp, variability in the catches of fish and shrimp, variability in fishing effort, number of observations, and correlations between fish and shrimp catches. Results indicated that only the mean per unit estimators provided statistically unbiased estimates, while all other methods overestimated bycatch. The mean of the individual fish to shrimp ratios, the method used in the South Atlantic Bight before the 1990s, gave the most biased estimates. Because of the statistically significant two- and 3-way interactions among parameters, it is unlikely that estimates generated by one method can be converted or corrected to estimates made by another method: therefore bycatch estimates obtained with different methods should not be compared directly.

Diving behavior of immature Steller sea lions (Eumetopias jubatus)

Understanding the ontogenetic relationship between juvenile Steller sea lions (Eumetopias jubatus) and their foraging habitat is key to understanding their relationship to available prey and ultimately their survival. We summarize dive and movement data from 13 young-of-the-year (YOY) and 12 yearling Steller sea lions equipped with satellite dive recorders in the Gulf of Alaska and Aleutian Islands (n=18), and Washington (n=7) from 1994 to 2000. A total of 1413 d of transmission (x =56.5 d, range: 14.5–104.1 d) were received. We recorded 222,073 dives, which had a mean depth of 18.4 m (range of means: 5.8−67.9 m; SD=16.4). Alaska YOY dived for shorter periods and at shallower depths (mean depth=7.7 m, mean duration=0.8 min, mean maximum depth=25.7 m, and maximum depth=252 m) than Alaska yearlings (x =16.6 m, 0=1.1 min, x = 63.4 m, 288 m), whereas Washington yearlings dived the longest and deepest (mean depth=39.4 m, mean duration=1.8 min, mean maximum depth=144.5 m, and maximum depth=328 m). Mean distance for 564 measured trips was 16.6 km; for sea lions ≤10 months of age, trip distance (7.0 km) was significantly less than for those >10 months of age (24.6 km). Mean trip duration for 10 of the 25 sea lions was 12.1 h; for sea lions ≤10 months of age, trip duration was 7.5 h and 18.1 h for those >10 months of age. We identified three movements types: long-range trips (>15 km and >20 h), short-range trips (<15 km and <20 h) during which the animals left and returned to the same site, and transits to other haul-out sites. Long-range trips started around 9 months of age and occurred most frequently around the assumed time of weaning, whereas short-range trips happened almost daily (0.9 trips/day, n=426 trips). Transits began as early as 7 months of age, occurred more often after 9 months of age, and ranged between 6.5 and 454 km. The change in dive characteristics coincided with the assumed onset of weaning. These yearling sea lion movement patterns and dive characteristics suggest that immature Steller sea lions are as capable of making the same types of movements as adults.

Modeling red sea urchin (Strongylocentrotus franciscanus) growth using six growth functions

The growth of red sea urchins (Strongylocentrotus franciscanus) was modeled by using tag-recapture data from northern California. Red sea urchins (n=211) ranging in test diameter from 7 to 131 mm were examined for changes in size over one year. We used the function Jt+1 = Jt + f(Jt) to model growth, in which Jt is the jaw size (mm) at tagging, and Jt+1 is the jaw size one year later. The function f(Jt), represents one of six deterministic models: logistic dose response, Gaussian, Tanaka, Ricker, Richards, and von Bertalanffy with 3, 3, 3, 2, 3, and 2 minimization parameters, respectively. We found that three measures of goodness of fi t ranked the models similarly, in the order given. The results from these six models indicate that red sea urchins are slow growing animals (mean of 7.2 ±1.3 years to enter the fishery). We show that poor model selection or data from a limited range of urchin sizes (or both) produces erroneous growth parameter estimates and years-to-fishery estimates. Individual variation in growth dominated spatial variation at shallow and deep sites (F=0.246, n=199, P=0.62). We summarize the six models using a composite growth curve of jaw size, J, as a function of time, t: J = A(B – e–Ct) + Dt, in which each model is distinguished by the constants A, B, C, and D. We suggest that this composite model has the flexibility of the other six models and could be broadly applied. Given the robustness of our results regarding the number of years to enter the fishery, this information could be incorporated into future fishery management plans for red sea urchins in northern California.

Estimates of survival probabilities for oceanic-stage loggerhead sea turtles (Caretta caretta) in the North Atlantic

Estimates of instantaneous mortality rates (Z) and annual apparent survival probabilities (Φ) were generated from catch-curve analyses for oceanic-stage juvenile loggerheads (Caretta caretta) in the waters of the Azores. Two age distributions were analyzed: the “total sample” of 1600 loggerheads primarily captured by sighting and dipnetting from a variety of vessels in the Azores between 1984 and 1995 and the “tuna sample” of 733 loggerheads (a subset of the total sample) captured by sighting and dipnetting from vessels in the commercial tuna fleet in the Azores between 1990 and 1992. Because loggerhead sea turtles begin to emigrate from oceanic to neritic habitats at age 7, the best estimates of instantaneous mortality rate (0.094) and annual survival probability (0.911) not confounded with permanent emigration were generated for age classes 2 through 6. These estimates must be interpreted with caution because of the assumptions upon which catch-curve analyses are based. However, these are the first directly derived estimates of mortality and survival probabilities for oceanic-stage sea turtles. Estimation of survival probabilities was identified as “an immediate and critical requirement” in 2000 by the Turtle Expert Working Group of the U.S. National Marine Fisheries Service.

The physiological effects of multiple forced submergences in loggerhead sea turtles (Caretta caretta)

Sea turtles are subjected to involuntary submergence and potential mortality due to incidental capture by the commercial shrimp fishing industry. Despite implementation of turtle excluder devices (TEDs) to reduce at-sea mortality, dead stranded turtles continue to be found in near-record numbers along the coasts of the western Atlantic Ocean and northern Gulf of Mexico. Although this mortality may be due to an increase in the number of turtles available to strand, one alternative explanation is that sea turtles are repetitively submerged (as one fishing vessel follows the path of another) in legal TEDs. In the present study, laboratory and field investigations were undertaken to examine the physiological effects of multiple submergence of loggerhead sea turtles (Caretta caretta). Turtles in the laboratory study were confined during the submersion episodes, whereas under field conditions, turtles were released directly into TED-equipped commercial fishing nets. Under laboratory and field conditions, pre- and postsubmergence blood samples were collected from turtles submerged three times at 7.5 min per episode with an in-water rest interval of 10, 42, or 180 min between submergences. Analyses of pre- and postsubmergence blood samples revealed that the initial submergence produced a severe and pronounced metabolic and respiratory acidosis in all turtles. Successive submergences produced significant changes in blood pH, Pco2, and lactate, although the magnitude of the acid-base imbalance was substantially reduced as the number of submergences increased. In addition, increasing the interval between successive submergences permitted greater recovery of blood homeostasis. No turtles died during these studies. Taken together, these data suggest that repetitive sub-mergence of sea turtles in TEDs would not significantly affect their survival potential provided that the animal has an adequate rest interval at the surface between successive submergences.

Population status, seasonal variation in abundance, and long-term population trends of Steller sea lions (Eumetopias jubatus) at the South Farallon Islands, California

We examined seasonal and annual variation in numbers of Steller (northern) sea lions (Eumetopias jubatus) at the South Farallon Islands from counts conducted weekly from 1974 to 1996. Numbers of adult and subadult males peaked during the breeding season (May–July), whereas numbers of adult females and immature individuals peaked during the breeding season and from late fall through early winter (September–December). The seasonal pattern varied significantly among years for all sexes and age classes. From 1977 to 1996, numbers present during the breeding season decreased by 5.9% per year for adult females and increased by 1.9% per year for subadult males. No trend in numbers of adult males was detected. Numbers of immature individuals also declined by 4.5% per year during the breeding season but increased by 5.0% per year from late fall through early winter. Maximum number of pups counted declined significantly through time, although few pups were produced at the South Farallon Islands. The ratio of adult females to adult males averaged 5.2:1 and declined significantly with each year, whereas no trend in the ratio of pups to adult females was discernible. Further studies are needed to determine if reduced numbers of adult females in recent years have resulted from reduced survival of juvenile or adult females or from changes in the geographic distribution of females.

Recovery of the Gulf of Maine--Georges Bank Atlantic herring (Clupea harengus) complex: perspectives based on bottom trawl survey data

NMFS bottom trawl survey data were used to describe changes in distribution, abundance, and rates of population change occurring in the Gulf of Maine–Georges Bank herring (Clupea harengus) complex during 1963–98. Herring in the region have fully recovered following severe overfishing during the 1960s and 1970s. Three distinct, but seasonally intermingling components from the Gulf of Maine, Nantucket Shoals (Great South Channel area), and Georges Bank appear to compose the herring resource in the region. Distribution ranges contracted as herring biomass declined in the late 1970s and then the range expanded in the 1990s as herring increased. Analysis of research survey data suggest that herring are currently at high levels of abundance and biomass. All three components of the stock complex, including the Georges Bank component, have recovered to pre-1960s abundance. Survey data support the theory that herring recolonized the Georges Bank region in stages from adjacent components during the late 1980s, most likely from herring spawning in the Gulf of Maine.

Origin of immature loggerhead sea turtles (Caretta caretta) at Hutchinson Island, Florida: evidence from mtDNA markers

Loggerhead sea turtles (Caretta caretta) are migratory, long-lived, and slow maturing. They are difficult to study because they are seen rarely and their habitats range over vast stretches of the ocean. Movements of immature turtles between pelagic and coastal developmental habitats are particularly difficult to investigate because of inadequate tagging technologies and the difficulty in capturing significant numbers of turtles at sea. However, genetic markers found in mitochondrial DNA (mtDNA) provide a basis for predicting the origin of juvenile turtles in developmental habitats. Mixed stock analysis was used to determine which nesting populations were contributing individuals to a foraging aggregation of immature loggerhead turtles (mean 63.3 cm straight carapace length [SCL]) captured in coastal waters off Hutchinson Island, Florida. The results indicated that at least three different western Atlantic loggerhead sea turtle subpopulations contribute to this group: south Florida (69%), Mexico (20%), and northeast Florida-North Carolina (10%). The conservation and management of these immature sea turtles is complicated by their multinational genetic demographics.