Sizes of walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius) consumed by the western stock of Steller sea lions (Eumetopias jubatus) in Alaska from 1999 to 2000

Prey-size selectivity by Steller sea lions (Eumetopias jubatus) is relevant for understanding the foraging behavior of this declining predator, but studies have been problematic because of the absence and erosion of otoliths usually used to estimate fish length. Therefore, we developed regression formulae to estimate fish length from seven diagnostic cranial structures of walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius). For both species, all structure measurements were related with fork length of prey (r2 range: 0.78−0.99). Fork length (FL) of walleye pollock and Atka mackerel consumed by Steller sea lions was estimated by applying these regression models to cranial structures recovered from scats (feces) collected between 1998 and 2000 across the range of the Alaskan western stock of Steller sea lions. Experimentally derived digestion correction factors were applied to take into account loss of size due to digestion. Fork lengths of walleye pollock consumed by Steller sea lions ranged from 3.7 to 70.8 cm (mean=39.3 cm, SD=14.3 cm, n=666) and Atka mackerel ranged from 15.3 to 49.6 cm (mean=32.3 cm, SD=5.9 cm, n=1685). Although sample sizes were limited, a greater proportion of juvenile (≤20 cm) walleye pollock were found in samples collected during the summer (June−September) on haul-out sites (64% juveniles, n=11 scats) than on summer rookeries (9% juveniles, n=132 scats) or winter (February−March) haul-out sites (3% juveniles, n=69 scats). Annual changes in the size of Atka mackerel consumed by Steller sea lions corresponded to changes in the length distribution of Atka mackerel resulting from exceptionally strong year classes. Considerable overlap (>51%) in the size of walleye pollock and Atka mackerel taken by Steller sea lions and the sizes of these species caught by the commercial trawl fishery were demonstrated.

Sizes of walleye pollock (Theragra chalcogramma) consumed by the eastern stock of Steller sea lions (Eumetopias jubatus) in Southeast Alaska from 1994 to 1999

Lengths of walleye pollock (Theragra chalcogramma) consumed by Steller sea lions (Eumetopias jubatus) were estimated by using allometric regressions applied to seven diagnostic cranial structures recovered from 531 scats collected in Southeast Alaska between 1994 and 1999. Only elements in good and fair condition were selected. Selected structural measurements were corrected for loss of size due to erosion by using experimentally derived condition-specific digestion correction factors. Correcting for digestion increased the estimated length of fish consumed by 23%, and the average mass of fish consumed by 88%. Mean corrected fork length (FL) of pollock consumed was 42.4 ±11.6 cm (range=10.0−78.1 cm, n=909). Adult pollock (FL>45.0 cm) occurred more frequently in scats collected from rookeries along the open ocean coastline of Southeast Alaska during June and July (74% adults, mean FL=48.4 cm) than they did in scats from haul-outs located in inside waters between October and May (51% adults, mean FL=38.4 cm). Overall, the contribution of juvenile pollock (≤20 cm) to the sea lion diet was insignificant; whereas adults contributed 44% to the diet by number and 74% by mass. On average, larger pollock were eaten in summer at rookeries throughout Southeast Alaska than at rookeries in the Gulf of Alaska and the Bering Sea. Overall it appears that Steller sea lions are capable of consuming a wide size range of pollock, and the bulk of fish fall between 20 and 60 cm. The use of cranial hard parts other than otoliths and the application of digestion correction factors are fundamental to correctly estimating the sizes of prey consumed by sea lions and determining the extent that these sizes overlap with the sizes of pollock caught by commercial fisheries.

Identification of formalin-preserved eggs of red sea bream (Pagrus major) (Pisces: Sparidae) using monoclonal antibodies

Catches of important commercial fish such as red sea bream, flat fish, and yellowtail are decreasing in Japan. In order to sustain these species it is especially important that their distribution and biomass at all life stages are known. However, information on the early life stages of these species is limited because identifying the eggs and larvae of such fish is sometimes extremely difficult.

Magnitude and distribution of sea turtle bycatch in the sea scallop (Placopecten magellanicus) dredge fishery in two areas of the northwestern Atlantic Ocean, 2001–2002

In May 2001, the National Marine Fisheries Service (NMFS) opened two areas in the northwestern Atlantic Ocean that had been previously closed to the U.S. sea scallop (Placopecten magellanicus) dredge fishery. Upon reopening these areas, termed the “Hudson Canyon Controlled Access Area” and the “Virginia Beach Controlled Access Area,” NMFS observers found that marine turtles were being caught incidentally in scallop dredges. This study uses the generalized linear model and the generalized additive model fitting techniques to identify environmental factors and gear characteristics that influence bycatch rates, and to predict total bycatch in these two areas during May-December 2001 and 2002 by incorporating environmental factors into the models. Significant factors affecting sea turtle bycatch were season, time-of-day, sea surface temperature, and depth zone. In estimating total bycatch, rates were stratified according to a combination of all these factors except time-of-day which was not available in fishing logbooks. Highest bycatch rates occurred during the summer season, in temperatures greater than 19°C, and in water depths from 49 to 57 m. Total estimated bycatch of sea turtles during May–December in 2001 and 2002 in both areas combined was 169 animals (CV=55.3), of which 164 (97%) animals were caught in the Hudson Canyon area. From these findings, it may be possible to predict hot spots for sea turtle bycatch in future years in the controlled access areas.

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.

Yield- and biomass-per-recruit analysis for rotational fisheries, with an application to the Atlantic sea scallop (Placopecten magellanicus)

A general model for yield-per-recruit analysis of rotational (periodic) fisheries is developed and applied to the sea scallop (Placopecten magellanicus) fishery of the northwest Atlantic. Rotational fishing slightly increases both yield- and biomass-per-recruit for sea scallops at FMAX. These quantities decline less quickly when fishing mortality is increased beyond FMAX than when fishing is at a constant rate. The improvement in biomass-per-recruit appears to be nearly independent of the selectivity pattern but increased size-at-entry can reduce or eliminate the yield-per-recruit advantage of rotation. Area closures and rotational fishing can cause difficulties with the use of standard spatially averaged fishing mortality metrics and reference points. The concept of temporally averaged fishing mortality is introduced as one that is more appropriate for sedentary resources when fishing mortality varies in time and space.

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

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.

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.

Developing a growth-transition matrix for the stock assessment of the green sea urchin (Strongylocentrotus droebachiensis) off Maine

The green sea urchin (Strongylocentrotus droebachiensis) is important to the economy of Maine. It is the state’s fourth largest fishery by value. The fishery has experienced a continuous decline in landings since 1992 because of decreasing stock abundance. Because determining the age of sea urchins is often difficult, a formal stock assessment demands the development of a size-structured population dynamic model. One of the most important components in a size-structured model is a growth-transition matrix. We developed an approach for estimating the growth-transition matrix using von Bertalanffy growth parameters estimated in previous studies of the green sea urchin off Maine. This approach explicitly considers size-specific variations associated with yearly growth increments for these urchins. The proposed growth-transition matrix can be updated readily with new information on growth, which is important because changes in stock abundance and the ecosystem will likely result in changes in sea urchin key life history parameters including growth. This growth-transition matrix can be readily incorporated into the size-structured stock assessment model that has been developed for assessing the green sea urchin stock off Maine.

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.