Analysis of the eastern Pacific yellowfin tuna fishery based on multiple management objectives

Vector optimization techniques were used to generate arbitrary segments of a policy frontier for a dynamic yellowfin tuna (Thunnus albacares) fishery model assuming fixed technology and considering four policy objectives: minimizing dolphin mortality, minimizing incidental catch (all species except dolphins), maximizing sustainable yield, and minimizing biological risk for the yellowfin tuna stock. Results show that along the policy frontier: (1) reducing incidental dolphin mortality increases the incidental catch of other species in a nonlinear way; (2) yield increases (subject to a biomass precautionary level) can only be obtained at the expense of higher levels of dolphin mortality and incidental catch; (3) biological risk increases as the level of tunas caught increases, but this increase depends on the type of fishery (longline fishing and three different modes of purse-seining: log-sets, dolphin-sets or school-sets) that dominates the fishing effort; (4) there is an indirect relationship between the dolphin mortality levels and those of biological risk; (5) there is a direct relationship between the incidental catch levels and biological risk. Catch obtained with dolphin-sets dominates the Pareto-optimal solutions with highest dolphin mortality levels but is associated with lower biological risk, whereas catch obtained with log-sets dominates in Pareto-optimal solutions with higher incidental catch and higher biological risk. In general, trade-offs or shadow prices among objectives are not linear, indicating that marginal costs vary along the policy frontier. Results of the trade-off analysis may provide useful information for decision-makers and other policy actors. Complete information about the preferences of the decision-makers regarding the objectives is necessary to recommend a specific management policy.     

Ecological metrics of biomass removed by three methods of purse-seine fishing for tunas in the eastern tropical Pacific Ocean

An ecosystem approach to fisheries management is a widely recognized goal, but describing and measuring the effects of a fishery on an ecosystem is difficult. Ecological information on the entire catch (all animals removed, whether retained or discarded) of both species targeted by the fishery and nontarget species (i.e., bycatch) is required. We used data from the well-documented purse-seine fishery for tunas (Thunnus albacares, T. obesus, and Katsuwonus pelamis) in the eastern tropical Pacific Ocean to examine the fishery's ecological effects. Purse-seine fishing in the eastern tropical Pacific is conducted in 3 ways that differ in the amount and composition of target species and bycatch. The choice of method depends on whether the tunas are swimming alone (unassociated sets), associated with dolphins (dolphin sets), or associated with floating objects (floating-object sets). Among the fishing methods, we compared catch on the basis of weight, number of individuals, trophic level, replacement time, and diversity. Floating-object sets removed 2-3 times as much biomass as the other 2 methods, depending on how removal was measured. Results of previous studies suggest the ecological effects of floating-object sets are thousands of times greater than the effects of other methods, but these results were derived from only numbers of discarded animals. Management of the fishery has been driven to a substantial extent by a focus on reducing bycatch, although discards are currently 4.8% of total catch by weight, compared with global averages of 7.5% for tuna longline fishing and 30.0% for midwater trawling. An ecosystem approach to fisheries management requires that ecological effects of fishing on all animals removed by a fishery, not just bycatch or discarded catch, be measured with a variety of metrics.     

Pregnancy patterns of pantropical spotted dolphins (Stenella attenuata) in the eastern tropical Pacific determined from hormonal analysis of blubber biopsies and correlations with the purse-seine tuna fishery

The northeastern offshore population of the pantropical spotted dolphin (Stenella attenuata) in the eastern tropical Pacific remains listed as depleted under the Marine Mammal Protection Act, and recent estimates of abundance show that recovery has been slow. One hypothesis for the slow recovery is that continued chase and encirclement by the tuna fishery negatively affects reproduction. Insufficient life-history sampling in this region over the last two decades makes traditional estimates of reproductive rates impossible. Here, we examine the current reproductive patterns of these dolphins by measuring blubber progesterone (BP) concentrations in biopsy samples to assess pregnancy state. BP was quantified in 212 biopsies from female offshore spotted dolphins sampled between 1998 and 2003 in the northeastern tropical Pacific, and we found that 11.5 % of the biopsied females (mature and immature) were pregnant. The relationship between pregnancy and fishery exposure was analyzed, and we found that pregnant females were exposed to significantly less fishery activity than non-pregnant ones (p = 0.022), suggesting that the fishery may have an inhibitive effect on pregnancy. Spatial analysis indicated that pregnancy was more aggregated than random (p < 0.05) at a scale up to 180-nmi, with the highest proportion pregnant in the mouth of the Gulf of California, an area with relatively low reported fishery activity.     

Vertical movements, behavior, and habitat of bigeye tuna (Thunnus obesus) in the equatorial eastern Pacific Ocean, ascertained from archival tag data

The results presented in this report are based on analyses of 16,721 days of data downloaded from 96 archival tags recovered from bigeye tuna (Thunnus obesus; 54–159 cm in length, 0.97–5.44 years of age) at liberty from 31 to 1,508 days in the equatorial eastern Pacific Ocean. Analyses of daily timed depth and temperature records resulted in the classification of the data into three daily behavior types: characteristic, associative (associated with floating objects), and other. There is a significant positive correlation between the proportion of time fish exhibit characteristic behavior and increasing length, and significant negative correlations between the proportion of time bigeye exhibit associative and other behavior with increasing length. For the smallest (54–80 cm) to largest (100–159 cm) length classes, the vertical habitats utilized when exhibiting non-associative behaviors were 99 and 98% of the time above the thermocline depth (60 m) during the night, at the same average depth of 34 m, and 60 and 72% of the time below the thermocline during the day at average depths of 163 and 183 m, respectively. For the same smallest to largest length classes, when exhibiting associative behavior, the average nighttime and daytime depths were 25 and 21, and 33 and 37 m, respectively. The apparent effects of the environment on the behavior of the fish are discussed.     

On the near-surface plankton of the eastern South Pacific Ocean

A study has been made of the plankton of the near-surface water layer (0 to 30 cm) of the eastern South Pacific Ocean in the region lying to the east of the meridian 90°W, between latitudes 5°N and 35°S. This region is influenced by the Peru Current: the current brings water from high latitudes, which results in a decrease in the number of species of the local fauna of copepods of the family Pontellidae, typical of tropical near-surface plankton. Some of the widely tropical and one bicentral species are absent or rare. Least affected by the Peru Current are the waters of equatorial structure in the northern part of the region. Here, 7 species of pontellids were recorded: the widely tropical Labidocera detruncata, Pontella tenuiremis, Pontellopsis regalis, the distant-neritic Pontella danae and Labidocera acuta, the bicentral Labidocera acutifrons, and the neritic Pontellopsis lubbockii. The dominant species among these are L. detruncata and L. acuta. To the west of the convergence, in the southern part of the region, live the southern central species Pontella valida and P. whiteleggei, with Pontellopsis regalis occurring occasionally. In these regions the copepod fauna is frequently dominated by pontellids. To the south of the boundary of the waters of equatorial structure, between the coast of South America and the line of convergence, lies a region most subjected to the effect of waters from high latitudes and of upwellings. It is inhabited by 2 pontellids only: Pontellopsis regalis and Labidocera acutifrons, but they too disappear close to the coast. In this particular region the copepods Calanus australis and Centropages brachiatus are common; they are found in a thicker water layer (0 to 200 m), and are often more abundant than the pontellids.     

Population structure of yellowfin tuna (Thunnus albacares) in the western Pacific Ocean, inferred from microsatellite loci

Five polymorphic microsatellite loci were examined in 1391 yellowfin tuna (Thunnus albacares) from eight regions of the western (Coral Sea, eastern Australia, Fiji, Indonesia, Philippines and Solomon Islands) and eastern (California and Mexico ) Pacific Ocean. Across all samples, numbers of alleles per locus ranged from 7 to 30 (mean: 17.0), and observed heterozygosities per locus ranged from 0.223 to 0.955 (mean: 0.593). Temporal collections were available for three areas: no significant temporal heterogeneity was observed for the Coral Sea (1991/1992 and 1995/1996 collections) or eastern Australia (1994/1995, 1995/1996, 1996/1997 and 1997/1998), but there was slight but significant heterogeneity at one locus (cmrTa-161) between the two Philippines collections (1994/1995 and 1996/1997). Genotypes generally showed a good fit to Hardy-Weinberg expectations within populations; only cmrTa-208 in the pooled Coral Sea population gave a significant deviation after Bonferroni correction for 40 tests, with a small but significant excess of homozygotes. Four loci showed no evidence of population differentiation following contingency Chi-squared and F_ST analyses. The fifth locus, cmrTa-161, showed small but significant differentiation (F_ST=0.002, P<0.001). This heterogeneity was largely a result of the Philippines 1994/1995 and Fiji collections; there was no correlation with geographic distance. The average F_ST across all five loci was very low (F_ST=0.002), but it was significant (P<0.001). It is unclear whether this low but significant differentiation reflects noise in the dataset, perhaps arising from experimental error, or real population differentiation. The finding of very limited population heterogeneity accords with most of the earlier allozyme and mitochondrial DNA studies of yellowfin tuna in the Pacific Ocean.     

Growth,dispersal, and identification of planktonic labrid and pomacentrid reef-fish larvae in the eastern Pacific Ocean

Planktonic larvae of six genera of labrid and pomacentrid reef fishes were captured in march 1985 in the eastern Pacific Ocean several hundred kilometers from the nearest reefs. The larvae were identified to genus by fin-ray counts as well as by comparison of their larval otolith morphology with that of known species. The larval otolith morphologies of known species were derived from measurements of the larval otolith embedded within the otoliths of settled juveniles (as delineated by the daily otolith-increment marks corresponding to the late larval period). The body morphology and melanophore patterns of the eastern Pacific labird and pomacentrid larvae closely matched those of congeneric larvae described from other oceans. Growth rates of larvae less than about 70 d old were similar between taxa (from 0.13 to 0.19 mm d^-1). After about 70 d in the plankton, labrid larvae grew much more slowly (0.06 mm d^-1 in Xyrichtys sp.). Labrid larvae had long larval durations (up to 131 d in Xyrichtys sp.), while the larval lives of the pomacentrids appeared to be shorter and much less variable. Larvae of many different ages occurred within the same water mass, and young cohorts of larvae appeared continuously over the sampling period. Some larvae were as young as 21 d, indicating that reef-fish larvae are capable of rapid long-distance dispersal (at least 18 km d^-1).     

Habitat,movements and environmental preferences of dusky sharks,Carcharhinus obscurus,in the northern Gulf of Mexico

The dusky shark (Carcharhinus obscurus) is the largest member of the genus Carcharhinus and inhabits coastal and pelagic ecosystems circumglobally in temperate, subtropical and tropical marine waters. In the western North Atlantic Ocean (WNA), dusky sharks are overfished and considered vulnerable by the International Union for the Conservation of Nature. As a result, retention of dusky sharks in commercial and recreational fisheries off the east coast of the United States (US) and in the northern Gulf of Mexico is prohibited. Despite the concerns regarding the status of dusky sharks in the WNA, little is known about their habitat utilization. During the summers of 2008–2009, pop-up satellite archival tags were attached to ten dusky sharks (one male, nine females) at a location where they have been observed to aggregate in the north central Gulf of Mexico southwest of the Mississippi River Delta to examine their movement patterns and habitat utilization. All tags successfully transmitted data with deployment durations ranging from 6 to 124 days. Tag data revealed shark movements in excess of 200 km from initial tagging locations, with sharks primarily utilizing offshore waters associated with the continental shelf edge from Desoto Canyon to the Texas/Mexican border. While most sharks remained in US waters, one individual moved from the northern Gulf of Mexico into the Bay of Campeche off the coast of Mexico. Sharks spent 87 % of their time between 20 and 125 m and 83 % of their time in waters between 23 and 30 °C. Since dusky sharks are among the most vulnerable shark species to fishing mortality, there is a recovery plan in place for US waters; however, since they have been shown to make long-distance migrations, a multi-national management plan within the WNA may be needed to ensure the successful recovery of this population.     

Allozyme and mitochondrial DNA variation in yellowfin tuna (Thunnus albacares) from the Pacific Ocean

Samples of yellowfin tuna (Thunnus albacares) collected in 1991 and 1992 from the western, central and eastern regions of the Pacific Ocean were examined for genetic variability. Four polymorphic allozyme loci (ADA ^*, FH ^*, GPI-S ^* and GPI-F ^*) were examined in all samples and a fifth polymorphism (GDA ^*) was examined in western and central samples only. Samples were also screened for mitochondrial DNA variation following restriction analysis by two enzymes (BcII and EcoRI) detecting polymorphic cut sites. Eighteen mtDNA haplotypes were revealed, with an overall nucleon diversity of 0.678. A subset of individuals screened for eight restriction enzymes had an overall nucleon diversity of 0.724 and a mean nucleotide diversity per sample of 0.359%. No significant spatial heterogeneity was detected for alleles at the ADA ^*, FH ^*, GPI-S ^* and GDA ^* loci nor for the mtDNA haplotypes. Significant heterogeneity was detected for GPI-F ^*. At this locus, the two eastern samples (southern California and northern Mexico) were not significantly different from each other but were significantly different (P<0.001) from the five western/central samples (Philippines, Coral Sea, Kiribati, Hawaii-91 and Hawaii-92). GPI-F ^* 100 was the most common allele in western and central regions, GPI-F ^* 75 the most common in eastern samples.     

Migration and habitat of white sharks (<Emphasis Type="Italic">Carcharodon carcharias</Emphasis>) in the eastern Pacific Ocean

Satellite telemetry studies of 20 adult and sub-adult white sharks (360–530 cm estimated total length (TL)) in the eastern North Pacific during 1999–2005 revealed long distance seasonal migrations from the coast of California to an offshore focal area 2,500 km west of the Baja Peninsula, as well as the Hawaii Islands. Three tags were recovered allowing detailed behavioral analyses, including one shark’s migration cycle from the coast to the offshore focal area and back. While near pinniped rookeries in autumn and winter, sharks avoided the surface and used water to 50 m depth, consistent with a silhouette-based hunting strategy. Offshore migrations were initiated during November–March and followed periods of decreasing pinniped abundance. Migrations were highly directed, taking 23 ± 5 days to reach the offshore focal area along similar paths among sharks and years, defining a migration corridor. Sharks exhibited a broad depth distribution (0–644 m) in the offshore focal area, and remained there for up to 167 days during spring and summer, though primary productivity and fishery data suggest that forage resources are scarcer there than in other regions of the eastern North Pacific. Archival data from one shark revealed intensive oscillatory movements while in the offshore focal area, a behavior that may be related to foraging or mating. Sharks traveling to Hawaii remained near the islands up to 122 days, potentially feeding on pelagic fishes and marine mammals that concentrate around the islands. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Regional differences in duration of the planktonic larval stage of reef fishes in the eastern Pacific Ocean

Regional variation in the duration of the planktonic larval phase of three species of reef fishes, Thalassoma lucasanum (Labridae), Stegastes flavilatus, and Microspathodon dorsalis (Pomacentridae) was investigated between 1982 and 1991 at several sites in the tropical eastern Pacific over a distance of 3500 km, encompassing virtually their entire range of distribution. Durations of the larval phase, determined from counts of daily otolith increments, were significantly different (1.3 to 1.6 x) between sites. Populations of all three species had a consistently shorter larval life at the most northern site, Cabo San Lucas (Mexico) compared to Panamá and the offshore islands of Galápagos and Cocos. Analyses of otolith increment width over the precompetent period revealed that this disparity in larval duration primarily reflected differences in larval growth rates: faster growing fish spent less time in the plankton. In T. lucasanum, some of the variation in larval duration between Panamá and offshore sites (Galápagos Islands and Cocos Island) may be accounted for by a higher frequency of individuals delaying metamorphosis at the offshore sites. These data indicate that conditions in the planktonic environment are not homogeneous throughout the tropical eastern Pacific and may have a profound effect on aspects of the larval ecology of reef fishes in this region.     

Biochemical genetic variation and evidence of speciation in Chthamalus barnacles of the tropical eastern Pacific Ocean

Gene-enzyme variation was studied electrophoretically within and between barnacle populations of the genus Chthamalus from 8 intertidal stations from central California to the Pacific coast of Panamá. Horizontal starch-gel electrophoresis separated and resolved 18 enzymes from 512 individual barnacles. A maximum of 25 gene-enzyme systems was interpretable from the resulting zymograms. Electrophoretic phenotypes and patterns of phenotypic variation generally conformed with those observed in other organisms. The amount of genetic variation within barnacle populations varied; average heterozygosity, for example, ranged from a low of 3.4% in a Mexican population of the C. fissus group to a high of 10.4% in C. anisopoma from the Gulf of California. Observed and expected average heterozygosities agreed in all population samples, indicating that these species are outbreeding. In contrast to the prediction stemming from a hypothesis that trophic stability regulates the amount of genetic variation in marine species, average heterozygosity tends to be positively correlated with latitude. Data from these and other barnacle species may support a hybrid environmental heterogeneity-trophic diversity model recently proposed to explain genetic variation in decapod crustaceans. Juxtaposition of individuals from different localities revealed numerous genetic differences among populations of the C. fissus group. At least three partially sympatric sibling species are separated by genetic distances as large as those observed between the C. fissus group and the distinct species C. dalli and C. anisopoma. A cladistic analysis places C. anisopoma close to the Mexican and Panamanian sibling species, with C. fissus from San Diego and C. dalli successively farther removed.     

Inter-island movements of scalloped hammerhead sharks (Sphyrna lewini) and seasonal connectivity in a marine protected area of the eastern tropical Pacific

Marine top predators are common at offshore bathymetric features such as islands, atolls, and seamounts, where most pelagic reef fish reside, while certain sharks perform inter-island movements between these formations. Scalloped hammerhead sharks are known to school in great numbers at small islands and seamounts in the eastern tropical Pacific (ETP) and are very susceptible to fisheries while moving into the open sea. It is, therefore, essential to understand hammerhead inter-island movements and environmental effects to provide baseline information for their conservation and management within and beyond an insular marine protected area. Movements of scalloped hammerheads were analyzed in the Galapagos Marine Reserve (GMR) and ETP, and environmental factors were linked to their movements. Hammerheads were tagged (N = 134) with V16 coded pingers (July 2006 to July 2010) in the northern Galapagos and detected at listening stations around four islands in the GMR and two isolated islands in the ETP, 700 and 1,200 km away. Hammerheads formed daytime schools at specific locations, but dispersed at night. Overall, more daytime than nighttime detections were recorded at all receivers in the northern Galapagos Islands, and more detections in the up-current sides of these islands. Hammerheads remained more days at the northern islands during part of the warm season (December–February) compared to the cool; however, fewer individuals were present in March–June. Movement modes were diel island excursions (24-h cycles) in the northern Galapagos and inter-island in the GMR and ETP at different scales: (1) short back-and-forth (<50 km, SBF), <5 days cycles, (2) medium distance (50–300 km, MDT), 5–20 days, and (3) long distance (>300 km, LDT), 15–52 days. The high degree of inter-island connectivity of hammerheads within the northern GMR is striking compared to the almost nil movement to the central GMR. A seasonal migratory pattern to locations offshore is indicated by (1) fewer hammerheads observed in the northern GMR during part of the warm season (March–June) and (2) evidence of LDT movements from the northern GMR to other islands in the ETP. LDT movements of mature female hammerheads are possibly associated with pupping areas. Our results indicate that currents, season, and individual behavior mainly drive inter-island movements of hammerheads at small (SBF) and medium (MDT) scales. These findings have important implications for the management of a highly mobile and endangered top predator within a marine protected area and beyond.     

Electrophoretic and biometric variability in the abyssal grenadier Coryphaenoides armatus of the western North Atlantic,eastern South Pacific and eastern North Pacific Oceans

Specimens of the abyssal grenadier Coryphaenoides armatus (Hector, 1875), from the western North Atlantic and eastern North Pacific Oceans were compared electrophoretically at 27 presumptive gene loci. At 6 of the 7 polymorphic loci there were only minor differences in allelic frequencies but a nearly fixed difference was found at one locus, phosphogluconate dehydrogenase. Eastern North Pacific grenadiers typically have a narrower interorbital space, a shorter dorsal interspace, more soft rays in the 1st dorsal fin (9–10 versus 8–9) and more pelvic fin rays (21–23 versus 18–21) than grenadiers from the western North Atlantic (as well as grenadiers from the eastern South Pacific, which were included in the biometric analysis). There is an apparent disjunction in the distribution of C. armatus in the eastern Pacific at the Gulf of Panamá which coincides with the change of morphology. It is suggested that North Pacific grenadiers comprise a subspecies, C. armatus variabilis Günther, 1878, which is morphologically distinct from the subspecies C. armatus armatus (Hector, 1875) of the other areas.     

Relationship between the growth and survival of larval Pacific bluefin tuna, Thunnus orientalis

We tested the hypothesis that a large body size and rapid growth rate affect the survival of larval Pacific bluefin tuna, Thunnus orientalis (PBT), and analyzed larval growth in relation to environmental conditions. Seven high density larval patches of PBT were tracked with reference buoys in the northwestern Pacific Ocean for 28–171 h in May–June from 2004 to 2008. The otolith radii and daily growth rates of the survivor larvae (collected on later tracking days of each tracking session) tended to be larger and more rapid, respectively, than those of original larvae (collected on earlier tracking days). A large body size was found to positively affect the survival of larval PBT, as did a rapid growth rate, even at an early larval stage (7 days after hatching). Generalized linear modeling showed that the otolith radius was influenced positively by the sea temperature, stratification parameter and food density, while the growth rate was influenced positively by the sea temperature and food density.     

Fecundity of a euphasiid crustacean,Nematoscelis difficilis,in the North Pacific Ocean

The number, size, and carbon and nitrogen contents of eggs in the pouches of a euphausiid crustacean, Nematoscelis difficilis Hansen, were examined. A clear linear relationship exists between the number of eggs in the pouch and the body weight of the maternal euphausiid. The eggs are not spherical in shape, and the size of eggs is greater in larger egg masses. The carbon content of the egg masses (50.2%) is higher than that of the bodies (40.7%). The carbon: nitrogen ratio (C/N) is also higher in the egg masses than in the bodies of euphausiids. The carbon content of the eggs is equivalent to 28.4% of the body carbon, the nitrogen content to 19.2% of the body nitrogen.     

Experimental effect of cold,La Niña temperatures on the survival of reef fishes from Gorgona Island (eastern Pacific Ocean)

The eastern tropical Pacific (ETP) reefs are affected at irregular times by extremely cold temperatures that occur principally during La Niña events. The effects of these low temperatures on the survival of reef fishes were experimentally assessed by determining the critical thermal minimum (CTM) of 15 reef fish species from Gorgona Island (ETP), and comparing these CTMs with the records of temperature during past La Niña events. Among species, mean CTMs ranged from 10.8°C to 16.3°C, which were lower than the coldest temperature recorded during the last La Niña event (18°C during La Niña 1998-1999). However, the observed ranges of CTM for two species (Thalassoma lucassanum and Eucinostomus gracilis) extended above 18°C. These results suggest that most of the reef fishes we studied are physiologically tolerant to the cold temperatures encountered during La Niña, though decreases in at least two populations may be expected as a result of the mortality of less tolerant individuals. Although tolerant to cold temperatures, reef fish populations may still experience negative changes during La Niña, because other determinants in population maintenance (e.g. reproduction and recruitment) are more temperature sensitive. The effects of other cold phenomena on reef fish survival are also discussed herein.