Movements,behavior, and habitat utilization of yellowfin tuna (<Emphasis Type="Italic">Thunnus albacares</Emphasis>) in the northeastern Pacific Ocean,ascertained through archival tag data

Sixty-eight yellowfin tuna, Thunnus albacares, (60-135 cm fork length) were caught and released with implanted archival tags offshore off Baja California, Mexico, during October 2002 and October 2003. Thirty-six fish (53%) were recaptured and the data were downloaded from all 36 recovered tags. Time at liberty ranged from 9 to 1,161 days, and the data were analyzed for the 20 fish that were at liberty for 154 or more days. The accuracy in the position estimates, derived from light-level longitude data and sea-surface temperatures (SSTs) based latitude, is about 0.41° in longitude and 0.82° in latitude, in this region. The movement paths, derived from position estimates, for the 20 yellowfin indicated that 19 (95%) remained within 1,445 km of their release locations. The estimated mean velocity along movement paths was 77 km/day. The southern and northern seasonal movement paths observed for yellowfin off Baja California are influenced by the seasonal movements of the 18°C SST isotherm. Cyclical movements to and from suitable spawning habitat (≥24°C SST) was observed only for mature fish. For the 12 fish that demonstrated site fidelity, the mean 95 and 50% utilization distributions were 258,730 km² and 41,260 km², respectively. Evaluations of the timed depth records resulted in discrimination of four distinct behaviors. When exhibiting type-1 diving behavior (78.1% of all days at liberty) the fish remained at depths less than 50 m at night and did not dive to depths greater than about 100 m during the day. Type-2 diving behavior (21.2% of all days at liberty) was characterized by ten or more dives in excess of 150 m during the day. Type-2 diving behavior is apparently a foraging strategy for fish targeting prey organisms of the deep-scattering layer during the day, following nighttime foraging within the mixed layer on the same prey. Yellowfin tuna exhibited occasional deep-diving behavior, and some dives exceeded 1,000 m, where ambient temperatures were less than 5°C. Surface-oriented behavior, defined as the time fish remained at depths less than 10 m for more than 10 min, were evaluated. The mean number and duration of surface-oriented events per day for all fish was 14.3 and 28.5 min, respectively. Habitat utilization of yellowfin, presented as monthly composite horizontal and vertical distributions, indicates confined geographical distributions, apparently resulting from an affinity to an area of high prey availability. The vertical distributions indicate greater daytime depths in relation to a seasonally deeper mixed layer and a greater proportion of daytime at shallower depths in relation to a seasonally shallower mixed layer.     

Genetic evidence for inter-oceanic subdivision of bigeye tuna (Thunnus obesus) populations

Bigeye tuna (Thunnus obesus Lowe, 1839) are a commercially important species of tuna found in the Atlantic, Indian, and Pacific oceans. To initiate an analysis of bigeye tuna population-structure, three PCR–RFLP assays were developed based on the published mtDNA control-region sequences of four bigeye tuna. Population analyses using these three restriction assays on a total of 248 individuals resulted in an array of 13 composite haplotypes. A total of 347 nucleotides of mtDNA control-region sequence was characterized for 11 of the 13 composite haplotypes. Phylogenetic analyses demonstrated that the DNA sequences belong to two monophyletic clades. However, only one of the three restriction assays was able to discriminate between the two clades. The other two assays were confounded by excessive homoplasy. Both parallel (independent occurrences of the same nucleotide change) and convergent (different nucleotide changes within the same restriction site) changes of restriction sites were observed. These results emphasize the importance of DNA sequence-analysis for the interpretation of restriction-site polymorphism data. Analyses of the frequency distribution indicated that samples of bigeye tuna from the Atlantic Ocean were genetically distinct from those found in the Indian and Pacific oceans. Thus, these results reject the null hypothesis of a single global population of bigeye tuna. Received: 16 June 1997 / Accepted: 25 July 1998     

Movement patterns of large bigeye tuna (Thunnus obesus) in the open ocean, determined using ultrasonic telemetry

 The horizontal and vertical movements of large bigeye tuna (Thunnus obesus Lowe, 1839; 25 to 50 kg) captured in the south Pacific Ocean (French Polynesia) were determined using pressure-sensitive ultrasonic transmitters. Bigeye tuna swam within the first 100 m below the surface during the night-time and at depths between 400 and 500 m during the daytime. The fish exhibited clear relationships with the sound scattering layer (SSL). They followed its vertical movements at dawn and dusk, and were probably foraging on the organisms of the SSL. Bigeye tuna did, however, make regular rapid upward vertical excursions into the warm surface layer, most probably in order to regulate body temperature and, perhaps, to compensate for an accumulated oxygen debt (i.e. to metabolize lactate). The characteristics of these dives differ from those reported from previous studies on smaller bigeye tuna (∼12 kg) near the main Hawaiian Islands. During the daytime, the large fish in French Polynesia made upward excursions approximately only every 2.5 h, whereas smaller fish in Hawaiian waters made upward excursions approximately every hour. Our data are the first observations on the role of body size in the vertical behavior of bigeye tuna. Received: 9 September 1998 / Accepted: 25 November 1999     

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.     

Simultaneous behavior of skipjack (Katsuwonus pelamis), bigeye (Thunnus obsesus), and yellowfin (T. albacares) tunas, within large multi-species aggregations associated with drifting fish aggregating devices (FADs) in the equatorial eastern Pacific Ocean

Ten separate experiments monitoring the simultaneous behaviors of 26 skipjack (Katsuwonus pelamis), 26 bigeye (Thunnus obesus), and 33 yellowfin (T. albacares) tunas within large multi-species aggregations associated with drifting fish aggregating devices (FADs) were investigated using ultrasonic telemetry in the equatorial eastern Pacific Ocean. Experiments were conducted during a research cruise aboard a chartered purse seine vessel. Purse seine sets were made on the tuna aggregations associated with FADs at the termination of six of the ten experiments. Seventeen of the 44 tagged tunas were not recaptured indicating the transient nature of the associative behavior of tunas with FADs. Although there was considerable overlap in the depths of the three species, by day and night, there were some species-specific differences and diel differences within species. While we documented spatial and temporal differences in the schooling behavior of the three tuna species, the differences do not appear sufficient such that modifications in purse seine fishing practices could effectively avoid the capture of small bigeye and yellowfin tunas, while optimizing the capture of skipjack tuna in purse seine sets on FADs.     

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.     

Movements of albacore tuna (Thunnus alalunga) in the South Pacific: Evidence from parasites

Parasites were collected from over 400 albacore (Thunnus alalunga) caught by surface trolling and longlining in the south-west Pacific between 1985 and 1988. Parasites found included 1 apicomplexan, 3 nematode species, 4 cestode species, 1 acanthocephalan, 12 digenean species and 3 copepod species. Twelve of these parasite species which could be accurately recognised and counted were used in the subsequent analyses. Parasite data from albacore caught around New Zealand show a decrease in prevalence of three didymozoid parasites with increasing fish length up to a fork length of 70 to 79 cm. The subsequent increase in prevalence of these didymozoids in large longline-caught fish is consistent with fish returning from spawning in tropical waters where re-infection is presumed to occur. Albacore collected at widely separate locations in the south-west Pacific have differences in parasite prevalence, supporting an hypothesis that juvenile albacore move south to New Zealand from the tropics and do not return until the onset of sexual maturity. Albacore appear to move along the subtropical convergence zone, as indicated by a decline in prevalence and abundance ofAnisakis simplex andHepatoxylon trichiuri from New Zealand to the central South Pacific. This is supported by tagging and seasonal movements of the fishery.     

Vertical movement and habitat of opah (Lampris guttatus) in the central North Pacific recorded with pop-up archival tags

Data from 11 pop-up archival transmitting tags attached to opah (Lampris guttatus, F. Lampridae) in the central North Pacific between November 2003 and March 2005 were used to describe their vertical movement and habitat. In the subtropical gyre northwest of the Hawaiian Islands, opah generally inhabited a 50–400 m depth range and 8–22°C temperatures. They were frequently found in depths of 50–150 m at night and in greater depths (100–400 m) during the day, but were constantly moving vertically within this broad range. At night, excursions below 200 m were not uncommon and during the day they were very likely to spend some time at depths <175 m. Their vertical speeds were generally <25 cm s⁻¹ but on one occasion an opah descended at a burst speed of 4 m s⁻¹. Vertical habitat use by individual opah apparently varied with local oceanographic conditions, but over a 24-h period the average temperature experienced was always in the narrow range of 14.7 to 16.5°C.

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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.     

Movements of bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) in the northwestern Atlantic Ocean recorded by pop-up satellite archival tags

Pop-up satellite archival tags were implanted into 68 Atlantic bluefin tuna (Thunnus thynnus Linnaeus), ranging in size from 91 to 295 kg, in the southern Gulf of Maine (n=67) and off the coast of North Carolina (n=1) between July 2002 and January 2003. Individuals tagged in the Gulf of Maine left that area in late fall and overwintered in northern shelf waters, off the coasts of Virginia and North Carolina, or in offshore waters of the northwestern Atlantic Ocean. In spring, the fish moved either northwards towards the Gulf of Maine or offshore. None of the fish crossed the 45°W management line (separating eastern and western management units) and none traveled towards the Gulf of Mexico or the Straits of Florida (known western Atlantic spawning grounds). The greatest depth recorded was 672 m and the fish experienced temperatures ranging from 3.4 to 28.7°C. Swimming depth was significantly correlated with location, season, size class, time of day, and moon phase. There was also evidence of synchronous vertical behavior and changes in depth distribution in relation to oceanographic features.Communicated by J.P. Grassle, New Brunswick     

Distribution and structure of heterotrophic protist communities in the northeast equatorial Pacific Ocean

The distribution and structure of heterotrophic protist communities and size-fractionated chlorophyll a were studied during the Korea Deep Ocean Study 98 (KODOS 98) research expedition (July 1998) in the northeast equatorial Pacific Ocean (5–11°N). Areas of convergence and divergence formed at the boundaries of the South Equatorial Current (SEC), North Equatorial Current (NEC), and North Equatorial Counter Current (NECC) during the expedition. Water column physicochemical characteristics significantly influenced the size structure of heterotrophic protist communities. Intense vertical mixing and high nutrient and chlorophyll a concentrations characterized SEC and NECC areas, which were affected by converging and diverging water masses, respectively. Nanophytoplankton dominated in SEC and NECC areas; both areas also had relatively high heterotrophic protist biomasses (average 743 µg C m^–2). NEC areas were characterized by a stratified vertical structure, low nutrient and chlorophyll a concentrations, and picophytoplankton dominance. The heterotrophic protist biomass in NEC areas averaged 414 µg C m^–2; nanoprotists (<20 µm) dominated the community. The nanoprotist biomass comprised 49–54% of the total heterotrophic protist biomass in SEC/NECC areas and 67–72% in NEC areas. The biomass of heterotrophic protists was higher in SEC/NECC areas than in NEC areas, but the relative importance of nanoprotists was greater in NEC areas than in SEC/NECC areas. Heterotrophic dinoflagellates were dominant components of the <20 µm and >20 µm size classes in both water columns. The biomass of heterotrophic protists significantly correlated with the net-, nano-, and picophytoplankton biomass in SEC/NECC areas and with the nano- and picophytoplankton biomass in NEC areas. Heterotrophic protists and phytoplankton also showed strong positive correlation in the study area. The size structure of the phytoplankton biomass coincided with that of heterotrophic protists; the heterotrophic protist biomass positively correlated with the protists prey source. These relationships suggest that the community structure of heterotrophic protists and the microbial food web depended on size classes within the phytoplankton biomass. Microzooplankton grazing and phytoplankton growth rates were higher in SEC/NECC areas than in NEC areas. In contrast, the potential primary production grazed by microzooplankton was relatively high in NEC areas (127.3%) compared with SEC/NECC areas (94.6%). Our results indicate that the relative importance and size structure of heterotrophic protists might vary according to two distinct water column structures.Communicated by T. Ikeda, Hakodate     

Behavior of bigeye (<Emphasis Type="Italic">Thunnus obesus</Emphasis>) and skipjack (<Emphasis Type="Italic">Katsuwonus pelamis</Emphasis>) tunas within aggregations associated with floating objects in the equatorial eastern Pacific

The horizontal and vertical movements of bigeye (Thunnus obesus Lowe, 1839) and skipjack (Katsuwonus pelamis Linnaeus, 1758) tunas within large multi-species aggregations associated with moored buoys or a drifting vessel were investigated, using ultrasonic telemetry and archival tags, along with sonar imaging, in the equatorial eastern Pacific Ocean (at 2°S–95°W and 2°N–95°W). Four sets of observations, each consisting of the concurrent monitoring of pairs of skipjack and/or bigeye with implanted acoustic or archival tags, were conducted in May 2002 and 2003. Ultrasonic telemetry data were not collected until 24 h or more after the fish were tagged and released, to avoid any abnormal behavior as a consequence of tagging. The pairs of acoustically tagged bigeye and skipjack, and also the entire aggregations, were primarily upcurrent of the moored buoy and downcurrent of the drifting vessel during the day. At night the aggregations were observed to be more diffuse, and the fish were feeding on organisms of the deep scattering layer. The aggregations returned to positions upcurrent of the buoy or downcurrent of the drifting vessel at dawn, commonly breezing at the surface within cohesive monospecific schools. The bigeye and skipjack had concurrent changes in depth records, occupying significantly greater mean depths at night than during the day, in most cases. When associated with a moored buoy, bigeye depth distributions were deeper during the day and night than those of skipjack, but bigeye depth distributions were shallower during the day and night than those of skipjack when associated with the drifting vessel. Simultaneous depth records of a large and a small bigeye with archival tags associated with a moored buoy also indicated diel changes in depth. The mean depth at night was significantly less than during the day for the larger bigeye, but the mean depth during the day was significantly less than during the night for the smaller bigeye. The mean depths during the day and night were significantly greater for the larger bigeye than the smaller.Electronic Supplementary Material  Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-004-1480-xCommunicated by J.P. Grassle, New Brunswick     

Vertical distribution and diurnal migration of some Cyclopoida (Copepoda) in the tropical region of the Pacific Ocean

The material was collected in the western equatorial part of the Pacific Ocean at a 24 h station on December 21 to 23, 1968. Collections were made by means of 2 Juday nets from 2 winches simultaneously. A series of hauls was made at 2 h intervals. The vertical distribution of 17 Cyclopoida species, belonging to 3 families (Oithonidae, Oncaeidae and Corycaeidae) was studied. From 0 to 300 m, the absolute number of Cyclopoida individuals was practically constant throughout the 24 h period. The bulk of the species performed no diurnal vertical migration, or migrated with low intensity in the usual way (i.e. moved upward at night and downward during the day). Only 2 Oncaeidae species performed significant diuranl vertical migration. No reversed migration in Cyclopoida was discovered. To characterize the vertical distribution of Cyclopoida, the distribution of the cores of populations (abundancies between 25 and 75%) was examined. During the 24 h period, the cores of populations of various Cyclopoida species were found at different depths; this steplike distribution was not disturbed in migrating species. The cores of populations differed both with habitat depth and time of ascent of the migrating species to the surface layer. This peculiarity, probably, tends to lessen the intensity of food competition between Cyclopoida species with similar nutritional habits.     

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.     

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.     

Near real time satellite tracking of striped marlin (Kajikia audax) movements in the Pacific Ocean

High-resolution satellite locations were obtained from striped marlin using Argos transmitters attached to the upper lobe of the caudal fin. Twenty-six striped marlin were tagged off New Zealand (2005–2007) and tracked as far as the central Pacific Ocean. Caudal fin mounted Argos tags generated 1,524 locations during a total of 659 tracking days [mean 25 (±21.24) days per fish and 2.3 (±2.30) locations per day]. 38% of locations have an estimated accuracy of ±1 km or better. Displacement rates from high quality locations ranged from 2.9 to 170.8 km in a 24 h period, with a mode at 20–30 km and a mean of 45 km/day. The caudal fin attachment methodology and antenna configuration was adjusted each season to improve transmission life and data quality, with the best results obtained in the last year of deployments (2007). The longest track duration was 102 days, with a total displacement of 4,959 km and a total track distance from all locations received of 6,850 km. Tag shedding and antenna failure appear to have limited the duration of tracks from SPOT tags. The high temporal and spatial resolution data revealed behaviours not previously observed in striped marlin, including associations to subsurface bathymetric features. High resolution location data such as these are useful inputs for statistical models used to investigate habitat selection and switching between different behavioural modes. The geolocations calculated using ukfsst estimates from PAT tag data had RMS errors of 1.01° latitude and 0.59° longitude when compared with SPOT tag Argos locations.     

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).