Annual estimates of the unobserved incidental kill of pantropical spotted dolphin (Stenella attenuata attenuata) calves in the tuna purse-seine fishery of the eastern tropical Pacific

We estimated the total number of pantropical spotted dolphin (Stenella attenuata) mothers killed without their calves (“calf deficit”) in all tuna purse-seine sets from 1973– 90 and 1996–2000 in the eastern tropical Pacific. Estimates were based on a tally of the mothers killed as reported by color pattern and gender, several color-pattern-based frequency tables, and a weaning model. Over the time series, there was a decrease in the calf deficit from approximately 2800 for the western-southern stock and 5000 in the northeastern stock to about 60 missing calves per year. The mean deficit per set decreased from approximately 1.5 missing calves per set in the mid-1970s to 0.01 per set in the late-1990s. Over the time series examined, from 75% to 95% of the lactating females killed were killed without a calf. Under the assumption that these orphaned calves did not survive without their mothers, this calf deficit represents an approximately 14% increase in the reported kill of calves, which is relatively constant across the years examined. Because the calf deficit as we have defined it is based on the kill of mothers, the total number of missing calves that we estimate is potentially an underestimate of the actual number killed. Further research on the mechanism by which separation of mother and calf occurs is required to obtain better estimates of the unobserved kill of dolphin calves in this fishery.

Testicular development in migrant and spawning bluefin tuna (Thunnus thynnus (L.)) from the eastern Atlantic and Mediterranean

Testis histological structure was studied in bluefin tuna (Thunnus thynnus) from the eastern Atlantic and Mediterranean during the reproductive season (from late April to early June). Testicular maturation was investigated by comparing samples from bluefin tuna caught on their eastward reproductive migration off Barbate (Strait of Gibraltar area) with samples of bluefin tuna fished in spawning grounds around the Balearic Islands. Histological evaluations of cross sections showed that the testis consists of two structurally different regions, an outer proliferative region where germ cells develop synchronously in cysts, and a central region made up of a well-developed system of ducts that convey the spermatozoa produced in the proliferative region to the main sperm duct. Ultrastructural features of the different stages of the male germ cell line are very similar to those described in other teleost species. The bluefin tuna testis is of the unrestricted spermatogonial testicular type, where primary spermatogonia are present all along the germinative portion of the lobules. All stages of spermatogenesis were present in the gonad tissue of migrant and spawning bluefin tuna, although spermatids were more abundant in spawning fish. The testis size was found to increase by a factor of four (on average) during migration to the Mediterranean spawning grounds, whereas the fat bodies (mesenteric lipid stores associated with the gonads) became reduced to half their weight, and the liver mass did not change significantly with sexual maturation. Linear regression analysis of the pooled data of migrant and spawning bluefin tuna revealed a significant negative correlation between the gonad index (IG) and the fat tissue index (IF), and a weaker positive correlation between the gonad index (IG) and the liver index (IL). Our analyses indicate that the liver does not play a significant role in the storage of lipids and that mesenteric lipid reserves constitute an important energy source for gametogenesis in bluefin tuna.

Estimating long-term growth-rate changes of southern bluefin tuna (Thunnus maccoyii) from two periods of tag-return data

Southern bluefin tuna (SBT) (Thunnus maccoyii) growth rates are estimated from tag-return data associated with two time periods, the 1960s and 1980s. The traditional von Bertalanffy growth model (VBG) and a two-phase VBG model were fitted to the data by maximum likelihood. The traditional VBG model did not provide an adequate representation of growth in SBT, and the two-phase VBG yielded a significantly better fit. The results indicated that significant change occurs in the pattern of growth in relation to a VBG curve during the juvenile stages of the SBT life cycle, which may be related to the transition from a tightly schooling fish that spends substantial time in near and surface shore waters to one that is found primarily in more offshore and deeper waters. The results suggest that more complex growth models should be considered for other tunas and for other species that show a marked change in habitat use with age. The likelihood surface for the two-phase VBG model was found to be bimodal and some implications of this are investigated. Significant and substantial differences were found in the growth for fish spawned in the 1960s and in the 1980s, such that after age four there is a difference of about one year in the expected age of a fish of similar length which persists over the size range for which meaningful recapture data are available. This difference may be a density-dependent response as a consequence of the marked reduction in the SBT population. Given the key role that estimates of growth have in most stock assessments, the results indicate that there is a need both for the regular monitoring of growth rates and for provisions for changes in growth over time (possibly related to changes in abundance) in the stock assessment models used for SBT and other species.

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.

Migration patterns of young Pacific bluefin tuna (Thunnus orientalis) determined with archival tags

We investigated the migration and behavior of young Pacific bluefin tuna (Thunnus orientalis) using archival tags that measure environmental variables, record them in memory, and estimate daily geographical locations using measured light levels. Swimming depth, ambient water temperature, and feeding are described in a companion paper. Errors of the tag location estimates that could be checked were –0.54° ±0.75° (mean ±SD) in longitude and –0.12° ±3.06° in latitude. Latitude, estimated automatically by the tag, was problematic, but latitude, estimated by comparing recorded sea-surface temperatures with a map of sea-surface temperature, was satisfactory. We concluded that the archival tag is a reliable tool for estimating location on a scale of about one degree, which is sufficient for a bluefin tuna migration study. After release, tagged fish showed a normal swimming behavioral pattern within one day and normal feeding frequency within one month. In addition, fish with an archival tag maintained weight-at-length similar to that of wild fish; however, their growth rate was less than that of wild fish. Of 166 fish released in the East China Sea with implanted archival tags, 30 were recovered, including one that migrated across the Pacific Ocean. Migration of young Pacific bluefin tuna appears to consist of two phases: a residency phase comprising more than 80% of all days, and a traveling phase. An individual young Pacific bluefin tuna was observed to cover 7600 km in one traveling phase that lasted more than two months (part of this phase was a trans-Pacific migration completed within two months). Many features of behavior in the traveling phase were similar to those in the residency phase; however the temperature difference between viscera and ambient temperature was larger, feeding was slightly more frequent, and dives to deeper water were more frequent.

Swimming depth, ambient water temperature preference, and feeding frequency of young Pacific bluefin tuna (Thunnus orientalis) determined with archival tags

We investigated the migration and behavior of young Pacific Bluefin tuna (Thunnus orientalis) using archival tags. The archival tag measures environmental variables, records them in its memory, and estimates daily geographical locations based on measured light levels. Of 166 archival tags implanted in Pacific bluefin tuna that were released at the northeastern end of the East China Sea from 1995 to 1997, 30 tags were recovered, including one from a fish that migrated across the Pacific. This article describes swimming depth, ambient water temperature, and feeding frequency of young Pacific bluefin tuna based on retrieved data. Tag performance, effect of the tag on the fish, and horizontal movements of the species are described in another paper. Young Pacific bluefin tuna swim mainly in the mixed layer, usually near the sea surface, and swim in deeper water in daytime than at nighttime. They also exhibit a pattern of depth changes, corresponding to sunrise and sunset, apparently to avoid a specific low light level. The archival tags recorded temperature changes in viscera that appear to be caused by feeding, and those changes indicate that young Pacific bluefin tuna commonly feed at dawn and in the daytime, but rarely at dusk or at night. Water temperature restricts their distribution, as indicated by changes in their vertical distribution with the seasonal change in depth of the thermocline and by the fact that their horizontal distribution is in most cases confined to water in the temperature range of 14−20°C.

The occurrence of yellowfin tuna (Thunnus albacares) at Espiritu Santo Seamount in the Gulf of Mexico) at Espiritu Santo Seamount in the Gulf of Mexico

Pelagic fishes are not evenly dispersed in the oceans, but aggregate at distinct locations in this vast and open environment. Nomadic species such as mackerels, tunas, and sharks form assemblages at seamounts (Klimley and Butler, 1988; Fontenau, 1991). Fishermen have recognized this behavior and have placed moorings with surface buoys in deep waters to provide artificial landmarks, around which fish concentrate and are more easily captured. These fish aggregating devices (termed FADs) are common in the tropical oceans (see review, Holland, 1996). In a sense, it may only be the larger size that separates a seamount from a man-made FAD.

Allozyme and RAPD variation in the eastern Pacific yellowfin tuna (Thunnus albacares)

Stock structure of eastern Pacific yellowfin tuna was investigated by analyzing allozymes and random amplified polymorphic DNAs (RAPDs) from 10 samples of 20–30 individuals each, collected between 1994 and 1996 from fishing vessels operating in the Inter-American Tropical Tuna Commission (IATTC) yellowfin regulatory area (CYRA). Allozyme analysis resolved 28 loci, eight of which were polymorphic under the 0.95 criterion: Aat-S*, Glud, Gpi-F*, Gpi-S*, La, Lgg, Pap-F*, and 6-Pgd, resulting in a mean heterozygosity over all allozyme loci of H = 0.052. Four polymorphic RAPD loci were selected for analysis, resulting in a mean heterozygosity of H = 0.43. Eight of 45 pairwise comparisons of allozyme allele frequencies among the ten samples showed significant differences after correction for multiple testing (P<0.0001), all of which involved comparisons with the Gulf of California sample. Confirmation of this signal of population structure would have management implications. No significant divergence in RAPD allele frequencies was observed among samples. Weir and Cockerham θ estimated for allozyme loci (θ=0.048; P<0.05) and RAPD loci (θ=0.030; P>0.05) revealed little population structure among samples. Mantel tests demonstrated that the genetic relationships among samples did not correspond to an isolation-by-distance model for either class of marker. Four of eight comparisons of coastal and offshore samples revealed differences of allele frequencies at the Gpi-F* locus (P<0.05), although none of these differences was significant after correction for multiple testing (P>0.001). Results are consistent with the hypothesis that the CYRA yellowfin tuna samples comprise a single genetic stock, although gene flow appears to be greater among coastal samples than between coastal and offshore samples.

Bycatch in the tuna purse-seine fisheries of the western Indian Ocean

Bycatch taken by the tuna purse-seine fishery from the Indian Ocean pelagic ecosystem was estimated from data collected by scientific observers aboard Soviet purse seiners in the western Indian Ocean (WIO) during 1986–92. A total of 494 sets on free-swimming schools, whale-shark-associated schools, whale-associated schools, and log-associated schools were analyzed. More than 40 fish species and other marine animals were recorded. Among them only two species, yellow-fin and skipjack tunas, were target species. Average levels of bycatch were 0.518 metric tons (t) per set, and 27.1 t per 1000 t of target species. The total annual purse-seine catch of yellowfin and skipjack tunas by principal fishing nations in the WIO during 1985–94 was 118,000–277,000 t. Nonrecorded annual bycatch for this period was estimated at 944–2270 t of pelagic oceanic sharks, 720–1877 t of rainbow runners, 705–1836 t of dolphinfishes, 507–1322 t of triggerfishes, 113–294 t of wahoo, 104–251 t of billfishes, 53–112 t of mobulas and mantas, 35–89 t of mackerel scad, 9–24 t of barracudas, and 67–174 t of other fishes. In addition, turtle bycatch and whale mortalities may have occurred. Because the bycatches were not recorded by some purse-seine vessels, it was not possible to assess the full impact of the fisheries on the pelagic ecosystem of the Indian Ocean. The first step to solving this problem is for the Indian Ocean Tuna Commission to establish a pro-gram in which scientific observers are placed on board tuna purse-seine and longline vessels fishing in the WIO.

Horizontal and vertical movements of juvenile bluefin tuna (Thunnus thynnus) in relation to oceanographic conditions of the western North Atlantic, determined with ultrasonic telemetry

We employed ultrasonic transmitters to follow (for up to 48 h) the horizontal and vertical movements of five juvenile (6.8–18.7 kg estimated body mass) bluefin tuna (Thunnus thynnus) in the western North Atlantic (off the eastern shore of Virginia). Our objective was to document the fishes’ behavior and distribution in relation to oceanographic conditions and thus begin to address issues that currently limit population assessments based on aerial surveys. Estimation of the trends in adult and juvenile Atlantic bluefin tuna abundance by aerial surveys, and other fishery-independent measures, is considered a priority. Juvenile bluefin tuna spent the majority of their time over the continental shelf in relatively shallow water (generally less then 40 m deep). Fish used the entire water column in spite of relatively steep vertical thermal gradients (≈24°C at the surface and ≈12°C at 40 m depth), but spent the majority of their time (≈90%) above 15 m and in water warmer then 20°C. Mean swimming speeds ranged from 2.8 to 3.3 knots, and total distance covered from 152 to 289 km (82–156 nmi). Because fish generally remained within relatively con-fined areas, net displacement was only 7.7–52.7 km (4.1–28.4 nmi). Horizontal movements were not correlated with sea surface temperature. We propose that it is unlikely that juvenile bluefin tuna in this area can detect minor horizontal temperature gradients (generally less then 0.5°C/km) because of the steep vertical temperature gradients (up to ≈0.6°C/m) they experience during their regular vertical movements. In contrast, water clarity did appear to influence behavior because the fish remained in the intermediate water mass between the turbid and phytoplankton-rich plume exiting Chesapeake Bay (and similar coastal waters) and the clear oligotrophic water east of the continental shelf.

Differences in diet of Atlantic bluefin tuna (Thunnus thynnus) at five seasonal feeding grounds on the New England continental shelf

The stomachs of 819 Atlantic bluefin tuna (Thunnus thynnus) sampled from 1988 to 1992 were analyzed to compare dietary differences among five feeding grounds on the New England continental shelf (Jeffreys Ledge, Stellwagen Bank, Cape Cod Bay, Great South Channel, and South of Martha’s Vineyard) where a majority of the U.S. Atlantic commercial catch occurs. Spatial variation in prey was expected to be a primary influence on bluefin tuna distribution during seasonal feeding migrations. Sand lance (Ammodytes spp.), Atlantic herring (Clupea harengus), Atlantic mackerel (Scomber scombrus), squid (Cephalopoda), and bluefish (Pomatomus saltatrix) were the top prey in terms of frequency of occurrence and percent prey weight for all areas combined. Prey composition was uncorrelated between study areas, with the exception of a significant association between Stellwagen Bank and Great South Channel, where sand lance and Atlantic herring occurred most frequently. Mean stomach-contents biomass varied significantly for all study areas, except for Great South Channel and Cape Cod Bay. Jeffreys Ledge had the highest mean stomach-contents biomass (2.0 kg) among the four Gulf of Maine areas and Cape Cod Bay had the lowest (0.4 kg). Diet at four of the five areas was dominated by one or two small pelagic prey and several other pelagic prey made minor contributions. In contrast, half of the prey species found in the Cape Cod Bay diet were demersal species, including the frequent occurrence of the sessile fig sponge (Suberites ficus). Prey size selection was consistent over a wide range of bluefin length. Age 2–4 sand lance and Atlantic herring and age 0–1 squid and Atlantic mackerel were common prey for all sizes of bluefin tuna. This is the first study to compare diet composition of western Atlantic bluefin tuna among discrete feeding grounds during their seasonal migration to the New England continental shelf and to evaluate predator-prey size relationships. Previous studies have not found a common occurrence of demersal species or a pre-dominance of Atlantic herring in the diet of bluefin tuna.

First record of a yellowfin tuna (Thunnus albacares) from the stomach of a longnose lancetfish (Alepisaurus ferox)

In 1987 we found a juvenile yellowfin tuna, Thunnus albacares (Bonnaterre, 1788), in the stomach of a longnose lancetfish, Alepisaurus ferox Lowe, 1833. Analysis of published information on lancetfish food habits (Haedrich, 1964, 1969; Haedrich and Nielsen, 1966; Parin, 1968; Parin et al., 1969; Fourmanoir, 1969; Grandperrin and Legand, 1970; Kubota and Uyeno, 1970; Legand et al., 1972; Kubota, 1973; Fujita and Hattori, 1976; Matthews et al., 1977) led us to conclude that this was the first record of a yellowfin tuna found in a lancetfish stomach.

Vertical and horizontal movements of southern bluefin tuna (Thunnus maccoyii) in the Great Australian Bight observed with ultrasonic telemetry

The vertical and horizontal movements of southern bluefin tuna (SBT), Thunnus maccoyii, in the Great Australian Bight were investigated by ultrasonic telemetry. Between 1992 and 1994, sixteen tuna were tracked for up to 49 h with depth or combined temperature-depth transmitting tags. The average swimming speeds (measured over the ground) over entire tracks ranged from 0.5 to 1.4 m/s or 0.5 to 1.4 body lengths/s. The highest sustained swimming speed recorded was 2.5 m/s for 18 hours. Horizontal movements were often associated with topographical features such as lumps, reefs, islands and the shelf break. They spent long periods of time at the surface during the day (nearly 30%), which would facilitate abundance estimation by aerial survey. At night, they tended to remain just below the surface, but many remained in the upper 10 m throughout the night. SBT were often observed at the thermocline interface or at the surface while travelling. A characteristic feature of many tracks was sudden dives before dawn and after sunset during twilight, followed by a gradual return to their original depth. It is suggested that this is a behavior evolved to locate the scattering layer and its associated prey when SBT are in waters of sufficient depth. SBT maintained a difference between stomach and ambient temperature of up to 9°C.

Bycatch of billfishes by the European tuna purse-seine fishery in the Atlantic Ocean

Billfishes are a component of offshore ecosystems; thus it is important to quantify the impact of the tuna fishery on these species in the world’s ocean. The aim of this study was to assess the bycatch of billfishes generated by the tropical tuna purse-seine fishery in the eastern Atlantic Ocean. Information on bycatch was collected by observers at sea during the European Union Bigeye Program. With a total of 62 observers’ trips, conducted on Spanish and French vessels between June 1997 and May 1999, this project is the biggest observer program ever carried out in the European tuna purse-seine fishery. This study showed that billfish bycatch by the purse seiners is very low (less than 0.021% of the total tuna catches and less than 10% of the total billfish catches currently reported). A Monte Carlo simulation was performed to account for some uncertainties in the fishing strategies of purse seiners operating in this ocean. One of the findings of this study indicated that the temporary moratorium on fishing with FADs (fish aggregating devices), adopted by the European purse-seine fishery in the eastern Atlantic Ocean, produced a decrease in incidental catches of marlins from 600–700 metric tons (t) to less than 300 t. In contrast, this trend was reversed for sailfishes, for which the bycatch increased from 25 t to 45 t. The difficulty of defining indices that express the conservation status in marine fishes and that gauge key ecosystem parameters and the need to promote an ecosystem approach for large-pelagic-resource management which takes into account biologic and socioeconomic criteria are briefly discussed.

Movements, behavior, and habitat selection of bigeye tuna (Thunnus obesus) in the eastern equatorial Pacific, ascertained through archival tags

Ninety-six bigeye tuna (88– 134 cm fork length) were caught and released with implanted archival (electronic data storage) tags near fish-aggregating devices (FADs) in the equatorial eastern Pacific Ocean (EPO) during April 2000. Twenty-nine fish were recaptured, and the data from twenty-seven tags were successfully downloaded and processed. Time at liberty ranged from 8 to 446 days, and data for 23 fish at liberty for 30 days or more are presented. The accuracy in geolocation estimates, derived from the light level data, is about 2 degrees in latitude and 0.5 degrees in longitude in this region. The movement paths derived from the filtered geolocation estimates indicated that none of the fish traveled west of 110°W during the period between release and recapture. The null hypothesis that the movement path is random was rejected in 17 of the 22 statistical tests of the observed movement paths. The estimated mean velocity was 117 km/d. The fish exhibited occasional deep-diving behavior, and some dives exceeded 1000 m where temperatures were less than 3°C. Evaluations of timed depth records, resulted in the discrimination of three distinct behaviors: 54.3% of all days were classified as unassociated (with a floating object) type-1 behavior, 27.7% as unassociated type-2 behavior, and 18.7% as behavior associated with a floating object. The mean residence time at floating objects was 3.1 d. Data sets separated into day and night were used to evaluate diel differences in behavior and habitat selection. When the fish were exhibiting unassociated type-1 behavior (diel vertical migrations), they were mostly at depths of less than 50 m (within the mixed layer) throughout the night, and during the day between 200 and 300 m and 13° and 14°C. They shifted their average depths in conjunction with dawn and dusk events, presumably tracking the deep-scattering layer as a foraging strategy. There were also observed changes in the average nighttime depth distributions of the fish in relation to moon phase.