Genetic structure of Patagonian toothfish (Dissostichus eleginoides) populations on the Patagonian Shelf and Atlantic and western Indian Ocean Sectors of the Southern Ocean

The genetic structure of Patagonian toothfish populations in the Atlantic and western Indian Ocean Sectors of the Southern Ocean (SO) were analysed using partial sequences of the mitochondrial 12S rRNA gene and seven microsatellite loci. Both haplotype frequency data (F _ST>0.906, P<0.01) and microsatellite genotype frequency data (F _ST=0.0141–0.0338, P<0.05) indicated that populations of toothfish from around the Falkland Islands were genetically distinct from those at South Georgia (eastern Atlantic Sector SO), around Bouvet Island (western Atlantic Sector SO) and the Ob Seamount (western Indian Ocean Sector of the SO). Genetic differentiation between these populations is thought to result from hydrographic isolation, as the sites are separated by two, full-depth, ocean-fronts and topographic isolation, as samples are separated by deep water. The South Georgia, Bouvet and Ob Seamount samples were characterised by an identical haplotype. However, microsatellite genotype frequencies showed genetic differentiation between South Georgia samples and those obtained from around Bouvet Island and nearby seamounts (F _ST=0.0037, P<0.05). These areas are separated by large geographic distance and water in excess of 3,000 m deep, below the distributional range of toothfish (<2,200 m). No significant genetic differentiation was detected between samples around Bouvet Island and the Ob Seamount although comparisons may have been influenced by low sample size. These localities are linked by topographic features, including both ridges and seamounts, that may act as oceanic “stepping stones” for migration between these populations. As for other species of deep-sea fish, Patagonian toothfish populations are genetically structured at the regional and sub-regional scales.     

Phylogeography of the Indo-Pacific parrotfish Scarus psittacus: isolation generates distinctive peripheral populations in two oceans

Phylogenetic, phylogeographic, population genetic and coalescence analyses were combined to examine the recent evolutionary history of the widespread Indo-Pacific parrotfish, Scarus psittacus, over a geographic range spanning three marine biogeographic realms. We sequenced 164 individuals from 12 locations spanning 17,000 km, from 55oE to 143oW, using 322 base pairs of mitochondrial control region (D-loop). S. psittacus displayed high haplotype (h = 0.83–0.98), but low nucleotide (<1%) diversity. Most (>83%) genetic variation was within populations. AMOVA revealed significant partitioning and identified five geographic groups. These included one central population and four populations peripheral to the centre. The central population occupied reefs from Western Australia to Tahiti and represented the central Indo-Pacific biogeographic realm. Cocos Keeling was distinct from central and western Indo-Pacific biogeographic realms occupying a position intermediate to these. Peripheral populations (Hawaii, Marquesas) represented the eastern Indo-Pacific biogeographic realm, while Seychelles represented the western Indo-Pacific biogeographic realm. All but the central population expanded (<163 kya). Whilst all populations experienced major sea level and SST changes associated with Pleistocene glaciation cycles, the genetic structure of the central population was relatively homogenous unlike the remaining genetically distinctive populations.     

Biogeographic patterns on shallow subtidal reefs in the western Indian Ocean

To resolve biogeographic limits and patterns on the east coast of Africa, the presence/absence and quantitative biomass data were collected from 55 shallow subtidal reefs along 4,800 km of coastline (5.2°–31.1°S). Multivariate analysis of distributions, trophic structure and biomass revealed two distinct marine provinces, the Tropical Indo-West Pacific and Subtropical Natal, with a transitional overlap (of ca. 120 km) located between Leven Point and Kosi Mouth on north-east South Africa. This region of overlap was one of three bioregions revealed by post hoc analyses. Biomass was unexpectedly highest in the tropics. ‘Auto-heterotrophs’ (species with autotrophic symbionts such as corals and clams), deposit feeders and grazers contributed significantly more biomass to the Tropical Indo-West Pacific Bioregion, whereas filter feeders dominated the Subtropical Natal Bioregion. ‘Auto-heterotrophs’ declined with latitude while filter feeders increased; soft corals made a defining contribution in the overlap bioregion. Possible underlying causes of these patterns include productivity, nutrient levels, riverine input, light penetration and temperature.     

Isotopic evidence of distinct feeding ecologies and movement patterns in two migratory predators (yellowfin tuna and swordfish) of the western Indian Ocean

Ecologists primarily use δ¹⁵N values to estimate the trophic level of organisms, while δ¹³C, and even recently δ¹⁵N, are utilized to delineate feeding habitats. However, many factors can influence the stable isotopic composition of consumers, e.g. age, starvation or isotopic signature of primary producers. Such sources of variability make the interpretation of stable isotope data rather complex. To examine these potential sources of variability, muscle tissues of yellowfin tuna (Thunnus albacares) and swordfish (Xiphias gladius) of various body lengths were sampled between 2001 and 2004 in the western Indian Ocean during different seasons and along a latitudinal gradient (23°S to 5°N). Body length and latitude effects on δ¹⁵N and δ¹³C were investigated using linear models. Both latitude and body length significantly affect the stable isotope values of the studied species but variations were much more pronounced for δ¹⁵N. We explain the latitudinal effect by differences in nitrogen dynamics existing at the base of the food web and propagating along the food chain up to top predators. This spatial pattern suggests that yellowfin and swordfish populations exhibit a relatively unexpected resident behaviour at the temporal scale of their muscle tissue turnover. The body length effect is significant for both species but this effect is more pronounced in swordfish as a consequence of their different feeding strategies, reflecting specific physiological abilities. Swordfish adults are able to reach very deep water and have access to a larger size range of prey than yellowfin tuna. In contrast, yellowfin juveniles and adults spend most of their time in the surface waters and large yellowfin tuna continue to prey on small organisms. Consequently, nitrogen isotopic signatures of swordfish tissues are higher than those of yellowfin tuna and provide evidence for different trophic levels between these species. Thus, in contrast to δ¹³C, δ¹⁵N analyses of tropical Indian Ocean marine predators allow the investigation of complex vertical and spatial segregation, both within and between species, even in the case of highly opportunistic feeding behaviours. The linear models developed in this study allow us to make predictions of δ¹⁵N values and to correct for any body length or latitude differences in future food web studies.     

The occurrence of Oratosquilla investigatoris (Crustacea: Stomatopoda) in the pelagic zone of the Gulf of Aden and the equatorial western Indian Ocean

Surface swarms of the stomatopod Oratosquilla investigatoris (Lloyd, 1907) were observed in the Gulf of Aden and the equatorial western Indian Ocean during 1967. Collections from inshore and offshore swarms were made in both areas, and from demersal trawl catches in the equatorial area. O. investigatoris was also collected from the stomachs of several species of oceanic pelagic and inshore fish from 1965 to 1967, and was also reported in the stomach contents of the lesser frigate bird. Further collections were made from strandings on two oceanic islands in the equatorial area. Indirect observations on the occurrence of the species resulted from increased predation on pelagic longline baits during seasos of apparent abundance of O. investigatoris in the equatorial region. At the surface, predation by the species was observed on fast-swiming fish, and the swimming behaviour was noted. A size range of 6 to 37 mm carapace length (CL) was recorded for the 4000 specimens collected, and there was no size difference between the sexes. Length-frequency analysis of the samples indicated a major mode at 19 to 21 mm CL and an increase in length of 3 mm over 3 months. The sex ratio was weighted in favour of females in the majority of collections. It was concluded that the pelagic swarming of O. investigatoris is an important ecological phenomenon, but not regular in the area, and the factors affecting it could not be deduced from the available data.     

Population structure of the black tiger prawn, Penaeus monodon, among western Indian Ocean and western Pacific populations

We examined the population structure of the black tiger prawn, Penaeus monodon Fabricius, 1798, in the Indo-West Pacific by analyzing the geographic distribution of elongation factor 1-alpha intron sequences from specimens collected during the winter and spring of 1997. Both the molecular phylogeny of alleles and F-statistics indicated very strong differentiation between populations from the western Indian Ocean and western Pacific. This pattern is concordant with other recent studies of marine species in this region, implying that the Indo-Australian Archipelago represents a biogeographic break between populations in the Indo-West Pacific. F _ST-values among populations in the western Indian Ocean also indicate structure within this region, whereas no structure was found among western Pacific populations. Nucleotide diversity was significantly lower in the western Indian Ocean populations than in the western Pacific, implying that the populations have regional differences in demographic history. Received: 16 November 1998 / Accepted: 26 May 1999     

Community interactions and zoogeography of the Indian Ocean Candaciidae (Copepoda: Calanoida)

Eighteen species of Candaciidae have been identified from collections made at 339 stations in the Indian Ocean. Most species are zonally distributed; however, on the eastern and western sides of the Indian Ocean, species ranges are extended north or south by boundary currents. Factor analysis was used to cluster phenotypically similar species based on 130 characters taken from the maxilla (a feeding appendage), the first swimming foot, and the fifth foot (a secondary sexual structure). Four morphological clusters were extracted. Clusterings based on separate factor analyses limited to characters from feeding or from sexually adapted appendages are in substantial agreement with the clusters based on composite morphology. Two geographically recurrent groups of species were also identified, one from equatorial waters, one from the central gyre. Each recurrent group is composed of species exceptionally different in body size and belonging to different morphological clusters. Niches of the species are compared using Hutchinson's multidimensional hypervolume as a model. An attempt is made to describe the niche of each species in terms of environmental variables measured at stations where the species was abundant. Environmental variables (dimensions) included in this study are space, time, temperature, food, oxygen and salinity. Mean niche adaptations of most of the species are separable from congeners along at least one niche dimension. It is proposed that community relationships among the Candaciidae developed to prevent both gamete loss and to avoid trophic competition. It is postulated that newly evolving species underwent contemporaneous displacement of both secondary sexual and trophic characters as a condition for sympatry, or diverged in their physiological adaptations to escape sympatric interference.Contribution No. 3686 of the Woods Hole Oceanographic Institution. This research was supported by NSF Grants GB 27405 and GA 43126 and is based on a doctoral dissertation submitted to the Curriculum in Marine Sciences of the University of North Carolina at Chapel Hill.     

Multi-marker estimate of genetic connectivity of sole (Solea solea) in the North-East Atlantic Ocean

A thorough knowledge on the genetic connectivity of marine populations is important for fisheries management and conservation. Using a dense population sampling design and two types of neutral molecular markers (10 nuclear microsatellite loci and a mtDNA cytochrome b fragment), we inferred the genetic connectivity among the main known spawning grounds of sole (Solea solea L.) in the North-East Atlantic Ocean. The results revealed a clear genetic structure for sole in the North-East Atlantic Ocean with at least three different populations, namely the Kattegat/Skagerrak region, the North Sea and the Bay of Biscay, and with indications for a fourth population, namely the Irish/Celtic Sea. The lack of genetically meaningful differences between biological populations within the southern North Sea is likely due to a large effective population size and sufficient connection (gene flow) between populations. Nevertheless, an isolation-by-distance pattern was found based on microsatellite genotyping, while no such pattern was observed with the cytochrome b marker, indicating an historical pattern prevailing in the latter marker. Our results demonstrate the importance of a combined multi-marker approach to understand the connectivity among marine populations at region scales.     

Notes on the Nerita (Archaeogastropoda) populations of Aldabra Atoll,Indian Ocean

Nerita undata, N. plicata, N. polita, N. albicilla and N. textilis are common intertidal gastropods on Aldabra Atoll. Each species prefers a different level on the shore and/or different degrees of exposure to wave action. Patterns of zonation, population size-frequency structure and biomass are given. N. undata exhibits 2 distinct ecophenotypes. Normal types occur on expsed to moderately sheltered shores but are replaced by type B in extremely sheltered conditions. Differences in shell morphology and population structure are also noted. Foraging by all species was confined to hours of darkness, generally at low tide. Activity of N. polita was induced slightly on overcast days. N. undata and N. textilis are cryptically coloured and often visible during the day. N. plicata is conspicuous, but is protected by a strong shell. N. polita and N. albicilla are often brightly and variably coloured, but both are concealed during daylight. No Nerita were found infected with trematodes, suggesting that the degree of infection was extremely low. Habitats of the Aldabran Nerita are compared with 3 species on Barbados, West Indies.     

Parasites and headless chaetognaths in the Indian Ocean

Typhloscolecid polychaetes, assigned to the genera Typhloscolex (Typhloscolex muelleri) and Travisiopsis (Travisiopsis dubia), feeding on chaetognaths are reported for the first time from the Indian Ocean. These parasitic/predatory polychaetes cause decapitation in the chaetognath Flacisagitta enflata, either by eating the head or by causing injuries to the neck or body. Data from eastern Africa to western India showed a peak of polychaete infestation and decapitation of F. enflata in the Seychelles. A 5% peak occurrence (turnover rate is unknown) of decapitated individuals indicates that this parasitism/predation may have an important impact on local F. enflata populations. The harpacticoid copepod Microsetella norwegica was found feeding on the head and the reproductive organs of F. enflata. Therefore, this copepod may cause both decapitation and castration. Flatworm parasites are reported from the body cavity of F. enflata. Digeneans probably belonging to the genera Aphanurus, Parahemiurus, Accacladocoelium and Opechona, along with the accacoeliid Cercaria owreae and didymozoid metacercariae, were found. Cestode larvae were also recovered. No nematodes were found inside chaetognaths in this study.Communicated by O. Kinne, Oldendorf/Luhe     

Home range and movement patterns of the Redlip Parrotfish (Scarus rubroviolaceus) in Hawaii

The activity and movement of the Redlip Parrotfish, Scarus rubroviolaceus, near Wawaloli reef, Hawaii (19°43′06″N, 156°03′11″W), was investigated with active and passive tracking techniques from April through November 2007. Site fidelity was determined and home range sizes were measured for 21 individuals. All individuals showed strong diurnal activity patterns and relatively stable home ranges. Terminal phase individuals had larger home range areas than initial phase fish, and home range size increased with depth. Occasional long forays from the home range were observed in many individuals. Forays often occurred in crepuscular hours and may represent migrations to/from nocturnal sheltering sites. Findings from this study underscore the importance of depth and nighttime shelter migrations as design considerations for effective marine protected areas (MPA) for large parrotfishes.     

Distribution of marine cladocerans in the Indian Ocean

Cladocera of the Indian Ocean have been studied, on the basis of samples collected during the International Indian Ocean Expedition (IIOE). Out of a total of 1927 samples, 552 contained Cladocera. Although the distribution of this group is mainly coastal, one species, Evadne tergestina, has been found in oceanic waters also, where it may establish seasonal populations. Evadne spinifera was recorded off the Australian and South African coasts in the southern hemisphere, and in the Northeastern part of the Arabian Sea coastal waters in the northern hemisphere. Podon polyphemoides was reported from a single station in South African waters. Penilia avirostris was found in coastal waters throughout the Indian Ocean. Short-term distribution patterns of E. tergestina and P. avirostris are discussed. The possible value of this group as food for pelagic fishes has been indicated. This work makes clear that adequate sampling in space and time is required along the Continental Shelf of the Indian Ocean in order to obtain more comprehensive information on this group.Visiting Scientist at Indian Ocean Biological Centre, Cochin, India under UNESCO contract SC/2077/70-20579.     

The swimming crab Charybdis smithii: distribution, biology and trophic role in the pelagic ecosystem of the western Indian Ocean

Surface “swarms” of the swimming crabs Charybdis smithii are still considered as an unusual phenomenon in the open Indian Ocean, although their dense pelagic aggregations were already reported in waters off the Indian coast and in the northern Arabian Sea. Based on an extensive large-scale data series taken over 45 years, we demonstrate that C. smithii is common in the pelagic provinces of the western Indian Ocean driven by the wind monsoon regime. Swimming crabs are dispersed by the monsoon currents throughout the equatorial Indian Ocean. They aggregate at night in the upper 150-m layer, where their estimated biomass derived from pelagic trawling data can exceed 130 kg km⁻². Abundance of C. smithii can reach >15,000 ind. km⁻² in July (i.e. the peak of the south-west monsoon), declines by 50-fold in March and is negligible in May. C. smithii is an important prey for more than 30 species of abundant epipelagic top predators. In turn, it feeds on mesopelagic species. This swimming crab is a major species of the intermediate trophic levels and represents a crucial seasonal trophic link in the open ocean ecosystem of the western Indian Ocean. Outbursts in pelagic waters of huge biomasses of ordinarily benthic crustaceans (C. smithii and Natosquilla investigatoris) are a remarkable feature of the Indian Ocean, although similar, but smaller, events are reported in the Pacific and Atlantic Oceans.

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Mitochondrial DNA variation in king mackerel (Scomberomorus cavalla) from the western Atlantic Ocean and Gulf of Mexico

King mackerel (Scomberomorus cavalla Cuvier) collected in 1992 and 1993 from 13 localities along the Atlantic coast of the southeastern USA and in the northern Gulf of Mexico were surveyed for variation in mitochondrial (mt)DNA and a nuclear-encoded dipeptidase locus (PEPA-2). Both polymorphic and fixed mtDNA restriction sites were identified and mapped using conventional and polymerase chain-reaction (PCR)-based methods. Heterogeneity in mtDNA haplotype frequencies was found only in comparisons of pooled haplotypes from Atlantic localities versus pooled haplotypes from Gulf localities. This finding indicates weak genetic divergence between king mackerel from the Atlantic and those from the Gulf. Frequencies of two PEPA-2 alleles essentially paralleled previous findings: one allele (PEPA-2a) was common among samples from western Gulf localities, whereas the other allele (PEPA-2b) was common among samples from Atlantic and eastern Gulf localities. There was considerable variation in PEPA-2 allele frequencies within broadly-defined regions. Variation in mtDNA haplotypes and PEPA-2 genotypes was independent, as was variation in mtDNA haplotypes with sex or age of individuals. Variation in PEPA-2 genotypes was not independent of sex or age of individuals. The latter result suggests that frequencies of PEPA-2 alleles in samples of king mackerel may stem, in part, from sex and age distributions of individuals within samples, and indicates that caution should be exercised in using allelic variation at PEPA-2 as a measure of population (stock) structure in king mackerel. The discordance in spatial patterning of mtDNA haplotypes versus PEPA-2 alleles across the Gulf (i.e. homogeneity in mtDNA haplotype frequencies versus heterogeneity in PEPA-2 allele frequencies) may be due to either female excess at several localities, sex-biased migration, or both. Observed patterns of genetic variation also are consistent with the hypothesis that king mackerel in the western Atlantic may have been subdivided during Pleistocene glaciation, and that the current distribution of PEPA-2 alleles may be a historical artefact. Received: 17 December 1996 / Accepted: 2 April 1997     

Phylogeography, historical demography, and the role of post-settlement ecology in two Hawaiian damselfish species

Coral reef fish generally have relatively sedentary juvenile and adult phases and a presumed highly dispersive pelagic larval phase, yet previous studies that have tried to relate pelagic larval duration (PLD) to population structure have given inconsistent results. In the present study, the population structures of two damselfishes, Stegastes fasciolatus and Dascyllus albisella, were examined using mitochondrial control region sequences. The two species have similar PLDs (∼25 and 27 days respectively), but consistently differ in their settlement preferences, habitat, and densities in populations throughout the Hawaiian Archipelago, from Hawaii north to Kure Atoll, and south to Johnston Atoll. Information on habitat preferences and population densities were collected between September 2000 and October 2002, and tissue samples for the genetic studies were collected between January and April 2004. Based on the differences in habitat and abundance of the two species, the expectation was that S. fasciolatus would have high genetic variability but little population structure compared to D. albisella, and this was largely confirmed. Stegastes fasciolatus had little population structure in most of the Hawaiian Islands, and D. albisella showed evidence of strong population structure throughout its range. An exception to this pattern was the large difference between the Kure Atoll population of S. fasciolatus and all others. These results suggest that the interaction of several biological factors (e.g. species-specific spawning habitat and season) with environmental factors (e.g. seasonal wind and current patterns) may have more influence on population structure than single life history characteristics, such as the PLD.     

Social organization and sexual pattern in the Mediterranean parrotfish Sparisoma cretense (Teleostei: Scaridae)

We examined the social organization and reproductive pattern of a population of Sparisoma cretense L. at Lampedusa Island (Italy). During the breeding season (July to September) individuals occur either in territorial or in non-territorial groups, which quantitatively differ with respect to use of space, sex-ratio and sexual activity. Territorial groups consist of one male plus one to three females, whereas in non-territorial groups as many as 54 fish share a common area. Among non-territorial groups, variations, in both sex-ratio and behavior, suggest they could represent either feeding aggregations or reproductive aggregations. Spawning, always in pairs, occurs daily in a short period of time before dusk and has been observed only in territorial groups. Group spawning has never been observed, but another alternative mating tactic, streaking on pair spawning, was recorded. In the non-breeding season fish do not aggregate in organized social units. Histological examination of gonads showed that adults are larger than 12 cm total length. Females have an asynchronous ovary, typical of species spawning several times during the breeding season. Males show secondary testes, indicating that their gonads develop ovaries as juveniles, which are later redifferentiated into functional testes. Histological and demographic data seem to indicate that, as in other species of this genus, prematurational sex-change occurs. The sexual pattern appears to be essentially gonochoristic, but the potential for sex-change is not excluded. Received: 5 January 1999 / Accepted: 22 June 1999     

Genetic distances among fishes of the genus Siganus (Siganidae) from the Western Pacific Ocean

Frequencies of electromorphs encoded by 14 polymorphic loci were used to estimate genetic distances among samples of the rabbitfishes Siganus argenteus, S. doliatus, S. fuscescens, S. guttatus, S. javus, S. punctatus, S. randalli, S. spinus, S. vermiculatus, and S. (Lo) vulpinus. Samples of the surgeonfish Naso lituratus (Acanthuridae) were included in the electrophoretic survey for the purpose of outgroup comparison. A matrix of genetic-distance estimates was used for construction of dendrograms by neighbor-joining, complete-linkage and single-linkage procedures, and the unweighted pair-group method of arithmetic averages (UPGMA). Cluster analysis revealed interspecific relationships that are concordant with several predictions intimated in the most recent taxonomic revision of this assemblage of fishes. S. randalli and S. vermiculatus, the pair postulated to be sibling species on the basis of strikingly similar external morphology, were found to exhibit the lowest levels of genetic divergence relative to all other pairs of study species. S. randalli, S. guttatus, and S. vermiculatus comprised one of three clusters discerned in all resultant topologies (i.e., ball clusters). S. argenteus, S. fuscescens, and S. spinus, three fusiform-bodied species, formed a second ball cluster that was clearly separable from the remainder of the siganids studied (which are all relatively deep-bodied). A third ball cluster contained S. punctatus and S. vulpinus: S. doliatus was depicted as being allied with S. punctatus and S. vulpinus in the neighbor-joining and complete-linkage topologies but not in topologies generated by UPGMA and single linkage. It is noteworthy that S. punctatus, S. vulpinus, and S. doliatus are the only three species included in the present study which purportedly school when young but form pair-bonds as adults. In all topologies, S. javus occupied branches stemming from furcation points between fusiform-bodied and deep-bodied species. Of all the siganids studied, S. argenteus was found to be least divergent genetically from N. lituratus. S. argenteus is the only siganid known to spawn pelagic eggs and undergo a specialized pelagic stage (features typical within the Acanthuridae). Hence, genetic relationships within the Siganidae and among the Siganidae and a representative acanthurid coincide with a host of predictions stemming from morphologic and life-history studies of the Acanthuroidei.     

An oceanographic interpretation of seabird distributions in the Indian Ocean

When sightings of seabirds in the Indian Ocean are analyzed in conjunction with oceanographic data, the distributions of certain species and higher taxa show significant correlation with the temperature and salinity of the surface waters. Closely related species occur preferentially over different types of surface water. Characteristic associations of species were also found corresponding to surface water-types. Seabirds are not exempted by their mobility from the constraints of the marine environment.     

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.     

Genetic variation in the neritic squid Loligo forbesi (Myopsida: Loliginidae) in the northeast Atlantic Ocean

Horizontal starch gel electrophoresis was employed to investigate levels of genetic differentiation between 13 samples of the neritic squid species Loligo forbesi Steenstrup obtained from throughout the majority of its known geographical range. Six enzyme loci identified in a preliminary study as being polymorphic were screened for variation between samples. No significant differences in allele distribution were detected between any of the samples obtained from the Faroe Bank in the north to Lisbon in the south, suggesting that squid throughout this range in the vicinity of the continental shelf are able to maintain panmixia, and effectively belong to a single population sharing a common gene pool. No clinal variation in allele distribution was detected throughout this range, a result which complements the findings of a detailed morphological companion study of the same individuals. Comparison of this homogenous European continental shelf population with squid from the Azores revealed highly significant (P<0.01) differences in allele distribution at five of the six polymorphic enzyme loci studied. A genetic identity value (I) equivalent to 0.93 over 33 loci was obtained. Analysis of F-statistics suggested migration rates between sites to be as low as one individual per five generations, a rate deemed insufficient under most models to prevent divergence by random genetic drift. The large distance and oceanic depths separating the Azores from continental Europe seem to present an effective barrier to gene flow to L. forbesi, a squid belonging to a family considered to be confined in distribution to relatively shallow, near coastal waters. The two populations of squid in the Azores and along the European continental shelf currently both ascribed to L. forbesi should therefore probably best be regarded as relative subspecies.