Population structure of short-beaked common dolphins (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) in the North Atlantic Ocean as revealed by mitochondrial and nuclear genetic markers

The understanding of population structure and gene flow of marine pelagic species is paramount to monitoring, management and conservation studies. Such studies are often hampered by the potentially high dispersal behavior of the species, the lack of obvious geographical barriers in the marine environment and the scarce sample availability. Short-beaked common dolphins (Delphinus delphis) are widespread in coastal and open-ocean habitats of the North Atlantic Ocean, nevertheless population structure and migratory patterns are poorly understood. Furthermore, concern has been raised about the status of the species because large numbers of dolphins have been taken incidentally in several fisheries throughout the North Atlantic in the past decades. In the present study, a large number of individual samples were obtained from seasonal and spatial aggregations of common dolphins from western (wNA) and eastern North Atlantic (eNA) regions, mostly using opportunistic sampling (i.e. from incidental entanglement in fishing gear or beach-cast carcasses). Genetic variability was investigated using nuclear (14 microsatellite loci) and mitochondrial (360 bp of the control region) genetic markers. Levels of genetic diversity were relatively high in all sampled areas and no evidence of recent reduction of effective population size (i.e. bottleneck) was detected at the nuclear loci. Significant population structure was detected between the two main regions (wNA and eNA) where it appeared to be more pronounced at mitochondrial (F _ST = 0.018, P < 0.001) than nuclear markers (F _ST = 0.005, P < 0.05), indicating the presence of at least two genetically distinct populations of common dolphins in the North Atlantic Ocean. In contrast, no significant genetic structure was detected between temporal aggregations of dolphins from within the same region, suggesting possible seasonal movement patterns at a regional scale. The observed levels of genetic differentiation between classes of markers are discussed here as a possible consequence of migratory patterns or recent population subdivision. An erratum to this article can be found at     

Genetic structure of leopard shark (<Emphasis Type="Italic">Triakis semifasciata</Emphasis>) populations in California waters

The leopard shark (Triakis semifasciata) is an important predator in coastal marine ecosystems of California, targeted by recreational and commercial fishermen and of specific interest in fisheries management. From October 2003 to August 2006, 169 leopard sharks were collected from the coast of California (between 40.750°N and 32.678°N) and analyzed for mitochondrial and nuclear genetic structure. Analyses of mtDNA control region sequences revealed relatively low levels of genetic variation (five haplotypes, average pairwise divergence π = 0.0067). In contrast, leopard sharks were highly polymorphic for inter simple sequence repeats (ISSRs), which characterize a broad range of the nuclear genome. The null hypothesis of panmixia in California waters was rejected for both genetic markers, and ISSRs displayed a statistically significant pattern of isolation by distance (IBD) across the species range (P = 0.002). A variety of analyses showed that divergence is most pronounced in the northernmost population of Humboldt Bay. Natal philopatry in T. semifasciata was tested using Siegel-Tukey tests on data partitioned by breeding site status, and sex-specific philopatry was tested by comparing IBD plots between sexes. Although there was some evidence for natal philopatry in leopard sharks (P = 0.038), and population divergence may be related to the proximity of breeding sites (P = 0.064), we found no support for sex-specific philopatry. In addition to identifying a novel set of highly variable genetic markers for use in shark population studies, these results may be used to better inform management decisions for leopard sharks in California.     

Evidence for restricted gene flow over small spatial scales in a marine snapping shrimp <Emphasis Type="Italic">Alpheus angulosus</Emphasis>

Despite the apparent absence of geographic barriers, connectivity among marine populations may be restricted by, for example, ecological or behavioral mechanisms. In such cases, populations may show genetic differentiation even over relatively small spatial scales. Here, mitochondrial sequence data from the cytochrome oxidase I (COI) gene and seven polymorphic microsatellite markers were used to investigate fine geographic scale population genetic structure in the snapping shrimp Alpheus angulosus, a member of the A. armillatus species complex, from collections in Florida, Jamaica, and Puerto Rico carried out from 1999 to 2005. The COI data showed a deep divergence that separated these samples into two mitochondrial clades, but this divergence was not supported by the microsatellite data. The COI data reflect past population divergence not reflected in extant population structure on the whole genome level. The microsatellite data also revealed evidence for moderate population structure between populations as close as ∼10 km, and no evidence for isolation by distance, as divergences between near populations were at least as strong as those between more broadly separated populations. Overall, these data suggest a role for restricted gene flow between populations, though the mechanisms that reduce gene flow in this taxon remain unknown.     

Microsatellite data reveal fine genetic structure in male Guiana dolphins (<Emphasis Type="Italic">Sotalia guianesis)</Emphasis> in two geographically close embayments at south-eastern coast of Brazil

A large macrogeographic differentiation has been observed among Sotalia guianensis populations along the South American coast. However, no genetic structure has been detected so far in closely distributed populations of this species, even though it has been observed in other cetaceans. Here, we examined the fine scale population structure for the largest populations of S. guianensis inhabiting Sepetiba and Paraty embayments at the south-eastern coast of Rio de Janeiro, Brazil. Analysis of mitochondrial DNA (mtDNA) control region sequences failed to detect variability among sequences. Conversely, evidence of significant male population structure was found on the basis of ten nuclear microsatellite loci. Surprisingly, the microsatellite markers were able to distinguish between individuals from the two embayments located 60 km apart. The results suggest that differences in habitat type and behavioral specializations are likely to explain the patterns of genetic structure. These findings should provide baselines for the management of communities exposed to increasing human-driven habitat loss.     

Song dialects do not restrict gene flow in an urban population of the orange-tufted sunbird, <Emphasis Type="Italic">Nectarinia osea</Emphasis>

Geographic variation in vocalizations is widespread in passerine birds, but its origins and maintenance remain unclear. In this study, we test the hypothesis that song dialect, a culturally transmitted trait, is related to the population genetic structure of the orange-tufted sunbird, Nectarinia osea. To address this, we compared mitochondrial DNA (mtDNA) sequence variation together with allele frequencies at five microsatellite loci from an urban population of sunbirds exhibiting two distinct song dialects on a microgeographic scale. Our findings reveal no association between dialect membership and genetic composition. All genetic measures, from both mitochondrial and nuclear DNA, indicate high levels of gene flow between both dialect populations. The low F _ST values obtained from mtDNA and microsatellite analysis imply that the variation among dialects does not account for more than 2%, at best, of the overall genetic variation found in the entire population. These measures fall well within the range of similar measures obtained in other studies of species exhibiting vocal dialects, most of which fail to detect any dialect-based genetic differentiation. The persistence of dialects in the orange-tufted sunbird may thus best be explained by dispersal of individuals across dialect boundaries and possibly from surrounding areas, followed by postdispersal vocal matching. Because genetic structuring appears weaker than cultural structure in this species, we discuss the behavioral mechanisms underlying dialect maintenance in the presence of apparent gene flow.     

Genetic evidence for the existence of cryptic species in an endangered clam <Emphasis Type="Italic">Coelomactra antiquata</Emphasis>

Coelomactra antiquata is a commercially important bivalve species, but has been suffering from severe population decline due to over-exploitation and the deterioration of environmental conditions. Previous genetic survey of C. antiquata conducted with allozymes combined with morphology revealed high levels of genetic differentiation between northern and southern populations which suggests a cryptic species might exist in C. antiquata. To test this hypothesis, amplified fragment length polymorphisms (AFLPs) and 16S rRNA gene sequence were used to re-evaluate the spatial genetic structure of six populations of C. antiquata along the coast of China. Both genetic markers display a sharp genetic break between the four northern populations (northern lineage) and two southern population (southern lineage). Large numbers of private alleles (AFLP) were found within the northern or southern populations and a deep divergence of about 6.5% in 16S rRNA gene sequence between the northern and southern lineages suggests the occurrence of potential cryptic or sibling species of C. antiquata. Applying previously published rates of mutation, divergence between the two lineages is estimated to have occurred approximately 3 million years ago and may be due to allopatric isolation during the middle Pliocene times. While no genetic differentiation was found within the northern or southern populations in both AFLP and 16S mtDNA markers, the results indicate that the northern and southern lineage should be managed separately and any translocation between the two areas should be avoided.     

Phylogeography of the sub-Antarctic notothenioid fish Eleginops maclovinus: evidence of population expansion

Phylogeography studies add insights into the geographic and evolutionary processes that underline the genetic divergence of populations. This work examines the geographic genetic structure of the Patagonian blennie, Eleginops maclovinus, a notothenioid (Perciformes) endemic to South American temperate and sub-Antarctic waters, using mitochondrial DNA cytochrome b sequences. We found 58 haplotypes in the analysis of 261 individual sequences of 833 base pairs in length. Among-population variance was very low (1.62%) and many haplotypes were shared between several populations across the species geographic range. Genetic differentiation was not consistent with a simple model of isolation by distance, possibly suggesting a lack of equilibrium between gene flow and local genetic drift. The analysis of mismatch distributions, neutrality tests, and the Bayesian Skyline Plot showed a pattern consistent with a recent population expansion event that may have taken place during the Middle Pleistocene.     

Phylogeographic structure of <Emphasis Type="Italic">Octopus vulgaris</Emphasis> in South Africa revisited: identification of a second lineage near Durban harbour

In a previous study that investigated genetic structure of Octopus vulgaris along the South African coast by sequencing the mitochondrial cytochrome oxidase III gene (COIII), all sequences generated were identical. Such a finding is unusual, because mitochondrial DNA mutates quickly, and several marine invertebrates present in southern Africa show considerable genetic variation and structure. We reanalysed the samples using two different mitochondrial markers, namely cytochrome oxidase I (COI) and the large ribosomal subunit (16S rRNA). Sequences of both these markers showed variation. The conclusion of the previous study, that South Africa’s O. vulgaris population is characterised by a lack of genetic structure along the coast, is rejected. Some specimens from Durban (southeast Africa) were genetically more different from those found in the remainder of the country than were specimens from other regions (Tristan da Cunha and Senegal). We suggest that the lineage in Durban may have been recently introduced.     

MtDNA population structure and gene flow in lingcod (<Emphasis Type="Italic">Ophiodon elongatus</Emphasis>): limited connectivity despite long-lived pelagic larvae

Lingcod, Ophiodon elongatus Girard, have a 3-month pelagic larval stage and are an important recreational and commercial species on the west coast of North America. Cytochrome-c oxidase I sequences from tissue samples were used to characterize population structure and infer patterns of gene flow from California to Alaska. No significant genetic structure was found when estimates of Wright’s F _ST (i.e., Φ_ST) were generated among all populations sampled. Nesting populations within regions, however, indicated that the inner coast of Washington State is distinct, a result corroborating previous allozyme work. Coalescent-based estimates of gene flow indicate that although migration can be high from an evolutionary perspective, nearly half of all comparisons among populations showed no gene flow in at least one direction. From an ecological perspective, moderate migration rates (Nm < 10) among most populations provide surprisingly limited connectivity at large (∼ 1,000 km) and small (∼100 km) spatial scales. Coalescent-based estimates also show that gene flow between the inner and the outer coasts is asymmetric, a result consistent with prevailing surface currents. Because the expected inter-locus variances for coalescent-based estimates of gene flow are likely large, future work will benefit from analyses of nuclear DNA markers. However, limited demographic connectivity on large spatial scales may help explain why stock recovery has been uneven, with greater recovery in the northern (87% rebuilt) than in the southern (24% rebuilt) fishery region, supporting a regional management strategy. These results suggest that despite a 3-month pelagic larval stage, some areas may be effectively closed with respect to both population dynamics and fishery management issues.     

Limited gene flow in the brooding coral Favia fragum (Esper, 1797)

Understanding population connectivity in corals is particularly important as these organisms are increasingly threatened by abiotic and biotic factors. This study examined the population genetic structure of the brooding coral Favia fragum across four locations in the Caribbean and Western Atlantic using mitochondrial and nuclear markers. Morphological features were also compared to test whether genetic diversity corresponds with skeletal morphology. When comparing across distantly related Caribbean and Bermudian locations, F _ST values were high and significant, indicating strong genetic structure. At a local scale, significant genetic structure was found among reefs in Panama, while no genetic structure was found among reefs within Barbados, Bermuda or Jamaica. Surprisingly, a single haplotype for each of the three markers examined was found in Bermuda, where samples varied significantly from all other locations in three out of four morphological features analyzed. These data indicate that gene flow of F. fragum may occur locally among reefs but is highly restricted at distant locations. Furthermore, isolated populations, such as that of Bermuda, must be self-seeding to maintain the observed genetic uniformity.     

Population structure of the common sole (<Emphasis Type="Italic">Solea solea</Emphasis>) in the Northeastern Atlantic and the Mediterranean Sea: revisiting the divide with EPIC markers

Spatial and temporal population genetic structures of the common sole, Solea solea, were studied in Northeastern Atlantic and Mediterranean Sea populations, using three polymorphic exon-primed intron-crossing (EPIC) markers. Results demonstrated significant multilocus differentiation among Eastern Mediterranean and a group composed by Western Mediterranean and Atlantic populations (θ = 0.150, P < 0.001), but also suggested unrecorded genetic differentiation of the Adriatic Sea population. No pattern of isolation-by-distance was recorded across the range covered by sampling, from the Kattegat to the Aegean Sea. Conversely to genetically structured Mediterranean populations, Atlantic populations ranging from Denmark to Portugal could be considered as representative of the same panmictic unit (θ = 0.009, not significant). Results further demonstrated stability of multilocus genetic structure among temporarily replicated cohort samples [0+, 1+, subadults] from several coastal and estuarine locations from Bay of Biscay, excepted for the amylase locus Am2B3-2 at one location (Pertuis d’Antioche). Despite coherence of such observed patterns of multilocus differentiation with previous allozymic surveys in sole, and with patterns generally obtained for other marine fish species, single-locus results from EPICs indicated divergent coalescence schemes supporting a complex response to ecology and history of sole’s populations. Results stress the use of nuclear genes such as EPIC markers to investigate population structure, but also historical, demographic, and possibly selective processes in marine fishes.     

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.     

Population differentiation in megrim (<Emphasis Type="Italic">Lepidorhombus whiffiagonis</Emphasis>) and four spotted megrim (<Emphasis Type="Italic">Lepidorhombus boscii</Emphasis>) across Atlantic and Mediterranean waters and implications for wild stock management

Megrim, Lepidorhombus whiffiagonis, and four spot megrim, Lepidorhombus boscii, are two marine fish species of high commercial interest. Despite their quite heavy exploitation little is known on the genetic structure of their populations. The present work aimed at characterizing the first seven microsatellites markers available for the two megrim species. These new markers were in a second step employed to describe the population structure of the two species among their almost entire habitat range (Atlantic and Mediterranean samples). Our study confirmed the existence of a strong genetic difference between Atlantic and Mediterranean megrim species already described in the literature for L. whiffiagonis on the basis of variations at ribosomal genes. Additionally our analysis gave the first evidences of a strong genetic differentiation among Atlantic populations in both megrim species (within Atlantic global F_ST in L. whiffiagonis and L. boscii were respectively 0.158 and 0.145). When describing megrim population structure, the comparison between allele-frequency-based tests (F_ST comparisons) and genotype-based inferences (Bayesian approach) gave evidences of a hierarchical structure of the populations. In conclusion, our work enlighten the existence of two different stocks within the Atlantic Ocean and one in the Mediterranean Sea that will clearly need to be managed separately. As the present results do not fully support the current megrim stock boundaries they will surely help to rethink megrim management policies in the future.     

Two markers and one history: phylogeography of the edible common sea urchin Paracentrotus lividus in the Lusitanian region

Benthic marine invertebrates with long-lived larvae are believed to have dispersal capabilities that contribute to maintaining genetic uniformity among populations over large geographical scales. However, both hydrological and biological factors may limit the actual dispersal of such larvae. We studied the population genetic structure of the edible common sea urchin Paracentrotus lividus (Lamarck, 1816), to explore its dispersal patterns in the Atlanto-Mediterranean region and, more specifically, to ascertain the role of the Strait of Gibraltar in shaping the genetic structure of this species. For this purpose, we analysed 158 individuals for the mitochondrial 16S rRNA gene and 151 of these for the nuclear single-copy intron adenine nucleotide transporter (ANT) from 16 localities from the Atlantic and Mediterranean basins, spanning over 4,000 km. Mitochondrial 16S rRNA shows higher genetic diversity in the Mediterranean than in the Atlantic and reveals a sharp break between the populations of both basins, probably as a consequence of the barrier imposed by the Almería–Orán hydrological front, situated east of the Strait of Gibraltar. Both markers suggest that a recent population expansion has taken place in both basins, most probably following the Messinian salinity crisis. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Molecular population structure of the kuruma shrimp <Emphasis Type="Italic">Penaeus</Emphasis><Emphasis Type="Italic"> japonicus</Emphasis> species complex in western Pacific

In a previous study on the kuruma shrimp Penaeus japonicus from the South China Sea, we detected high genetic divergence between two morphologically similar varieties (I and II) with distinct color banding patterns on the carapace, indicating the occurrence of cryptic species. In the present study, we clarify the geographical distribution of the two varieties in the western Pacific by investigating the genetic differentiation of the shrimp from ten localities. Two Mediterranean populations are also included for comparison. Based on the mitochondrial DNA sequence data, the shrimps are separated into two distinct clades representing the two varieties. Variety I comprises populations from Japan and China (including Taiwan), while variety II consists of populations from Southeast Asia (Vietnam, Singapore and the Philippines), Australia and the Mediterranean. Population differentiation is evident in variety II, as supported by restriction profiles of two mitochondrial markers and analysis of two microsatellite loci. The Australian population is genetically diverged from the others, whereas the Southeast Asian and Mediterranean populations show a close genetic relationship. Variety I does not occur in these three localities, while a small proportion of variety II is found along the northern coast of the South China Sea and Taiwan, which constitute the sympatric zone of the two varieties. The present study reveals high genetic diversity of P. japonicus. Further studies on the genetic structure of this species complex, particularly the populations in the Indian Ocean and Mediterranean, are needed not only to understand the evolutionary history of the shrimp, but also to improve the knowledge-based fishery management and aquaculture development programs of this important biological resource.     

Contrasting population structures in two sympatric fishes in the Baltic Sea basin

Detailed multispecies studies on the patterns of genetic variability and differentiation in marine environments are still rare. Using mitochondrial and nuclear genetic markers, we compared genetic variability and population structuring of threespine (Gasterosteus aculeatus) and ninespine (Pungitius pungitius) sticklebacks from the same eleven marine and six freshwater locations within the Baltic Sea basin. Analyses of both marker types revealed a significantly lower degree of genetic structuring in both marine and freshwater populations of threespine than those ninespine sticklebacks. Isolation-by-distance (IBD) was detected across the marine populations in both species, suggesting spatially limited gene flow. However, the levels of genetic diversity and differentiation across the localities were uncorrelated between the two species in both marine and freshwater environments. Accordingly, estimates of effective population sizes were larger and migration rates were higher for three- than for ninespine sticklebacks. Hence, ninespine stickleback populations from the Baltic Sea basin appear to be subject to stronger genetic drift than sympatric threespine sticklebacks, and the proximate reason for this difference is likely to be found from autecological differences between the two species. In accordance with the earlier studies, genetic variability was higher and the degree of genetic differentiation was lower in marine than in freshwater populations in both species.     

Temporal stability of genetic population structure in the New Zealand snapper, <Emphasis Type="Italic">Pagrus auratus</Emphasis>, and relationship to coastal currents

Genetic diversity and population structure of snapper (Pagrus auratus, Bloch and Schneider), a coastal demersal sparid fish, were determined using six nuclear microsatellite loci and SSCP (single strand conformational polymorphism) analysis of themitochondrial (mt) DNA D-loop in samples collected across the range of the species in New Zealand. Microsatellite data showed similar results to allozyme data collected in the late 1970s that found differentiation between the north-east and southern populations. In addition, an isolated population of snapper in Tasman Bay was identified. The two data sets provide evidence for the temporal stability of the genetic population structure of snapper over 22 years, with differentiation over relatively small spatial scales separated by oceanographic boundaries rather than isolation by distance. In contrast to nuclear markers, mtDNA did not reveal any significant genetic heterogeneity among samples.     

Sink populations in carnivore management: cougar demography and immigration in a hunted population.

Carnivores are widely hunted for both sport and population control, especially where they conflict with human interests. It is widely believed that sport hunting is effective in reducing carnivore populations and related human-carnivore conflicts, while maintaining viable populations. However, the way in which carnivore populations respond to harvest can vary greatly depending on their social structure, reproductive strategies, and dispersal patterns. For example, hunted cougar (Puma concolor) populations have shown a great degree of resiliency. Although hunting cougars on a broad geographic scale (> 2000 km2) has reduced densities, hunting of smaller areas (i.e., game management units, < 1000 km2), could conceivably fail because of increased immigration from adjacent source areas. We monitored a heavily hunted population from 2001 to 2006 to test for the effects of hunting at a small scale (< 1000 km2) and to gauge whether population control was achieved (lambda < or = 1.0) or if hunting losses were negated by increased immigration allowing the population to remain stable or increase (lambda > or = 1.0). The observed growth rate of 1.00 was significantly higher than our predicted survival/fecundity growth rates (using a Leslie matrix) of 0.89 (deterministic) and 0.84 (stochastic), with the difference representing an 11-16% annual immigration rate. We observed no decline in density of the total population or the adult population, but a significant decrease in the average age of independent males. We found that the male component of the population was increasing (observed male population growth rate, lambda(OM) = 1.09), masking a decrease in the female component (lambda(OF) = 0.91). Our data support the compensatory immigration sink hypothesis; cougar removal in small game management areas (< 1000 km2) increased immigration and recruitment of younger animals from adjacent areas, resulting in little or no reduction in local cougar densities and a shift in population structure toward younger animals. Hunting in high-quality habitats may create an attractive sink, leading to misinterpretation of population trends and masking population declines in the sink and surrounding source areas.     

Genetic structure of natural populations of California red abalone (<Emphasis Type="Italic">Haliotis rufescens</Emphasis>) using multiple genetic markers

Mitochondrial cytochrome oxidase subunit one (COI) sequence, nuclear microsatellites, and amplified fragment length polymorphisms (AFLPs) were used to evaluate connectivity among nine red abalone (Haliotis rufescens) populations sampled between August 1998 and November 2003 along approximately 1,300 km of California coastline from Crescent City (41°46′N, 124°12′W) to San Miguel Island (34°02′N, 120°22′W). COI sequences and microsatellite genotypes did not show significant genetic divergence among nine sampled populations. A subset of five populations spanning the geographic range of the study was scored for 163 polymorphic AFLP markers. Of these, 41 loci showed significant divergence (P < 0.001) among populations. Still, no AFLP markers were diagnostic for any of the study populations, and assignment tests did not consistently assign individuals to the correct population. Although the AFLP data are the first to suggest there is significant genetic differentiation among California red abalone populations, the discordance between the different genetic markers needs further study before unambiguous conclusions can be drawn with respect to connectivity among the populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.