Population differentiation of the Atlantic spotted dolphin (Stenella frontalis) in the western North Atlantic, including the Gulf of Mexico

Information about the genetic population structure of the Atlantic spotted dolphin [Stenella frontalis (G. Cuvier 1829)] in the western North Atlantic would greatly improve conservation and management of this species in USA waters. To this end, mitochondrial control region sequences and five nuclear microsatellite loci were used to test for genetic differentiation of Atlantic spotted dolphins in the western North Atlantic, including the Gulf of Mexico (n=199). Skin tissue samples were collected from 1994–2000. Significant heterozygote deficiencies in three microsatellite loci within samples collected off the eastern USA coast prompted investigation of a possible Wahlund effect, resulting in evidence for previously unsuspected population subdivision in this region. In subsequent analyses including three putative populations, two in the western North Atlantic (n=38, n=85) and one in the Gulf of Mexico (n=76), significant genetic differentiation was detected for both nuclear DNA (R _ST=0.096, P≤0.0001) and mitochondrial DNA (Φ _ST=0.215, P≤0.0001), as well as for all pair-wise population comparisons for both markers. This genetic evidence for population differentiation coupled to known biogeographic transition zones at Cape Hatteras, North Carolina and Cape Canaveral, Florida, USA, evidence of female philopatry, and preliminary support for significant genetic differences between previously documented morphotypes of Atlantic spotted dolphins in coastal and offshore waters all indicate that the biology and life history of this species is more complex than previously assumed. Assumptions of large, panmictic populations might not be accurate in other areas where S. frontalis is continuously distributed (e.g., eastern Atlantic), and could have a detrimental effect on long-term viability and maintenance of genetic diversity in this species in regions where incidental human-induced mortality occurs.

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Population structure of red porgy, <Emphasis Type="Italic">Pagrus pagrus</Emphasis>, in the Atlantic Ocean

The red porgy, Pagrus pagrus (L.), is a protogynous sparid associated with reefs and hard bottom habitat throughout the warm-temperate Atlantic Ocean. In this study, the degree of geographic population differentiation in Atlantic populations was examined with microsatellite and mitochondrial DNA markers (mtDNA). Six microsatellite loci were amplified and scored in 690 individuals from the eastern North Atlantic (Crete, Madeira, and Azores), western North Atlantic (North Carolina to Florida, and the eastern Gulf of Mexico), and Brazil. At two loci, fixed allelic differences were found among the three major geographic areas, while frequency differences were observed at three other loci. The DNA of 371 individuals was amplified at the mtDNA control region, and 526 bp were sequenced. Tamura–Nei’s D was used as a measure of nucleotide diversity and divergence: diversity averaged 0.011 within samples, while the corrected divergence averaged 0 between samples within the same area and 0.061 between samples from different areas. Transversion haplotype minimum spanning networks, nucleotide divergence, and F _ST values all show that the western Atlantic samples were more closely related to each other than any was to samples from the eastern North Atlantic. Within the western North Atlantic, no significant population differentiation was observed, and within the eastern North Atlantic, only the Azores sample showed detectable differences from Crete and Madeira. These data indicate general homogeneity within large areas, and deep divisions between these areas. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Phylogeography of surfclams,<Emphasis Type="Italic"> Spisula solidissima</Emphasis>, in the western North Atlantic based on mitochondrial and nuclear DNA sequences

The Atlantic surfclam, Spisula solidissima (Dillwyn), is broadly distributed in sandy sediments of the western North Atlantic between the Gulf of St. Lawrence and the Gulf of Mexico. In the United States, a substantial commercial fishery between Long Island and Cape Hatteras harvests offshore populations of one subspecies, S. s. solidissima. A smaller coastal form, S. s. similis Say (also known as S. s. raveneli Conrad), has a partially sympatric geographic distribution, but differs in several life-history characteristics. DNA sequence variation in mitochondrial cytochrome oxidase I (COI) and in introns at two nuclear calmodulin loci was examined to measure genetic divergence between the two subspecies and to test for population structure among populations of S. s. solidissima. Surfclams were collected from seven localities between 1994 and 2001. Based on both mitochondrial and nuclear DNA variation, the two subspecies of S. solidissima are reciprocally monophyletic, with a net COI divergence of 13.9%, indicating long-term reproductive isolation. The only significant differentiation among populations of S. s. solidissima (based on an AMOVA analysis of COI sequences) was between the Gulf of St. Lawrence and more southerly populations. A long internal branch in the S. s. solidissima genealogy coupled with low haplotype diversity in the northern-most population suggests that populations north and south of Nova Scotia have been isolated from each other in the past, with gene exchange more recently. Populations of S. s. similis from Atlantic and Gulf of Mexico coasts had a net COI divergence of 9.2%. Thus, diversification of Spisula spp. clams in the western North Atlantic involved an early adaptive divergence between coastal and offshore forms, with later barriers to dispersal emerging in the offshore form from north to south and in the coastal form between Atlantic and Gulf of Mexico populations.     

Satellite tracking reveals a dichotomy in migration strategies among juvenile loggerhead turtles in the Northwest Atlantic

Few data are available on the movements and behavior of immature Atlantic loggerhead sea turtles (Caretta caretta) from their seasonal neritic foraging grounds within the western north Atlantic. These waters provide developmental habitat for loggerheads originating from several western Atlantic nesting stocks. We examined the long-term movements of 23 immature loggerheads (16 wild-caught and seven headstart turtles) characterizing their seasonal distribution, habitat use, site fidelity, and the oceanographic conditions encountered during their migrations. We identified two movement strategies: (1) a seasonal shelf-constrained north–south migratory pattern; and (2) a year-round oceanic dispersal strategy where turtles travel in the Gulf Stream to the North Atlantic and their northern dispersal is limited by the 10–15°C isotherm. When sea surface temperatures dropped below 20°C, neritic turtles began a migration south of Cape Hatteras, North Carolina (USA) where they established fidelity to the waters between North Carolina’s Outer Banks and the western edge of the Gulf Stream along outer continental shelf. Two turtles traveled as far south as Florida. Several turtles returned to their seasonal foraging grounds during subsequent summers. Northern movements were associated with both increased sea surface temperature (>21°C) and increased primary productivity. Our results indicate strong seasonal and interannual philopatry to the waters of Virginia (summer foraging habitat) and North Carolina (winter habitat). We suggest that the waters of Virginia and North Carolina provide important seasonal habitat and serve as a seasonal migratory pathway for immature loggerhead sea turtles. North Carolina’s Cape Hatteras acts as a seasonal “migratory bottleneck” for this species; special management consideration should be given to this region. Six turtles spent time farther from the continental shelf. Three entered the Gulf Stream near Cape Hatteras, traveling in the current to the northwest Atlantic. Two of these turtles remained within an oceanic habitat from 1 to 3 years and were associated with mesoscale features and frontal systems. The ability of large benthic subadults to resume an oceanic lifestyle for extended periods indicates plasticity in habitat use and migratory strategies. Therefore, traditional life history models for loggerhead sea turtles should be reevaluated.     

Temporal stability and spatial divergence of mitochondrial DNA haplotype frequencies in red drum (Sciaenops ocellatus) from coastal regions of the western Atlantic Ocean and Gulf of Mexico

Restriction-site variation in mitochondrial (mt) DNA was assayed among 1675 red drum (Sciaenops ocellatus Linnaeus) sampled from 20 localities along the southeastern coast of the USA (western Atlantic) and the Gulf of Mexico (Gulf). Up to four consecutive year-classes (cohorts) were sampled at most localities. Nucleotide-sequence divergence among 170 mtDNA haplotypes identified ranged (in percentage) from 0.184 to 1.913, with a mean (±SD) of 0.887 ± 0.300. Comparisons of mtDNA haplotype frequencies across year-classes within localities were non-significant, indicating temporal stability of breeding components within localities. Significant heterogeneity in mtDNA haplotype frequencies was found across all localities, between (pooled) samples from the western Atlantic and the Gulf, and among geographically spaced, regional groupings in the Gulf. Genetic divergence between subpopulations of red drum in the western Atlantic and Gulf follows a pattern exhibited in other marine fishes, and probably stems from physical (historical environmental heterogeneity, absence of suitable habitat, and current patterns) and, perhaps, behavioral factors. Genetic differences among red drum in the Gulf appear to be due largely to an isolation-by-distance effect that is attributable to behavioral factors. The latter may include female philopatry to natal bays or estuaries, limited offshore (coastwise) movement of females relative to their natal bay or estuary, or both. Genetic divergence among red drum in the Gulf occurs despite high gene flow (estimated as the number of genetic effective migrants in an island mode). Conservation and management of red drum should be based on the premise that strategies for a given bay or estuary will impact geographically proximal bays or estuaries more than distal ones. Trajectories of correlograms in spatial autocorrelation analysis suggest a geographic neighborhood size, relative to genetic migration of red drum from a bay or estuary, of roughly 500 to 600 km. Received: 22 July 1998 / Accepted: 19 November 1998     

Anguilliform larvae collected off North Carolina

The distinctive larval stage of eels (leptocephalus) facilitates dispersal through prolonged life in the open ocean. Leptocephali are abundant and diverse off North Carolina, yet data on distributions and biology are lacking. The water column (from surface to 1,293 m) was sampled in or near the Gulf Stream off Cape Hatteras, Cape Lookout, and Cape Fear, North Carolina during summer through fall of 1999–2005, and leptocephali were collected by neuston net, plankton net, Tucker trawl, and dip net. Additional samples were collected nearly monthly from a transect across southern Onslow Bay, North Carolina (from surface to 91 m) from April 2000 to December 2001 by bongo and neuston nets, Methot frame trawl, and Tucker trawl. Overall, 584 tows were completed, and 224 of these yielded larval eels. The 1,295 eel leptocephali collected (combining all methods and areas) represented at least 63 species (nine families). Thirteen species were not known previously from the area. Dominant families for all areas were Congridae (44% of individuals, 11 species), Ophichthidae (30% of individuals, 27 species), and Muraenidae (22% of individuals, ten species). Nine taxa accounted for 70% of the overall leptocephalus catches (in order of decreasing abundance): Paraconger caudilimbatus (Poey), Gymnothorax ocellatus Agassiz complex, Ariosoma balearicum (Delaroche), Ophichthus gomesii (Castelnau), Callechelys muraena Jordan and Evermann, Letharchus aliculatus McCosker, Rhynchoconger flavus (Goode and Bean), Ophichthus cruentifer (Goode and Bean), Rhynchoconger gracilior (Ginsburg). The top three species represented 52% of the total eel larvae collected. Most leptocephali were collected at night (79%) and at depths > 45 m. Eighty percent of the eels collected in discrete depth Tucker trawls at night ranged from mean depths of 59–353 m. A substantial number (38% of discrete depth sample total) of larval eels were also collected at the surface (neuston net) at night. Daytime leptocephalus distributions were less clear partly due to low catches and lower Tucker trawl sampling effort. While net avoidance may account for some of the low daytime catches, an alternative explanation is that many species of larval eels occur during the day at depths > 350 m. Larvae of 21 taxa of typically shallow water eels were collected at depths > 350 m, but additional discrete depth diel sampling is needed to resolve leptocephalus vertical distributions. The North Carolina adult eel fauna (estuary to at least 2,000 m) consists of 51 species, 41% of which were represented in these collections. Many species of leptocephali collected are not yet known to have juveniles or adults established in the South Atlantic Bight or north of Cape Hatteras. Despite Gulf Stream transport and a prolonged larval stage, many of these eel leptocephali may not contribute to their respective populations.     

Population genetics of the American oyster Crassostrea virginica along the Atlantic coast and the Gulf of Mexico

An examination by protein-gel electrophoresis of 19 different geographical populations of the American oyster Crassostrea virginica (Gmelin) was conducted along the Atlantic coast and the Gulf of Mexico. Estimates were made of levels of genetic variation and similarity among the populations based on 32 structural loci. The percentage of loci polymorphic ranged from 46.9 to 65.6% along the Atlantic coast while the estimate ranged from 54.8% to 68.8% on the Gulf of Mexico. The percentage of loci heterozygous ranged from 18.6 to 23.6% along the Atlantic coast and from 20.0 to 25.4% in the Gulf of Mexico. The genetic similarities between all contiguous populations from Cape Cod, Massachusetts to Corpus Christi, Texas were estimated as 99%, while the same estimate between Corpus Christi and Brownsville, Texas was computed as 93%, indicating a major transition in genetic structure for the Brownsville population of the Laguna Madre. The study revealed that the migration of planktonic oyster larvae is predominantly in a westerly direction along the Gulf of Mexico and that gene flow appears to be disrupted in the region of the Laguna Madre. Evidence has been presented for single gene selection at the Lap-2 and Pgi loci in the form of macrogeographical clines in allele frequencies with changing environmental conditions.     

Mitochondrial DNA and electronic tracking reveal population structure of Atlantic bluefin tuna (Thunnus thynnus)

Population subdivision was examined in Atlantic bluefin tuna (Thunnus thynnus) through sequencing of the control region of the mitochondrial genome. A total of 178 samples from the spawning grounds in the Gulf of Mexico, Bahamas and Mediterranean Sea were analyzed. Among the samples from these locations were 36 electronically tagged bluefin tuna that were tagged in the North Atlantic and subsequently traveled to one of these known spawning grounds during the spawning season. Bluefin tuna populations from the Gulf of Mexico and the Mediterranean Sea were found to be genetically distinct based on Φ_st, and sequence nearest neighbor analyses, showing that these two major spawning areas support independent stocks. Sequence nearest neighbor analysis indicated significant population subdivision among the Gulf of Mexico, western Mediterranean and eastern Mediterranean Sea. However, it was not possible to find significant pairwise differences between any sampling areas when using all samples. If only samples that had a high likelihood of assignment to a specific spawning site were used (young of the year, spawning adults), the differentiation increased among all sampling areas and the Western Mediterranean Sea was distinct from the Eastern Mediterranean Sea and the Gulf of Mexico. It was not possible to distinguish samples from the Bahamas from those collected at any of the other sampling sites. These data support tagging results that suggested distinctness of the Gulf of Mexico, Eastern and Western Mediterranean Sea spawning areas. This level of stock differentiation is only possible if Atlantic bluefin tuna show strong natal homing to individual spawning grounds.     

Mitochondrial and nuclear DNA analysis of population subdivision among young-of-the-year Spanish mackerel (Scomberomorus maculatus) from the western Atlantic and Gulf of Mexico

The degree of population subdivision among collections of young-of-the-year (YOY) Spanish mackerel Scomberomorus maculatus sampled from along the Atlantic coast of the United States (Atlantic) and Gulf of Mexico (Gulf) was examined. Young-of-the-year were collected from spawning areas to minimize possible mixing among putative spawning stocks that may occur at later life history stages along migratory pathways or at over-wintering grounds. Restriction fragment length polymorphism (RFLP) analysis of the mitochondrial DNA (mtDNA) ND4 region and a nuclear actin intron was conducted. Collections in each of two years were analyzed to minimize bias associated with sampling at a single point in space or time. Substantial variation was detected at ND4 (haplotype diversity=0.81) and at the actin intron locus (gene diversity=0.21). Significant differences in gene diversity or allele frequencies were not detected among temporal samples at any of three locations. A homogeneous distribution of genetic variance among samples from widely spaced geographic regions was consistent with the hypothesis that Spanish mackerel comprise a single intermingling genetic stock. Power analysis showed that the mitochondrial analysis from this study was much more likely to detect population subdivision than analysis of the nuclear actin locus, despite lower mtDNA sample size. Received: 3 January 2000 / Accepted: 28 August 2000     

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     

Mitochondrial DNA differentiation and population structure in red drum (Sciaenops ocellatus) from the Gulf of Mexico and Atlantic Ocean

Variation in mitochondrial (mt)DNA was examined among 473 red drum (Sciaenops ocellatus) sampled in 1988 and 1989 from nearshore localities in the northern Gulf of Mexico (Gulf) and the Atlantic coast of the southeastern United States (Atlantic). Data were combined with those from a previous study to generate a total of 871 individuals sampled from 11 localities in the Gulf and 5 localities in the Atlantic. Individuals assayed were from the 1986 and 1987 year-classes. A total of 118 composite mtDNA genotypes (haplotypes) was found. The percentage nucleotide sequence divergence among the 118 haplotypes ranged from 0.184 to 1.913, with a mean (±SE) of 0.878±0.004. MtDNA nucleon diversities and intrapopulational nucleotide-sequence divergence values were similar over all Gulf and Atlantic localities, and were high relative to most fish species surveyed to date. These data indicate that the perceived decline in red drum abundance appears not to have affected the genetic variability base of the species. Significant heterogeneity in the frequencies of at least four haplotypes was detected between pooled samples from the Gulf vs pooled samples from the Atlantic. No heterogeneity was found among localities from the Gulf or localities from the Atlantic. High levels of gene flow among all localities were inferred from F _ST values (a measure of the variance in mtDNA haplotype frequencies) and from Slatkin's qualitative and quantitative analyses. Parsimony and phenetic analyses revealed no strong evidence for phylogeographic cohesion of localities, although there was weak support for cohesion of four of five localities from the Atlantic. These data indicate that the red drum population is subdivided, with weakly differentiated subpopulations (stocks) occurring in the northern Gulf and along the Atlantic coast of southeastern USA. Spatial autocor-relation analysis and heterogeneity tests of haplotype frequencies among regions within the Gulf supported the hypothesis of increased gene flow among neighboring localities; i.e., migration of individuals within the Gulf may be inversely related to geographic distance from an estuary or bay of natal origin. Estimates of evolutionary effective female-population size indicate that the red drum subpopulations may be large.     

Speciation in western Atlantic stone crabs (genus Menippe): the role of geological processes and climatic events in the formation and distribution of species

Electrophoretically detectable variation in 38 proteins and color morphology were used to determine the evolutionary relationships of crabs of the genus Menippe (Xanthidae) in the southeastern USA. Both allele frequencies (=genotype) and color morphology (=phenotype) showed that one species, Menippe mercenaria, is probably a taxonomic supergroup composed of two taxa (semispecies). One taxon (the western Gulf form) is distributed from northwest Florida westward through Texas. The second (the peninsular Florida form) ranges through the Florida peninsula from northwest to east central Florida, and in North Carolina. The taxa appear to have hybridized in two discrete regions: in the Gulf of Mexico (northwest Florida) and in the Atlantic Ocean (east central Florida to South Carolina). The agreement of patterns of geographic variation in genotype and phenotype with the geological record and estimated times of divergence based on genetic distances suggests that the observed patterns are the product of the influence of Late Cenozoic changes in climate and geology. The Atlantic zone of hybridization was formed prior to the closure of the seaway across north Florida connecting the Gulf of Mexico and the Atlantic, and the northwest Florida zone at some time subsequent to the closure. The present distribution, and the location of zones of hybridization, between the two semispecies of M. mercenaria illustrate the importance of the interaction of historical geological and climatic events with ecological boundaries in determining the distribution and interactions of shallow water marine species.     

Morphological and genetic evidence for vicariance and refugium in Atlantic and Gulf of Mexico populations of the hermit crab Pagurus longicarpus

The number and wide variety of southeastern United States marine taxa with significant differentiation between Gulf of Mexico and Atlantic Ocean populations suggests that these taxa may have experienced major vicariance events, whereby populations were subdivided by geological or ecological barriers. The present study compared variation in morphology, allozymes, and mtDNA in Gulf of Mexico and western Atlantic populations of the longwrist hermit crab Pagurus longicarpus Say collected during 1997 and 1998. Combined Atlantic populations had significantly fewer denticles on the second segment of the third maxilliped than did Gulf of Mexico populations, and the mean ratio of dactyl length to propodus length was significantly greater in the Atlantic crabs than in the Gulf of Mexico crabs. Allozyme allele frequencies at three loci showed genetic differentiation between a Gulf of Mexico population and two Atlantic populations. Analysis of mtDNA sequence data revealed a clear reciprocal monophyly between Gulf and Atlantic populations, with an estimated divergence age of ~0.6 million years ago. This estimated age of divergence is significantly more recent than an age previously estimated for its congener Pagurus pollicaris (~4 million years ago), suggesting that species with a similar genetic break between Gulf and Atlantic populations may not necessarily share an identical history. Surprisingly, there is evidence of geographic subdivision within Atlantic populations of P. longicarpus along the east coast of North America. This differentiation is especially strong between Nova Scotia and southern populations, suggesting that the Nova Scotia population may represent survivors from a northern refugium during the last glacial maximum.     

Otolith record of age, growth, and ontogeny in larval and pelagic juvenile Stephanolepishispidus (Pisces: Monacanthidae)

Juveniles of the planehead filefish Stephano-lepishispidus (Pisces: Monacanthidae) (Linnaeus, 1766) are a major component of the Sargassum spp. community, yet little is known of their ecology. In this study, the otolith record of age, growth, and ontogeny in S. hispidus was examined. Juveniles caught off Beaufort Inlet, North Carolina (USA) on 30 June 1996 were marked with alizarin complexone and reared in a flow-through, outdoor tank for up to 19 days. Examination of marked otoliths at several time intervals showed that increment formation was not significantly different than one increment per day, and thus, increment number was used to estimate age. Depth-distribution, morphology, and meristics of larvae and juveniles collected (1990–1992) between Cape Romain, South Carolina, and Cape Hatteras, North Carolina, were examined to identify the timing of the larval to juvenile transition. All indicators suggested the transition occurred between 17 and 20 days. Mean otolith increment widths exhibited a marked change at about 20 days, coinciding with the timing of the larval to juvenile transition and a change in the depth distribution from bottom to surface waters. Increment width of individual juveniles, however, did not exhibit the same pattern; only 40% conformed to the pattern identified for all fish. Thus, the record of the larval to juvenile transition is clear at the population level, but unresolved at the individual level. Received: 1 November 1999 / Accepted: 18 December 2000     

Genetic structure of Atlantic and Gulf of Mexico populations of sea bass,menhaden, and sturgeon: Influence of zoogeographic factors and life-history patterns

To assess the influence of zoogeographic factors and life-history parameters (effective population size, generation length, and dispersal) on the evolutionary genetic structure of marine fishes in the southeastern USA, phylogeographic patterns of mitochondrial DNA (mtDNA) were compared between disjunct Atlantic and Gulf of Mexico populations in three coastal marine fishes whose juveniles require an estuarine or freshwater habitat for development. Black sea bass (Centropristis striata), menhaden (Brevoortia tyrannus andB. patronus) and sturgeon (Acipenser oxyrhynchus) samples were collected between 1986 and 1988. All species showed significant haplotype frequency differences between the Atlantic and Gulf, but the magnitude and distribution of mtDNA variation differed greatly among these taxa: sea bass showed little within-region mtDNA polymorphism and a clear phylogenetic distinction between the Atlantic and Gulf; menhaden showed extensive within-region polymorphism and a paraphyletic relationship between Atlantic and Gulf populations; and sturgeon exhibited very low mtDNA diversity both within regions and overall. Evolutionary effective sizes of the female populations (N _f (e)) estimated from the mtDNA data ranged fromN _f (e) = 50 (Gulf of Mexico sturgeon) toN _f (e) = 800 000 (Atlantic menhaden), and showed a strong rank-order agreement with the current-day census sizes of these species. The relationship betweenN _f (e) and the estimated times of divergence (t) among mtDNA lineages (from conventional clock calibrations) predicts the observed phylogenetic distinction between Atlantic and Gulf sea bass, as well as the paraphyletic pattern in menhaden, provided the populations have been separated by the same long-standing zoogeographic barriers thought to have influenced other coastal taxa in the southeastern USA. However, vicariant scenarios alone cannot explain other phylogenetic aspects of the menhaden (and sturgeon) mtDNA data and, for these species, recent gene flow between the Atlantic and Gulf coasts is strongly implicated. These data are relevant to management and conservation issues for these species.Please address all requests for reprints to Dr. J. C. Avise     

Genetic structure of summer flounder (Paralichthys dentatus) populations north and south of Cape Hatteras

Genetic divergence among samples of summer flounder (Paralichthys dentatus, Linnaeus) was estimated to evaluate the effect of Cape Hatteras, a well-known zoogeographic barrier in the western Atlantic Ocean, on intraspecific gene flow. Previous studies based on meristic and morphometric variation and tagging data suggested that gene flow might be limited among summer flounder populations north and south of Cape Hatteras, although the high vagility of both juveniles and adults suggests otherwise. We analyzed genetic diversity revealed in the mitochondrial DNA (mtDNA) control region in samples of juveniles and adults collected in 1992 to 1996 from coastal sites from Buzzard's Bay, Massachusetts to Charleston, South Carolina. In contrast to previous morphological studies, analyses of mtDNA variation reveal no significant population subdivision centered around Cape Hatteras. Received: 17 September 1997 / Accepted: 8 September 1998     

Genetic evidence of cryptic speciation within hammerhead sharks (Genus Sphyrna)

Surveys of genetic variation within cosmopolitan marine species often uncover deep divergences, indicating historical separation and potentially cryptic speciation. Based on broad geographic (coastal eastern North America, Gulf of Mexico, western Africa, Australia, and Hawaii) and temporal sampling (1991–2003), mitochondrial (control region [CR] and cytochrome oxidase I [COI]) and nuclear gene (lactate dehydrogenase A intron 6 [LDHA6]) variation among 76 individuals was used to test for cryptic speciation in the scalloped hammerhead, Sphyrna lewini (Griffith and Smith). CR and COI gene trees confirmed previous evidence of divergence between Atlantic and Indo-Pacific scalloped hammerhead populations; populations were reciprocally monophyletic. However, the between-basin divergence recorded in the mtDNA genome was not reflected in nuclear gene phylogenies; alleles for LDHA6 were shared between ocean basins, and Atlantic and Indo-Pacific populations were not reciprocally monophyletic. Unexpectedly, CR, COI, and LDHA6 gene trees recovered a deep phylogenetic partition within the Atlantic samples. For mtDNA haplotypes, which segregated by basin, average genetic distances were higher among Atlantic haplotypes (CR: D _HKY=0.036, COI: D _GTR=0.016) than among Indo-Pacific haplotypes (CR: D _HKY=0.010, COI: D _GTR=0.006) and approximated divergences between basins for CR (D _HKY=0.036 within Atlantic; D _HKY=0.042 between basins). Vertebral counts for eight specimens representing divergent lineages from the western north Atlantic were consistent with the genetic data. Coexistence of discrete lineages in the Atlantic, complete disequilibrium between nuclear and mitochondrial alleles within lineages and concordant partitions in genetic and morphological characters indicates reproductive isolation and thus the occurrence of a cryptic species of scalloped hammerhead in the western north Atlantic. Effective management of large coastal shark species should incorporate this important discovery and the inference from sampling that the cryptic scalloped hammerhead is less abundant than S. lewini, making it potentially more susceptible to fishery pressure.     

Experimental interbreeding between geographically separated populations of the marine wood-boring isopod Limnoria tripunctata with preliminary indications of hybrid vigor

Populations of the marine wood-boring isopod Limnoria tripunctata Menzies were cultivated and interbred in the laboratory in order to determine whether geographically separated populations would interbreed. Two series of interbreeding studies were conducted. In the first, field populations from 11 Atlantic and Caribbean sites and 3 Pacific sites were interbred with a North Western Atlantic population from Beaufort, North Carolina (USA). Viable crossings, to at least F ₁, occurred in all but one case—that of a BeaufortxChatham (Massachusetts, USA) cross. In the second series of experiments, field populations from 9 Atlantic and Caribbean sites were crossed with populations from St. Teresa, Florida, Gulf of Mexico. In this case also, all crosses except one produced viable offspring, not only to the F ₁, but to the F ₂ generations as well. The St. Teresa x Chatham cross was not successful. Geographic distance was not a factor in the success of crossbreeding between disjunct populations. Chatham populations (mainly) are now believed to represent a valid but previously obscure species, Limnoria tuberculata Sowinski, and not a variant of L. tripunctata Menzies. Enhanced viality or vigor, as determined from rate of wood destruction, population growth rate and survival, was indicated in several crossings, but this was not necessarily correlated with distance between populations. Observations were made on the manner in which the species has achieved a worldwide distribution, and a hypothesis is put forward to explain the evolution of species related to L. tripunctata.Contribution from Zoology Department, Duke University, Durham, North Carolina, and Duke University Marine Laboratory, Beaufort, North Carolina, and the Department of Oceanography, No. 289, Florida State University, Tallahassee, Florida. Support in part from NSF Grant No. GB-6898.     

Population genetics of the blue crabCallinectes sapidus: modest population structuring in a background of high gene flow

To determine the genetic population structure of blue crabs (Callinectes sapidus Rathbun), electrophoretic allozyme analysis was performed on 750 individuals collected from 16 nearshore locations ranging from New York to Texas, USA. Twenty enzymes and non-enzymatic proteins coded by 31 presumptive loci were examined. Twenty-two loci were either monomorphic or polymorphic at less than theP ₉₅ level; alleles for these polymorphic loci were geographically dispersed. Allele frequencies for three of the remaining polymorphic loci were homogeneous over all populations, as were levels of polymorphism and heterozygosity. Phenograms generated by the UPGMA (unweighted pair-group method using arithmetic averages) and distance Wagner methods exhibited no geographic pattern in the clustering of populations. Estimates ofN _em (effective number of migrants per generation between populations) indicated substantial gene flow, with aalues sufficiently high to infer panmixia between all blue crab populations from New York to Texas. However, despite this high level of gene flow, two striking patterns of geographic differentiation occurred: genetic patchiness and clinal variation. Allele frequencies atEST-2, GP-1, IDHP-2, DPEP-1, DPEP-2, andTPEP exhibited genetic patchiness on local and range-wide geographic scales, and allele frequencies atEST-2 varied temporally. Genetic patchiness in blue crabs is likely to be the result of the pre-settlement formation and subsequent settlement of genetically heterogeneous patches of larvae; allele frequencies of those larval patches may then be further modified through ontogeny by localized selection. In the Atlantic Ocean, a regional latitudinal cline ofEST-2 allele frequencies was superimposed on the range-wide genetic patchiness exhibited by that locus. This pattern against a background of high gene flow is highly likely to be maintained by selection. In estuaries along the Atlantic Ocean coast, a combination of low adult long-distance migration and a high retention rate of locally spawned larvae could serve to segregate populations and allow for the development of the geographic cline inEST-2. The Gulf of Mexico showed no apparent cline, perhaps due to long-distance migration of females in some regions of the Gulf, or to masking by genetic patchiness. These results emphasize the importance of both ecological and evolutionary time scales and structuring mechanisms in determining genetic population structure.     

Genetic studies in marine fishes

Nine polymorphic loci were found among 42 presumptive protein-coding gene loci surveyed among 474 red drum (Sciaenops ocellatus) sampled in 1987 from 13 nearshore and 1 offshore localities from the Atlantic coast of the southeastern USA and the northern Gulf of Mexico. The mean number of alleles over the polymorphic loci was 3.8, and the average heterozygosity over all loci examined was estimated as 0.047. These data indicate that red drum have normal levels of genetic variability. Wright'sF1 _ST values (the standardized variance of allele frequencies between samples) over all polymorphic loci ranged from 0.009 to 0.027 (meanF1 _ST =0.019), and estimates of the effective number of migrants (N _e m) per generation using Wright's island model ranged from 9.0 to 27.5. High levels of gene flow among the red drum samples were also indicated by Slatkin's qualitative analysis using conditional average allele frequencies. Nei's estimates of genetic distance between pairs of samples ranged from 0.000 to 0.009, indicating a high degree of nuclear gene similarity among all samples. Highly significant heterogeneity in allele frequencies at the locus for adenosine deaminase was detected between red drum sampled from the Atlantic and those sampled from the Gulf and among red drum sampled from the Gulf.