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.     

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.     

Population genetics of the nurse shark (Ginglymostoma cirratum) in the western Atlantic

The nurse shark, Ginglymostoma cirratum, inhabits shallow, tropical, and subtropical waters in the Atlantic and the eastern Pacific. Unlike many other species of sharks, nurse sharks are remarkably sedentary. We assayed the mitochondrial control region and eight microsatellite loci from individuals collected primarily in the western Atlantic to estimate the degree of population subdivision. Two individuals from the eastern Atlantic and one from the Pacific coast of Panama also were genotyped. Overall, the mtDNA haplotype (h = 48 ± 5%) and nucleotide (π = 0.08 ± 0.06%) diversities were low. The microsatellite data mirror the mitochondrial results with the average number of alleles ([`(N)]<sub>A</sub> \bar{N}_{A}  = 9) and observed heterozygosity ([`(H)]_O \bar{H}_{O}  = 0.58) both low. The low levels of diversity seen in both the mtDNA and the microsatellite may be due to historical sea level fluctuations and concomitant loss of shallow water habitat. Eight of the 10 pair-wise western Atlantic F _ST estimates for mtDNA indicated significant genetic subdivision. Pair-wise F _ST values for the microsatellite loci indicated a similar pattern as the mtDNA. The western Atlantic population of nurse sharks is genetically subdivided with the strongest separation seen between the offshore islands and mainland Brazil, likely due to deep water acting as a barrier to dispersal. The eastern and western Atlantic populations were closely related. The eastern Pacific individual is quite different from Atlantic individuals and may be a cryptic, sister species.     

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.     

Population structure and genetic variation of lane snapper (<Emphasis Type="Italic">Lutjanus synagris</Emphasis>) in the northern Gulf of Mexico

Lane snappers (Lutjanus synagris), sampled from eight localities in the northern Gulf of Mexico (Gulf) and one locality along the Atlantic coast of Florida, were assayed for allelic variation at 14 nuclear-encoded microsatellites and for sequence variation in a 590 base-pair fragment of the mitochondrially encoded ND-4 gene (mtDNA). Significant heterogeneity among the nine localities in both microsatellite allele and genotype distributions and mtDNA haplotype distributions was indicated by exact tests and by analysis of molecular variance (AMOVA). Exact tests between pairs of localities and spatial analysis of molecular variance (SAMOVA) for both microsatellites and mtDNA revealed two genetically distinct groups: a Western Group that included six localities from the northwestern and northcentral Gulf and an Eastern Group that included three localities, one from the west coast of Florida, one from the Florida Keys, and one from the east (Atlantic) coast of Florida. The between-groups component of molecular variance was significant for both microsatellites (Φ _CT = 0.016, P = 0.009) and mtDNA (Φ _CT = 0.208, P = 0.010). Exact tests between pairs of localities within each group and spatial autocorrelation analysis did not reveal genetic heterogeneity or an isolation-by-distance effect among localities within either group. MtDNA haplotype diversity was significantly less (P < 0.0001) in the Western Group than in the Eastern Group; microsatellite allelic richness and gene diversity also were significantly less in the Western Group (P = 0.015 and 0.013, respectively). The difference in genetic variability between the two groups may reflect reduced effective population size in the Western Group and/or asymmetric rates of genetic migration. The relative difference in variability between the two groups was substantially greater in mtDNA and may reflect one or more mtDNA selective sweeps; tests of neutrality of the mtDNA data were consistent with this possibility. Bayesian analysis of genetic demography indicated that both groups have experienced a historical decline in effective population size, with the decline being greater in the Western Group. Maximum-likelihood analysis of microsatellite data indicated significant asymmetry in average, long-term migration rates between the two groups, with roughly twofold greater migration from the Western Group to the Eastern Group. The difference in mtDNA variability and the order-of-magnitude difference in genetic divergence between mtDNA and microsatellites may reflect different demographic events affecting mtDNA disproportionately and/or a sexual and/or spatial bias in gene flow and dispersal. The spatial discontinuity among lane snappers in the region corresponds to a known zone of vicariance in other marine species. The evidence of two genetically distinct groupings (stocks) has implications for management of lane snapper resources in the northern Gulf.     

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     

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.     

DNA sequence analysis identifies genetically distinguishable populations of harp seals (Pagophilus groenlandicus) in the Northwest and Northeast Atlantic

Harp seals (Pagophilus groenlandicus Erxleben, 1777) comprise three populations based upon whelping areas in the Greenland Sea, White Sea, and Northwest Atlantic. The last comprises two subpopulations, one whelping in the Gulf of St. Lawrence (“Gulf ”) and one on the pack ice of the southern Labrador/northern Newfoundland coastal shelf (“Front”). A total of 40 female seals from the four whelping areas were collected during the 1990 and 1992 whelping seasons. DNA sequence variation was examined in a 307 bp region of the mitochondrial cytochrome b gene. Eleven variable nucleotide positions defined 13 genotypes: a significant fraction of the genotypic variance (F _ST=0.12, or 0.09 as measured by Weir's coancestry coefficient θ) is attributable to differentiation between Northwest and Northeast Atlantic populations. There was no significant differentiation between the two whelping areas in the Northwest Atlantic, or between the Greenland Sea and White Sea. These findings suggest significant reproductive isolation exists between trans-Atlantic breeding populations. Received: 18 January 1999 / Accepted: 22 February 2000     

Genetic heterogeneity among blacktip shark,<Emphasis Type="Italic"> Carcharhinus limbatus</Emphasis>, continental nurseries along the U.S. Atlantic and Gulf of Mexico

Genetic population structure of the blacktip shark, Carcharhinus limbatus, a commercially and recreationally important species in the southeast U.S. shark fishery, was investigated using mitochondrial DNA control region sequences. Neonate blacktip sharks were sampled from three nurseries, Pine Island Sound, Terra Ceia Bay, and Yankeetown, along the Gulf of Mexico coast of Florida (Gulf) and one nursery, Bulls Bay, on the Atlantic Ocean coast of South Carolina (Atlantic). Sequencing of the complete mitochondrial control region of 169 neonates revealed 10 polymorphic sites and 13 haplotypes. Overall haplotype diversity and percent nucleotide diversity were 0.710 and 0.106%, respectively. Haplotype frequencies were compared among nurseries to determine if the high mobility and seasonal migrations of adult blacktip sharks have maintained genetic homogeneity among nurseries in the Atlantic and Gulf. Chi-square analysis and AMOVA did not detect significant structuring of haplotypes among the three Gulf nurseries, P(²)=0.294, _ST=–0.005 to –0.002. All pairwise AMOVA between Gulf nurseries and the Atlantic nursery detected significant partitioning of haplotypes between the Gulf and Atlantic (_ST=0.087–0.129, P<0.008), as did comparison between grouped Florida Gulf nurseries and the Atlantic, _CT=0.090, P<0.001. Based upon the dispersal abilities and seasonal migrations of blacktip sharks, these results support the presence of philopatry for nursery areas among female blacktip sharks. Our data also support the treatment of Atlantic and Gulf blacktip shark nursery areas as separate management units.Communicated by P.W.Sammarco, Chauvin     

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.     

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.     

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     

Allozyme and mitochondrial DNA variation in Cuban populations of the shrimp Farfantepenaeus notialis (Crustacea: Decapoda)

We investigated the genetic diversity among populations of the shrimp Farfantepenaeus notialis, the most abundant penaeid species around Cuba. A total of 25 allozyme loci were analyzed in samples of shrimps from seven localities at the south central platform of the island (Ana María Gulf). Samples from three of these localities and from Batabanó Gulf and Guacanayabo Gulf at the south west and south east platforms of the island, respectively, were also characterized at the mtDNA level through sequence variation of a 2027 bp segment including part of the COI and COIII genes. Of the 25 allozyme loci studied 9 were polymorphic: Akp2, Akp3, AmyB, Est3, Gdh, GP7, and Per1, 2 and 3. In contrast to mtDNA, the pattern of allozyme variation among localities revealed strong population structuring at Ana María Gulf, with significant F _st in all pairwise comparisons. The magnitude of F _st estimates as well as the grouping pattern obtained by a UPGMA analysis based on a distance matrix indicated that the level of differentiation was concordant with the geographical position of the localities and the hydrographic regime. Homogeneity of mtDNA suggested that differentiation of allozyme loci might be due to more recent events rather than historical isolation of the sampled populations. Ana María and Guacanayabo Gulf populations were differentiated by mtDNA from Batabanó Gulf, at the southwestern end of the island. The analysis showed three restriction site differences among them, suggesting genetic isolation of the two regions. The present results also suggest that an artificial introduction of larvae from Tunas de Zaza into Batabanó Gulf, in an effort to repopulate this fishing region, may have been ineffective. Received: 13 December 1999 / Accepted: 2 October 2000     

Origin of the antitropical distribution pattern in marine mussels (Mytilus spp.): routes and timing of transequatorial migration

Many marine species, including mussels in the Mytilus edulis species group (i.e. M. edulis L., M. galloprovincialis Lamarck, and M. trossulus Gould), have an antitropical distribution pattern, with closely related taxa occurring in high latitudes of the northern and southern hemispheres but being absent from the tropics. We tested four hypotheses to explain the timing and route of transequatorial migration by species with antitropical distributions. These hypotheses yield different predictions for the phylogenetic relationship of southern hemisphere taxa relative to their northern counter-parts. The three Mytilus species were used to test these hypotheses since they exhibit a typical antitropical distribution and representative taxa occur in both the Pacific and Atlantic. Two types of mtDNA lineages were found among populations of mussels collected from the southern hemisphere between 1988 and 1996; over 90% of the mtDNA lineages formed a distinct subclade which, on average, had 1.4% divergence from haplotypes found exclusively in northern Atlantic populations of M. galloprovincialis. These data indicate that southern hemisphere mussels arose from a migration event from the northern hemisphere during the Pleistocene via an Atlantic route. The remainder of the southern hemisphere lineages (<10%) were very closely related to mtDNA haplotypes found in both M. edulis and M. galloprovincialis in the northern hemisphere, suggesting a second, more recent migration to the southern hemisphere. There was no evidence that southern hemisphere mussels arose from Pacific populations of mussels. Received: 8 December 1998 / Accepted: 8 November 1999     

Molecular genetic variation in tarpon (<Emphasis Type="Italic">Megalops atlanticus</Emphasis> Valenciennes) in the northern Atlantic Ocean

The tarpon (Megalops atlanticus) is a highly valued game fish and occasional food fish in the eastern and western Atlantic Ocean. Tarpon have a high capacity for dispersal, but some regional biological differences have been reported. In this study we used two molecular genetic techniques—protein electrophoresis of nuclear DNA loci, and restriction fragment length polymorphism analysis of the mitochondrial DNA (mtDNA)—to assess this species population genetic structure in the eastern (coastal waters off Gabon and Sierra Leone, Africa) and western (coastal waters off Florida, Caribbean Sea) Atlantic Ocean north of the equator. Genetic differentiation was observed between tarpon from Africa and tarpon from the western Atlantic Ocean. A unique allele and haplotype, significant differences in allozyme allele and mtDNA haplotype frequencies between the African and western Atlantic samples, and significant F_ST analyses suggest that levels of gene flow between tarpon from these two regions is low. Among the western Atlantic Ocean collections, genetic diversity values and allele and haplotype frequencies were similar. AMOVA analyses also showed a degree of genetic relatedness among most of the western Atlantic Ocean collections: however, some significant population structuring was detected in the allozyme data. A regional jackknifed F_ST analysis indicated the distinction of the Costa Rica population from the other western Atlantic populations and, in pairwise analyses, F_ST values tended to be higher (i.e., genetic relatedness was lower) when the Costa Rican sample was paired with any of the other western Atlantic samples. These data suggest that Costa Rican tarpon could be partially isolated from other western Atlantic tarpon populations. Ultimately, international cooperation will be essential in the management of this species in both the eastern and western Atlantic Ocean.Communicated by P.W. Sammarco, Chauvin     

Population divergence in the sinistral whelks of North America,with special reference to the east Florida ecotone

This study evaluated models of species relationships among sinistral whelks in the genus Busycon in the Atlantic and Gulf of Mexico. Gene frequencies at eight polymorphic allozyme loci, shell morphology, anatomy, and partial DNA sequences for the cytochrome c oxidase I (COI) mitochondrial gene were examined in eight populations, ranging from New Jersey to the Yucatan peninsula, and from the dextrally coiled sister taxon Busycon carica (Gmelin, 1791). Whelks were collected in 1997 and 1998. The maximum COI sequence divergence recorded among 32 sinistral individuals was 1.96%, which together with the absence of any gross or qualitative morphological differences, suggested all eight populations should be considered conspecific. High levels of divergence between the allopatric western Atlantic and Gulf of Mexico populations, as revealed by fixed or nearly fixed differences at several allozyme-encoding loci were interpreted as evidence that the east Florida ecotone constitutes a significant barrier to gene flow. Size trimming also revealed several significant quantitative differences in shell and radular morphology between the three pooled Atlantic populations and five pooled Gulf populations. The Yucatan sample was the most distinctive conchologically, with heavy spines and tumid ridges, possibly related to stone crab predation. Based on the evidence all left-handed whelks of North America should be referred to the oldest available nomen, Busycon perversum (Linné, 1758), with three subspecies, B. perversum perversum along the Yucatan peninsula, B. perversum sinistrum (Hollister, 1958) in the northern and eastern Gulf of Mexico, and B. perversum laeostomum (Kent, 1982) in the Atlantic.Communicated by J.P. Grassle, New Brunswick     

Allozyme and mitochondrial DNA variation in orange roughy,Hoplostethus atlanticus (Teleostei: Trachichthyidae): little differentiation between Australian and North Atlantic populations

Allozyme and mitochondrial DNA (mtDNA) genetic variation was compared in orange roughy (Hoplostethus atlanticus Collett) collected from waters off southern Australia and from waters about 22 000 km away in the North Atlantic west of Scotland. Samples were screened for 11 polymorphic allozyme loci and with 9 restriction enzymes. Significant heterogeneity between the two areas was detected for three allozyme loci (ADA ^*, CK ^* and GPI-1 ^*), and the overall G _ST (gene-diversity statistic) value of 1% was small but significant. Significant mtDNA haplotype heterogeneity was observed after ²- of haplotype frequencies but not after a G _ST analysis. Nucleotide sequence-diversity analysis showed very low net divergence (0.0023%) between the two samples. The Australian orange roughy had a lower allozyme heterozygosity and a lower mitochondrial DNA nucleon diversity than the North Atlantic sample. The very limited, although significant, allozyme and mitochondrial DNA heterogeneity between these areas suggests that there is some gene flow between these two populations. The species appears to be widespread, with its presence reported from the southern Pacific, southern Indian, and northern and southern Atlantic Oceans, and it is likely that gene flow between the antipodes is mediated by stepping-stone exchange between adjacent populations rather than by direct migration.     

A comparative seasonal study of two populations of Hypnea musciformis from the East and West Coasts of Florida,USA I. Growth and chemistry

Dry weight levels of the red alga Hypnea musciformis (Wulfen) Lamouroux from Atlantic and Gulf of Mexico coastal sites in florida, USA were lowest in the late winter and early spring, increasing through the summer and highest in the fall. There was a two-month lag in the Gulf coast population's dry weight pattern, indicating differing growth patterns. Chlorophyll a, phycoerythrin and phycoerythrin/chlorophyll a ratios were highest in the winter and lowest in the summer for both populations. Total pigment levels for H. musciformis from the Atlantic coast site were significantly greater than those of the Gulf coast. Protein and carbohydrate percentages were inversely related in both populations, with carbohydrate levels highest in summer and protein levels highest in winter. The Gulf coast population contained significantly more protein than the Atlantic coast plants. Carrageenan levels were highest in spring and lowest in fall, the Atlantic coast population generally had higher levels than the Gulf coast population. The differences in seasonal patterns and levels of the chemical constituents were reflected by distinct morphological characteristics for each population. The Atlantic coast population was larger, darker, more coarse in texture and possessed more crozier branch tips than the Gulf coast plants. These distinctions represent acclimitization responses that relate to habitat differences.     

Understanding and Estimating Effective Population Size for Practical Application in Marine Species Management

Abstract: Effective population size (N_e) determines the strength of genetic drift in a population and has long been recognized as an important parameter for evaluating conservation status and threats to genetic health of populations. Specifically, an estimate of N_e is crucial to management because it integrates genetic effects with the life history of the species, allowing for predictions of a population's current and future viability. Nevertheless, compared with ecological and demographic parameters, N_e has had limited influence on species management, beyond its application in very small populations. Recent developments have substantially improved N_e estimation; however, some obstacles remain for the practical application of N_e estimates. For example, the need to define the spatial and temporal scale of measurement makes the concept complex and sometimes difficult to interpret. We reviewed approaches to estimation of N_e over both long‐term and contemporary time frames, clarifying their interpretations with respect to local populations and the global metapopulation. We describe multiple experimental factors affecting robustness of contemporary N_e estimates and suggest that different sampling designs can be combined to compare largely independent measures of N_e for improved confidence in the result. Large populations with moderate gene flow pose the greatest challenges to robust estimation of contemporary N_e and require careful consideration of sampling and analysis to minimize estimator bias. We emphasize the practical utility of estimating N_e by highlighting its relevance to the adaptive potential of a population and describing applications in management of marine populations, where the focus is not always on critically endangered populations. Two cases discussed include the mechanisms generating N_e estimates many orders of magnitude lower than census N in harvested marine fishes and the predicted reduction in N_e from hatchery‐based population supplementation.     

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.