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.     

Negligible evidence for regional genetic population structure for two shark species <Emphasis Type="Italic">Rhizoprionodon acutus</Emphasis> (Rüppell, 1837) and <Emphasis Type="Italic">Sphyrna lewini</Emphasis> (Griffith &; Smith, 1834) with contrasting biology

Biodiversity of sharks in the tropical Indo-Pacific is high, but species-specific information to assist sustainable resource exploitation is scarce. The null hypothesis of population genetic homogeneity was tested for scalloped hammerhead shark (Sphyrna lewini, n = 237) and the milk shark (Rhizoprionodon acutus, n = 207) from northern and eastern Australia, using nuclear (S. lewini, eight microsatellite loci; R. acutus, six loci) and mitochondrial gene markers (873 base pairs of NADH dehydrogenase subunit 4). We were unable to reject genetic homogeneity for S. lewini, which was as expected based on previous studies of this species. Less expected were similar results for R. acutus, which is more benthic and less vagile than S. lewini. These features are probably driving the genetic break found between Australian and central Indonesian R. acutus (F-statistics; mtDNA, 0.751–0.903, respectively; microsatellite loci, 0.038–0.047 respectively). Our results support the spatially homogeneous monitoring and management plan for shark species in Queensland, Australia.     

Hidden ancient diversification in the circumtropical lancelet Asymmetron lucayanum complex

The tropical lancelet Asymmetron lucayanum (= Epigonichthys lucayanus) is distributed from the western Indian Ocean to the central Pacific Ocean, and the western Atlantic Ocean. Molecular phylogenetic analysis of mitochondrial cytochrome c oxidase subunit I (COI) sequences (1,035 bp) of A. lucayanum (80 specimens from seven localities) showed clearly that this species is genetically distinguished into three major groups of geographical populations based on neighbor-joining tree using maximum likelihood distance (HKY model with invariable sites and gamma correction), suggesting the existence of three cryptic species. Our genetic data show that (1) inter-oceanic divergence time between Clade B (the West-Central Pacific) and Clade C (the Atlantic) (d = 6.6%, ca. 12 million years ago) was smaller than intra-oceanic divergence time between Clade A (the Indo-West Pacific) and Clade B (d=39.5%, ca. 100 million years ago); (2) there are two cryptic species in the West Pacific in sympatry; and (3) high gene flow is implied between the Maldives and the Ryukyus in Clade A (10,000 km distance), the Philippines and Hawaii in Clade B (8,500 km distance), and Barbados and Bermuda in Clade C (2,200 km distance).     

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     

Mitochondrial cytochrome b variation in sleeper sharks (Squaliformes: Somniosidae)

Sleeper sharks are a poorly studied group of deep-sea sharks. The subgenus, Somniosus, contains three morphologically similar species: S. microcephalus found in the Arctic and North Atlantic; S. pacificus in the North Pacific; and S. antarcticus in the Southern Ocean. These sharks have been reported mainly in temperate to polar waters and occasionally in subtropical locations. They have not been recorded in tropical waters. This study investigates the relationships among the accepted species of Somniosus through analysis of mitochondrial cytochrome b sequence variation. Seventy-five samples were examined from four sampling locations in the North Pacific, Southern Ocean and North Atlantic. Twenty-one haplotypes were found. A minimum spanning parsimony network separated these haplotypes into two divergent clades, an S. microcephalus and an S. pacificus/antarcticus clade, strongly supporting the distinction of S. microcephalus as a separate species from the Pacific sleeper shark species. Analysis of genetic structure within the S. pacificus/antarcticus clade (analysis of molecular variance, allele frequency comparisons, and a nested clade analysis) showed limited or no differences amongst three populations. Further examination of genetic variation at more variable mtDNA and nuclear markers is needed to examine the species status of S. pacificus and S. antarcticus.     

Hidden genetic diversity in a key model species of coral

A total of 43 colonies of the scleractinian coral Pocillopora damicornis from lagoonal and reef slope sites in the western Indian Ocean (WIO) region were genetically characterised at one nuclear and two mitochondrial sequence markers and six microsatellite loci. Both mitochondrial and microsatellite data support the existence of two reciprocally monophyletic clusters (F- and NF-types) and provide evidence of the existence of two cryptic species of P. damicornis on reefs in WIO region and put current morphological delineation and geographical boundaries of P. damicornis and Pocillopora molokensis into question. The results add to ongoing studies on the phylogeny and phylogeography within the genus Pocillopora, which all point towards a range of unresolved morphological and molecular species boundaries. Nuclear phylogenies derived from the present and previously published sequences show evidence for incomplete lineage sorting and/or introgressive hybridisation between Pocillopora morphospecies. However, the two WIO types largely remain in separate clusters, further supporting the theory that these represent two different species.     

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.     

Multiple trans-Arctic passages in the red alga Phycodrys rubens: evidence from nuclear rDNA ITS sequences

In order to investigate how episodes of geological and climatic change have influenced the distribution and evolutionary diversification of Arctic to cold temperate-North Atlantic seaweed species, intraspecific genetic variation was analyzed among isolates of the sublittoral, benthic red alga Phycodrys rubens (collected between June 1992 and January 1994). Rooted phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer (ITS) sequences and the plastid encoded Rubisco spacer sequences suggest that P. rubens invaded the North Atlantic from the Pacific shortly after the opening of the Bering Strait (3 to 3.5 million years ago), colonizing both the western and eastern Atlantic coasts. Based on these data we further hypothesize that P. rubens survived along the European coasts during the more recent Pleistocene glaciations, while becoming locally extinct along the North American Atlantic coasts. Following retraction of the last ice sheet, the western Atlantic coast was colonized a second time from the Pacific. The presence of two distinct genetic types (based on ITS and Rubisco sequences) along the European coasts is postulated to be a result of isolation and subsequent differentiation. This is likely because ice-free areas are known to have existed in northern Scotland and Norway during the last glaciation. The presence of an East Atlantic genetic type along the West Atlantic coast is believed to be a recent introduction (caused by human activity) of P. rubens to Newfoundland.     

Phylogeography of the ocean quahog (Arctica islandica): influences of paleoclimate on genetic diversity and species range

The ocean quahog, Arctica islandica (Linnaeus, 1767), is a commercially important bivalve found on continental shelves throughout much of the North Atlantic. To assess genetic subdivision in this species, we sequenced 385 nucleotides of the mitochondrial cytochrome b (cyt b) gene from 83 specimens collected from 12 localities between September 1998 and July 1999 (based on preliminary data, the Internal Transcribed Spacers, ITS, of the nuclear ribosomal repeat were not useful). The cyt b data delimited 11 haplotypes with 0.26 to 8.1% nucleotide difference (coded by 36 variable nucleotide positions) among them. Only three haplotypes were detected in 39 specimens collected along the USA coastline, compared to five haplotypes from nine Icelandic individuals. The western Atlantic populations ranging from Penobscot Bay (Maine, USA) to southern Virginia showed relatively low diversity and appeared genetically similar in that region. Based on the presence of shared haplotypes, AMOVA analyses, and phylogenetic reconstructions, Icelandic populations appear to be more genetically similar to western Atlantic populations than eastern Atlantic populations. Specimens from the Faroe Islands (n=4) show mixed affinities. These data are consistent with the hypothesis that a warm Holocene climatic optimum (ca. 7,500 years BP), and not glacial refugia, shaped the present-day genetic structure in A. islandica. Received: 18 January 2000 / Accepted: 26 June 2000     

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

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

Phylogeography of the marine interstitial nemertean <Emphasis Type="Italic">Ototyphlonemertes parmula</Emphasis> (Nemertea,Hoplonemertea) reveals cryptic diversity and high dispersal potential

We conducted a phylogeographic study of the meiofaunal nemertean Ototyphlonemertes parmula, an apparent species complex from the littoral zone of coarse-grained beaches, using a 494-bp fragment of the mitochondrial cytochrome oxidase 3 gene (cox3). Six populations from the Gulf and Atlantic coasts of Florida, two from New England, and one from the Caribbean were sampled in March and August 2005. Three major lineages were identified, separated by cox3 sequence divergence of 16–18%, with partially overlapping ranges. Tests for hybridization using ISSR markers detected nuclear gene exchange within but not between the major mitochondrial lineages, indicating the presence of cryptic species. One lineage dominating the Atlantic coast of Florida shows no evidence of geographic structuring. Another lineage shows a phylogenetic break between the Atlantic and Gulf coasts, suggesting that unsuitable habitat may act as a barrier to dispersal. Long-distance migration is evidenced by shared haplotypes between Florida and the eastern Caribbean. Overall, the widespread distribution of individual haplotypes and lack of structuring within geographic regions contrast with O. parmula’s strongly sediment-bound lifestyle. We speculate that dispersal of adults by storms and/or sediment transport may be more important than few and potentially short-lived planktonic larvae to explain geographic diversity in O. parmula and may be important for meiofauna in general.     

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 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     

Deconstructing an assemblage of “turtle” barnacles: species assignments and fickle fidelity in Chelonibia

Barnacles in the genus Chelonibia are commensal with a variety of motile marine animals including sea turtles, crustaceans, and sirenians. We conducted a worldwide molecular phylogenetic survey of Chelonibia collected from nearly all known hosts to assess species relationships, host-fidelity, and phylogeographic structure. Using DNA sequences from a protein-coding mitochondrial gene (COI), a mitochondrial rRNA gene (12S), and one nuclear rRNA gene (28S), we found that of four species, three (C. testudinaria, C. patula, and C. manati) are genetically indistinguishable. In addition, we show each utilizes a rare androdioecious mode of reproduction involving complemental males. In contrast, the fourth species (C. caretta), which is hermaphroditic and specializes on turtles, is genetically distinct—leading to the conclusion that the three former taxa are morphotypes of the same species and should be synonymized under C. testudinaria. Phylogenetic analysis resulted in three geographic clades (Atlantic, Indian Ocean/western Pacific, and eastern Pacific) with haplotype parsimony networks revealing no shared haplotypes among geographic regions. Analysis of molecular variance detected significant differences among sequences by region (p < 0.005); conversely, there were no significant differences among sequences when grouped by host or taxonomic designation. Average pairwise genetic distances were lower between the eastern Pacific and Atlantic clades (0.053 ± 0.006) than between the eastern Pacific and Indian Ocean/western Pacific clades (0.073 ± 0.008), suggesting Atlantic and eastern Pacific populations were connected more recently, perhaps until the rise of the Isthmus of Panama. Host use by Chelonibia morphotypes is discussed along with speculation on possible ancestral hosts and support for a “turtle-first” hypothesis.     

Geographic and bathymetric patterns of mitochondrial 16S rRNA sequence divergence among deep-sea amphipods,Eurythenes gryllus

The physical uniformity of the deep sea suggests a lack of absolute barriers to faunal dispersal, and thus genetic homogeneity in broadly distributed species is expected. The deep-sea amphipod Eurythenes gryllus Lichtenstein (Crustacea: Lysianassoidea) is considered a panoceanic, cold-water stenotherm, with a vertical depth distribution from 184 to 6500 m. We surveyed mtDNA sequence diversity in E. gryllus to assess genetic diversity and population structure in different oceans and across traditionally defined bathyal and abyssal zones. DNA sequences (437 nucleotides length) from the mitochondrial large-subunit ribosomal RNA gene (16S rRNA) of 95 individuals, collected between 1982 and 1990 from 14 locations in the central North Pacific (including multiple samples on the slope of a seamount), castern and western North Atlantic, and the Arctic Ocean, were obtained. Our analysis of DNA sequence diversity indicates (1) genetic homogeneity among sites within the same depth zone at the scale of ocean basins; and (2) genetically divergent, cryptic taxa distributed at different depths, with the greatest diversity in the bathyal zone. These observations suggest that ecological and physical conditions are important isolating mechanisms that may lead to speciation in this group.     

Molecular population genetics of male and female mitochondrial genomes in European mussels <Emphasis Type="Italic">Mytilus</Emphasis>

The doubly uniparental system of mitochondrial inheritance (DUI) is best known in marine mussels Mytilus. Under DUI there are two types of mitochondrial DNA (mtDNA). The female type (F) is transmitted to offspring of both genders and the male type (M) exclusively to sons; consequently two distinct mtDNA lineages exist. The M lineage evolves under more relaxed selection than the F lineage resulting in higher polymorphism within the M lineage. Though this polymorphism is expected to make inferences on fine population structure easier using M instead of F data, no comprehensive comparative data exist to support this claim. We sequenced a 1,205 bp fragment of M and F mtDNA comprising parts of the COIII and ND2 genes, and analysed 204 individuals representing three Mytilus species: M. edulis, M. galloprovincialis and M. trossulus from 13 European sampling sites. A clear distinction between Mediterranean and Atlantic populations was found with both M and F data, but much better geographic differentiation was found within the Atlantic using F rather than M data. In particular, Atlantic M. galloprovincialis can be differentiated from Atlantic M. edulis, and further subdivision of Atlantic M. edulis is possible using the F data but not the M data. Multiple tests of selection were carried out to attempt to explain this paradox. We concluded that the overall pattern of polymorphism is consistent with strong purifying selection; not only is this selection relaxed in the M lineage in comparison with the F lineage, but it is also more frequently interrupted by periodic selective sweeps within the M lineage.     

Sympatric size variants of the microcopepod Oncaea venusta exhibit distinct lineages in DNA sequences

We investigated the phylogenetic relationships among different size groups of the pelagic microcopepod Oncaea venusta Philippi, 1843, by comparing the patterns of genetic variation of specimens collected at five locations of the Indo-West Pacific Ocean. Phylogenetic analyses were based on sequence data obtained from two DNA markers: A 310 bp fragment of the mitochondrial cytochrome b (cyt b) gene and a 480 bp fragment of the nuclear internal transcribed spacer 1 (ITS1). The cyt b sequences showed a much higher level of variation than those from ITS1, but the conclusion from both genes was concordant. Four genetic clades could be differentiated. A small- and a large-size group were unambiguously assigned to two distinct clades or lineages. Unexpectedly, the medium-sized individuals could be divided into another two different genetic clades. All four lineages were supported by high bootstrap values. The high levels of sequence divergence under sympatric conditions indicated that at least the two main groups, the large and the small one, may be assigned to different species. For the medium-size group additional morphological studies and more sensitive nuclear markers are required to clarify their taxonomic status.     

Poor-data and data-poor species stock assessment using a Bayesian hierarchical approach

Appropriate inference for stocks or species with low-quality data (poor data) or limited data (data poor) is extremely important. Hierarchical Bayesian methods are especially applicable to small-area, small-sample-size estimation problems because they allow poor-data species to borrow strength from species with good-quality data. We used a hammerhead shark complex as an example to investigate the advantages of using hierarchical Bayesian models in assessing the status of poor-data and data-poor exploited species. The hammerhead shark complex (Sphyrna spp.) along the Atlantic and Gulf of Mexico coasts of the United States is composed of three species: the scalloped hammerhead (S. lewini), the great hammerhead (S. mokarran), and the smooth hammerhead (S. zygaena) sharks. The scalloped hammerhead comprises 70-80% of the catch and has catch and relative abundance data of good quality, whereas great and smooth hammerheads have relative abundance indices that are both limited and of low quality presumably because of low stock density and limited sampling. Four hierarchical Bayesian state-space surplus production models were developed to simulate variability in population growth rates, carrying capacity, and catchability of the species. The results from the hierarchical Bayesian models were considerably more robust than those of the nonhierarchical models. The hierarchical Bayesian approach represents an intermediate strategy between traditional models that assume different population parameters for each species and those that assume all species share identical parameters. Use of the hierarchical Bayesian approach is suggested for future hammerhead shark stock assessments and for modeling fish complexes with species-specific data, because the poor-data species can borrow strength from the species with good data, making the estimation more stable and robust.     

Close genetic similarity of Atlantic and Pacific skipjack tuna (Katsuwonus pelamis) demonstrated with restriction endonuclease analysis of mitochondrial DNA

Restriction endonuclease analysis of mitochondrial DNA indicated a surprisingly high degree of genetic similarity between skipjack tuna (Katsuwonus pelamis) from the Atlantic and Pacific Oceans. The present results (1983) support the findings of previous morphological and electrophoretic studies. Evidently, since the uplift of the Panamá land bridge about 3.1 million years ago, there has been continued genetic contact between Atlantic and Pacific skipjack tuna, presumably via the Southern Ocean.     

Cryptic species and population structuring of the Atlantic and Pacific seabob shrimp species, Xiphopenaeus kroyeri and Xiphopenaeus riveti

Seabob shrimps of the genus Xiphopenaeus are important fishery resources along the Atlantic and Pacific coasts of Central and South America. The genus was considered to comprise two species: the Atlantic Xiphopenaeus kroyeri (Heller, Sitzungsber Math Naturwiss cl kaiserliche Akad Wiss Wien 45:389–426, 1862), and the Pacific Xiphopenaeus riveti (Bouvier, Bull Mus Hist Nat Paris 13:113–116, 1907). In a recent review, Xiphopenaeus was regarded as a monotypic genus, on the basis that no clear morphological differences could be found between Pacific and Atlantic specimens (Pérez Farfante and Kensley, Mem Mus Nat Hist Nat Paris 175:1–79, 1997). In the present work, nuclear (allozymes), and mitochondrial (Cytochrome Oxidase I) genes were used to demonstrate the validity of X. riveti and reveal the presence of two cryptic species of Xiphopenaeus within X. kroyeri in the Atlantic Ocean. The high levels of molecular divergence among these species contrast with their high morphological resemblance. Interspecific sequence divergences (Kimura 2-parameter distance) varied from 0.106 to 0.151, whereas intraspecific distances ranged from 0 to 0.008 in Xiphopenaeus sp. 1, from 0 to 0.003 in Xiphopenaeus sp. 2, and from 0.002 to 0.005 in X. riveti. In addition, five diagnostic allozyme loci were found between sympatric samples of Xiphopenaeus sp. 1 and 2 along the Brazilian coast. The results suggest that Xiphopenaeus sp. 2 from the Atlantic is more closely related to the Pacific X. riveti than to the Atlantic Xiphopenaeus sp. 1. Furthermore, a high level of genetic structuring (Xiphopenaeus sp. 1: F _ST =0.026; P<0.05; Xiphopenaeus sp. 2: F _ST =0.055; P<0.01) was found in the Brazilian Xiphopenaeus populations, indicating the presence of different genetic stocks in both Atlantic species. These findings have important commercial implications as they show that the fisheries of the two Atlantic species must be managed separately, and that each one is comprised of different populations.Communicated by O. Kinne, Oldendorf/Luhe