Ancient divergences and recent connections in two tropical Atlantic reef fishes<Emphasis Type="Italic"> Epinephelus adscensionis</Emphasis> and<Emphasis Type="Italic"> Rypticus saponaceous</Emphasis> (Percoidei: Serranidae)

Reef habitats of the tropical Atlantic are separated by river outflows and oceanic expanses that may preclude larval dispersal or other population connections in shorefishes. To examine the impact of these habitat discontinuities on the intraspecific phylogeography of reef-associated species we conducted range-wide surveys of two amphi-Atlantic reef fishes that have dispersive pelagic larval stages. Based on 593 bp of mtDNA cytochrome b from the rock hind Epinephelus adscensionis and 682 bp from the greater soapfish Rypticus saponaceous (n=109 and 86, respectively), we found evidence of relatively ancient separations as well as recent surmounting of biogeographic barriers by dispersal or colonization. Rock hind showed slight but significant population genetic differentiation across much of the tropical Atlantic Ocean (_ST=0.056), but deep divergence between the southeastern United States and seven other localities from the Bahamas to the south, central and east Atlantic (mean pairwise d=0.040, overall _ST=0.867). The geographic distribution of the two rock hind lineages is highly unusual in genetic studies of Caribbean Sea reef fishes, because those lineages are separated by less than 250 km of open water within a major biogeographic region. In contrast, highly significant population genetic structure was observed among greater soapfish from the SW Caribbean, Brazil, and mid-Atlantic ridge (_ST=0.372), with a deep evolutionary separation distinguishing putative R. saponaceous from West Africa (mean pairwise d=0.044, overall _ST=0.929). Both species show evidence for a potential connection between the Caribbean and Brazilian provinces. While widespread haplotype sharing in rock hind indicates that larvae of this species cross oceanic expanses of as much as 2000 km, such a situation is difficult to reconcile with the isolation of populations in Florida and the Bahamas separated by only 250 km. These findings indicate that populations of some species in disjunct biogeographic zones may be isolated for long periods, perhaps sufficient for allopatric speciation, but rare gene flow between zones may preclude such evolutionary divergence in other species.Communicated by P.W. Sammarco, Chauvin     

Temperature effects on behavior and survival of marine invertebrates exposed to variations in hydrostatic pressure

Investigation into the pressure resistance of 8 intertidal species belonging to the genera, Uca, Sesurma, Talorchestia and Littorina from the Northern Gulf of Mexico and Panama sheds light on the relationships between temperature and hydrostatic-pressure resistance. Generally, increasing temperature increases the pressure required to elicit reversible reactions such as increased activity and tetany, or paralysis, wheroas increasing temperature generally evokes the irreversible response of death (LD₅₀) at a decreasing pressure. Tropical stenotherms tend to be more sensitive to hydrostatic pressure than eurythermal-temperate species at the same or similar temperatures.     

Nuclear and mitochondrial DNA markers indicate unidirectional gene flow of Indo-Pacific to Atlantic bigeye tuna (<Emphasis Type="Italic">Thunnus obesus</Emphasis>) populations,and their admixture off southern Africa

A sharp genetic break separates Atlantic from Indo-Pacific bigeye tuna (Thunnus obesus) populations, as the frequencies of two major mitochondrial (mt) DNA types ( and ) found in this species are different across the tip of southern Africa. The level of nucleotide divergence between mtDNA types and is of the same order as that between reproductively isolated taxa. To further investigate the genetic structure of bigeye tuna over its distribution range and in the contact zone off southern Africa, bigeye tuna samples collected between 1992 and 2001 (including samples from a previous mtDNA survey) were characterized for four nuclear DNA loci and for mtDNA. Nuclear markers did not support the hypothesis that and mitochondria characterize sibling species. Significant allele-frequency differences at one intronic locus (GH2) and one microsatellite locus (µ208) were found between Atlantic and Indo-Pacific samples, although the level of nuclear genetic differentiation (Weir and Cockerhams =0.025 to 0.042) was much lower than in mtDNA ( =0.664 to 0.807). Probabilistic Bayesian assignment of individuals to a population confirmed that southern African bigeye tuna samples represent a simple mixture of individuals from Atlantic and Indian stocks that do not interbreed, with a higher contribution from Indian Ocean individuals (about 2/3 vs. 1/3).Communicated by M.S. Johnson, Crawley     

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.     

Comparison of allozyme and mitochondrial DNA spatial differentiation in the limpet Patella vulgata

Allozyme and restriction enzyme analysis of mtDNA was used to study variation in samples from British populations of the marine limpet Patella vulgata in two regions. South Wales and Northeast England. Allozyme analysis revealed significant differences in allele frequencies among samples. However F _ST (population differentiation) values were no higher between than within regions, indicating that genetic heterogeneity was localised and not related to geographic separation. For mtDNA, samples from South Wales exhibited higher haplotype diversity values than samples from Northeast England. In addition there were substantial differences in the haplotype distribution between regions. The value of , the haploid analogoue of F _ST , was low within regions (=0.09) but high between regions (=0.44). The estimated difference in migration rate for allozymes and mtDNA exceed the neutral expectation, unless it is assumed that there are influential differences in the magnitude of female and male gametic dispersal.     

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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High levels of genetic subdivision in peripherally isolated populations of the atherinid fish Craterocephalus capreoli in the Houtman Abrolhos Islands,Western Australia

The atherinid fish Craterocephalus capreoli Rendahl is abundant in the Houtman Abrolhos Islands, 70 km off the Western Australia coast and 250 km south of the southern limit of the range of the species along the mainland. Electrophoretic examination of 7 allozyme loci at 17 sites in the Houtman Abrolhos revealed a substantially lower level of polymorphism than found in an earlier study of the species in its mainland distribution, with many of the uncommon alleles and some common ones missing. There is a very high degree of genetic subdivision among the populations in the Houtman Abrolhos, measured by a mean F _ST of 0.437 over a distance of 35 km. This F _ST (standardized variance in allelic frequencies) is six times that found previously among populations along the mainland coast over distances up to 850 km. The subdivision of populations in the Houtman Abrolhos is similar within one island group on a scale up to 12 km, and between two groups that are separated by 15 km of deep water. Significant differences in allelic frequencies were found between populations from the open shore and enclosed lagoons less than 800 m apart, but the overall levels of subdivision were similar for the two types of environment. Previous work had shown high levels of genetic subdivision in the Houtman Abrolhos for a gastropod with direct development. The results for C. capreoli demonstrate that the archipelago favours subdivision even for a species with potentially much greater mobility and different life history.     

Allozyme and mitochondrial DNA evidence of population subdivision in the purple sea urchin Strongylocentrotus purpuratus

Despite high potential for dispersal, the purple sea urchin Strongylocentrotus purpuratus was found to have significant genetic subdivision among locations. Ten geographic locations along the coast of California and Baja California were sampled between 1994 and 1995. Samples from some locations included both adult and recruit urchins. Allozyme analyses revealed a genetic mosaic, where differentiation over short geographic distances could exceed differentiation over much larger distances. Significant allozyme differentiation was found among subpopulations of adults (standardized variance, F _ST =0.033), among subpopulations of recruits (F _ST =0.037), and between adults and recruits from the same location. DNA-sequence data for the mitochondrial cytochrome oxidase I gene also showed significant heterogeneity among locations, with a mild break in haplotype frequencies observed 300 km south of Point Conception. California. Repeated sampling over time is necessary to determine whether these patterns of differentiation are stable and to begin to understand what forces produce them.     

Genetic structure of dictyoceratid sponge populations on the western Coral Sea reefs

Allozyme variation at six polymorphic loci was examined in foliose dictyoceratid sponges from isolated reefs in the western Coral Sea. Four major genetic groups corresponding to the species Phyllospongia lamellosa, P. alcicornis, Carterospongia flabellifera and Collospongia auris were examined. A further two rare morphotypes from individual reefs formed genetic outliers to the P. lamellosa group, and may represent further taxa related to P. lamellosa. Gene frequencies in individual reef populations were largely in Hardy-Weinberg equilibrium, suggesting that random mating occurred in local populations of all four common species. Genetic variability was high and observed heterozygosities within populations ranged from 0.13 to 0.40. All four taxa showed significant genetic differentiation among populations (F _ST=0.05 to 0.36). Genetic distances (Nei's D) among populations within species ranged from 0 to 0.723 and increased with increasing geographical separation. There was evidence that genetic differentiation between populations to the north and to the south of the southern limit of the South Equatorial Current (SEC) divergence was greater than expected on the basis of their geographical separation. The SEC divergence may form a partial barrier to gene flow among populations of these ecologically important sponges on the submerged Queensland Plateau. Levels of migration among populations of three of the species was less than those required to prevent divergence of the populations through genetic drift (Nm<1). Restricted migration among populations may provide a mechanism to explain the occurrence of highly divergent populations of dictyoceratid sponges whose specific identity is not clear, and may allow them additionally to develop partial reproduction isolation from other populations.     

Evolutionary genetics of marine Mysis spp. (Crustacea: Mysidacea)

Inter-and intraspecific allozyme differentiation in the mysid crustacean genus Mysis in the North Atlantic region was studied in order to evaluate earlier concepts of evolutionary and systematic relationships and to assess patterns of subdivision within widespread taxa. The results support a relatively ancient divergence of the marine and non-marine species of the genus, and are generally in line with the current subgeneric tridivision into Mysis s.str., Michteimysis and Auricomysis. However, the North American littoral species M. gaspensis should be returned to subgenus Mysis s.str. from its present position in Michteimysis with M. mixta. The closest observed affinities within Mysis s.str. were between M. gaspensis and the freshwater M. relicta group, and between M. oculata and M. litoralis. Intraspecific differentiation among North European coastal populations of M. oculata and M. litoralis was moderately strong (F _ST0.1), suggesting population bottlenecks and limited dispersal in the post-glacial time. On the other hand, ransoceanic differences were not essentially greater, indicating the systematic homogeneity and long-term dispersal capacity in the marine species. This contrasts with the strong genetic and systematic fragmentation earlier found within the circumboreal M. relicta species group.     

Genetic differentiation of bicolor damselfish (Eupomacentrus partitus) populations in the Florida Keys

Electrophoretic variation in proteins encoded by 23 loci revealed substantial genetic differentiation among populations of bicolor damselfish (Eupomacentrus partitus) collected from four coral reefs in the Florida Keys, USA, during 1986–1988. Genetic differentiation was concentrated between a sample collected from Little Grecian Rocks Reef (LGR) and the remaining samples, including fish from a reef only 600 m distant (Grecian Rocks Reef). Genetic distinction of the LGR sample derived from significantly heterogenous allelic frequencies at six of eight polymorphic loci. Aco-1 (aconitase); Ada (adenosine deaminase); Gpi-2 (glucosephosphate isomerase); Ldh-2 and Ldh-3 (lactate dehydrogenase); and Me-1 (malic enzyme); nevertheless, differentiation at cytosolic aconitase (Aco-1) far exceeded that observed for other loci (fixation index, F _ST=0.482), and differences in Aco-1 allele frequencies were largely responsible for large genetic distances (0.20) between LGR and the other reefs. Paradoxically, estimates of numbers of migrants exchanged between reefs per generation (mN _e=17.47) indicated the potential for extensive gene flow. The extent of genetic differentiation among these populations is evaluated relative to models of population genetic structure based on equilibrium between gene flow and natural selection or genetic drift.     

Genetic differentiation in two sibling species of the brackish-water polychaete Hediste japonica complex (Nereididae)

Genetic divergence among ten populations of small- and large-egg forms of the brackish-water polychaete Hediste japonica complex was investigated on 14 isozyme loci by electrophoretic analysis. The two forms were distinguishable by complete allele substitutions at five loci, resulting in high genetic differentiation (Nei's D: 0.533 to 0.662). No genetic evidence of hybridization between the two forms was detected in sympatric populations in three rivers. These results indicate that the two forms are reproductively isolated, clearly showing that the two forms are distinct species. The genetic differentiation among populations was higher in the large-egg form (D: 0.005 to 0.111, G _ST: 0.435) than that in the small-egg form (D: 0.000 to 0.001, G _ST: 0.020). This genetic difference between the two forms seems to be attributable to a difference in their life histories. The average expected heterozygosity was low in populations of both the large-egg form (0.005 to 0.068) and the small-egg form (0.014 to 0.038) in comparison with other marine invertebrates. Received: 11 April 1997 / Accepted: 8 September 1997     

Sibling species in interstitial flatworms: a case study using<Emphasis Type="Italic"> Monocelis lineata</Emphasis> (Proseriata: Monocelididae)

The genetic relationships between morphologically indistinguishable marine and brackish-water populations of Monocelis lineata (O.F. Müller, 1774) (Proseriata: Monocelididae) were analysed by means of allozyme electrophoresis. Fifteen samples of M. lineata (13 from the Mediterranean and two from the Atlantic) from coastal marine and brackish-water habitats were examined for variation at 18 loci. Eleven loci were polymorphic in at least one population of M. lineata. Low levels of within-population genetic variability were found, with average observed and expected heterozygosity values ranging from H_o=0.015±0.015 to 0.113±0.044, and from H_e=0.028±0.028 to 0.138±0.054, respectively. The occurrence of a number of private alleles indicated a marked genetic divergence among populations of M. lineata, with Rogers genetic distances ranging from D_R=0.003 to 0.676 and a highly significant F_ST value (0.918±0.012, P<0.001). UPGMA (unweighted pair-group method using arithmetic average) cluster analysis and multidimensional scaling showed a clear genetic divergence between marine and brackish-water populations. Moreover, Atlantic and Mediterranean populations were sharply separated. Our results suggest that M. lineata is a complex of sibling species.Communicated by R. Cattaneo-Vietti, Genova     

Zooplankton of the Bristol Channel and Severn Estuary. The distribution of four copepods in relation to salinity

The seasonal variations in distribution and abundance of the common zooplankton species in the Bristol Channel and Severn Estuary were related to the salinity regimes observed over the period November 1973 to February 1975. The dominant constituents in all regions were the calanoid copepods, which reached maximum densities in July: approximately 100 times their winter levels. Four zooplankton assemblages were recognised using an objective classification program which computed similarity coefficients and used group-average sorting. The assemblages existed along the salinity gradient observed from the Severn Estuary to the Celtic Sea. The assemblages were classified as true estuarine, estuarine and marine, euryhaline marine and stenohaline marine and were characterized by the copepods Eurytemora affinis (Poppe) (<30S), Acartia bifilosa var. inermis (rose) (27 to 33.5S), Centropages hamatus (Lilljeborg) (31 to 35S) and Calanus helgolandicus (Claus) (>33S), respectively.     

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     

Resource use by five sympatric parrotfishes in the San Blas Archipelago,Panama

Resource use by five sympatric species of parrotfish was quantified in the San Blas Archipelago of the Republic of Panama from March to August 1987. Detailed observations of parrotfishes on patch reefs and surrounding seagrass beds showed that they partition resources with respect to habitat, food and size, but not time. Although parrotfishes shared resources, the proportions of each resource used differed significantly among species. Scarus iserti (Bloch) scraped filamentous microalgae that grew from eroded coral pavement on lower slopes of patch reefs and in halos, the area of sparse vegetation surrounding reefs. Sparisoma viride (Bonnaterre) foraged on upper slopes of patch reefs where they mostly took bites from dead coral and associated algae. S. aurofrenatum (Cuvier and Valenciennes) had the broadest diet, which consisted mostly of seagrasses and macro- and microalgae that were attached to dead coral on lower reef slopes and in halos. Although S. chrysopterum (Bloch and Schneider) commonly occurred on patch reefs, it primarily foraged in seagrass beds that surround them. S. rubripinne (Cuvier and Valenciennes) was distributed most widely, ranging from seagrass beds to reef crests, where it took bites from seagrasses, dead coral and macroalgae. Juveniles of all species occurred on lower slopes or in halos where they scraped filamentous microalgae from coral pavement. As they matured, parrotfishes moved into other habitats changing access to different types of food. All of these parrotfishes fed throughout the daytime, and resource use did not differ between morning and afternoon.     

Local and regional genetic connectivity in a Caribbean coral reef fish

Coupled bio-physical models of larval dispersal predict that the Costa Rica–Panama (CR–PAN) reefs should constitute a demographically isolated region in the western Caribbean. We tested the hypothesis that CR–PAN coral reef fish populations would be isolated from Mesoamerican Barrier Reef System (MBRS) populations. To test that, we assessed population genetic structure in bicolor damselfish (Stegastes partitus) from both regions. Adult fish were genotyped from five reefs in CR–PAN and from four reefs along the MBRS at 12 microsatellite loci. Between-region F _ST (F _ST = 0.0030, P < 0.005) and exact test (x ² = 74.34, df = 18, P < 0.0001) results indicated that there is weak but significant genetic differentiation between regions, suggesting some restriction in connectivity along the Central American coastline, as predicted by bio-oceanographic models. Additionally, there is among-site genetic structure in the CR–PAN region, relative to the MBRS and between regions, suggesting higher self-recruitment within CR–PAN. This finding may be explained by differences in habitat characteristics.     

Evolutionary genetics of the Mytilus edulis complex in the North Atlantic region

Genetic relationships among Mytilus populations throughout the North Atlantic region, including the Mediterranean and the Baltic Sea, were studied using enzyme electrophoresis. Three distinct groups of populations, each of a remarkably wide distribution, can be recognised on the basis of their multilocus allelic composition: (1) M. galloprovincialis L. of the Mediterranean and western Europe; (2) a genetically distinct form of M. edulis Lmk. from both the Baltic Sea and some localities in the Canadian Maritime Provinces (here provisionally termed the trossulus type mussel); and (3) the traditional Atlantic M. edulis populations of northwestern European coasts and most of eastern North America. These groups are regarded as representing three relatively old evolutionary lineages, which all deserve separate and equal systematic status. The main part of the differentiation at most of the loci studied is accounted for by this major systematic pattern, but considerable geographical differentiation within each of the three principal groups was also detected. At single loci, different electromorphs were found to prevail in disjunct populations of M. galloprovincialis (Mediterranean/Britain) and of the trossulus-type mussel (Baltic/eastern Canada). Within the Atlantic M. edulis, a major part of the differentiation is transoceanic. At one locus (Ap), geographic differentiation appeared to be relatively independent of the systematic boundaries; the possible role of interlineage hybridisation in contact areas in regulating the pattern of geographical variation is discussed.     

Genetic variation and biochemical systematics in the marine bryozoan Alcyonidium mytili

For many years the taxonomic position of the marine ctenostome bryozoan Alcyonidium mytili Dalyell (1848) has been in doubt, with some authorities regarding it as being conspecific with A. polyoum (Hassall). We have set out to resolve this problem using enzyme electrophoresis to examine the genetics of the population structure in the two species. The results show beyond reasonable doubt that sympatric populations of the two species are indeed from quite separate and non-interbreeding gene pools. An examination of various allopatric populations shows a new species of A. mytili to exist sympatrically with the other at Langstone Harbour, Hampshire, England. This species can be readily distinguished on morphological as well as genetic ground. Samples of A. mytili from Guernsey also show substantial genetic differences from other populations examined. It is tentatively concluded that this population also represents a separate gene pool and consequently should be recognised as a third species of A. mytili.     

Does displacement of spawning time occur in the sibling species Cerastoderma edule and C. lamarcki?

The spawning cycles of the sibling species Cerastoderma edule (L.) and C. lamarcki (Reeve) were studied in allopatric and sympatric Danish populations from 1978–1980. The results do not support earlier findings of reproductive character displacement in sympatric populations of the two species. New records of hybrid formation and intermediate forms are presented and discussed.