Genetic relationships of pilchards (genus: Sardinops) with anti-tropical distributions

The pilchards (genus Sardinops), one of the major components of the world's harvest of fishes, is distributed sporadically in the temperate zone on a global scale. Mitochondrial DNA variation in 95 pilchards from nine worldwide localities in five current systems was examined using ten restriction enzymes. Although several opinions have been proposed on the global population structure of the genus Sardinops, our results clearly rejected the hypothesis of gene flow among geographically distant populations on a global scale. Results indicate that the Japanese pilchards and the Australian and South African pilchards were derived from the eastern Pacific populations, following the speciation of Sardinops in the Pacific, after the formation of the Isthmus of Panama. An intermixing of the northeastern and southeastern Pacific populations during the Pleistocene glacial maxima is suggested from their genetic affinities.     

Population genetics of the fissiparous holothurians Stichopus chloronotus and Holothuria atra (Aspidochirotida): a comparison between the Torres Strait and La Réunion

Collections of about 50 individuals from each of five populations of the fissiparous holothurian species Stichopus chloronotus and four populations of Holothuria atra were made in 1999. These populations were located in the Torres Strait (western Pacific) and La Réunion (western Indian Ocean). Allozyme electrophoretic surveys of five (S. chloronotus) and six (H. atra) loci were conducted to compare patterns of asexual reproduction and to investigate connectivity between regions separated by large geographic distances. Deviations from genotype frequencies expected under Hardy-Weinberg equilibrium, mostly heterozygote excesses, were observed in all populations of both species. The maximum contribution of sexual reproduction (calculated as the maximum number of sexually produced individuals: sample size=N*/N_i) was similar for all S. chloronotus (58-64%) and H. atra (76-92%) populations, and on the same level as previously reported for midshelf reefs of the Great Barrier Reef. The higher values in the latter species indicated greater contributions of asexual reproduction to S. chloronotus populations. Variability was strongly reduced in S. chloronotus populations at La Réunion, with only one locus being variable in that population. When the dataset was reduced to one representative per multi-locus genotype per population to reduce the effect of asexual reproduction on calculations on gene flow, F_ST values were not significantly different from zero, suggesting high gene flow between these regions. However UPGMA cluster analyses using Rogers' genetic distance, roughly clustered populations by region. In the case of H. atra, pooled populations within each region were significantly different from those of the other region. Thus, although some restrictions in gene flow and greater genetic distances between the regions may exist, those differences are distinctly less than those reported in previous studies on echinoderms over similar geographic scales. Despite the importance of asexual reproduction for the maintenance of local population size, this study also confirmed that the potential for widespread dispersal mediated by sexually produced larvae is large.     

Phylogenetic relationships of deep-sea mussels of the genus<Emphasis Type="Italic"> Bathymodiolus</Emphasis> (Bivalvia: Mytilidae)

We examined phylogenetic relationships among three Bathymodiolus species in Japanese waters and Bathymodiolus spp. from the Manus Basin by two different approaches. Two-dimensional gel electrophoresis allowed us to compare 263–407 (average=318) proteins, giving comprehensive information on genetic distances among the species. The neighbor-joining tree presented two clusters: (1) B. japonicus and B. platifrons and (2) B. septemdierum and B. sp. Members of the first cluster contain methanotrophic endosymbiotic bacteria and members of the second cluster contain thioautotrophic endosymbionts. DNA sequencing of a fragment (415 bp) of mitochondrial cytochrome c oxidase subunit I (COI) provided a neighbor-joining tree with the same topology as that derived from protein analysis. Inspection of intraspecific variation in COI in B. japonicus and B. platifrons revealed no genetic differentiation between mussel populations of either species from cold-water seeps versus hydrothermal vents, suggesting high adaptability of these Bathymodiolus species to deep-sea chemosynthetic environments. Our results indicated genetic exchanges between mussels from distant localities, suggesting that a limited dispersal capability of the larvae is not the likely factor leading to speciation events in these Bathymodiolus species.Communicated by T. Ikeda, Hakodate     

Minimal genetic variation among samples of six species of coral reef fishes collected at La Parguera,Puerto Rico,and Discovery Bay,Jamaica

Intraspecific genetic variation among samples of six species of reef fishes,Chromis cyanea, Stegastes partitus, S. planifrons, S. leucostictus, S. dorsopunicans, andThalassoma bifasciatum collected over a 2 wk period in 1990 at La Parguera, Puerto Rico, USA and Discovery Bay, Jamaica, was evaluated using starch-gel electrophoresis. On average, products of 33 protein-coding loci were resolved in each species. Levels of polymorphism (0.95 criterion) ranged from 3.1% inS. dorsopunicans to 42.4% inC. cyanea. Estimates of genetic divergence among samples and indices of genetic subdivision were small in all six study species: mean genetic distances ranged from 0.000 to 0.002 and mean fixation indices ranged from 0.004 to 0.035. Estimates of numbers of migrants per generation (mN _e) ranged from 5.1 to 11.6, indicating that substantial genetic exchange probably occurs over the relatively large geographic distance (ca. 1000 km) separating coral reef communities of La Parguera and Discovery Bay. The estimates ofmN _e may be biased by a sampling strategy involving only two localities, and should therefore be interpreted with caution. With inferences based solely on allozyme frequency data under a primary assumption of neutrality, genetic substructuring of populations of the six study species on a macrogeographic scale appears virtually nonexistent.     

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

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

Genetic diversity and genetic structure in the brown alga Halidrys dioica (Fucales: Cystoseiraceae) in Southern California

Genetic diversity and genetic structure in a population of the brown seaweed Halidrys dioica Gardner were evaluated in five sites in southern California, USA, in 1991, using isozyme electrophoresis. H. dioica is relatively long-lived and has an outcrossing mating system and floating reproductive fronds with the potential for longdistance gamete dispersal. Because these characteristics are hypothetically important in determining genetic diversity and structure, we predicted that genetic diversity would be high and genetic structure would be exhibited only at relatively large geographic scales in H. dioica populations. The data were consistent with the prediction: genetic diversity (% polymorphic loci, no. of alleles/locus, average expected heterozygosity) was high compared to that of other seaweed species. Genetic structure (Wright's F statistics, Nei's genetic distance, point-pattern analysis of alleles) was low within and among distinct rocky reefs over 4 km of coast but high in subpopulations separated by 90 km. Life-history characteristics may be useful predictors of genetic diversity and structure in seaweed populations, but information on selection regimes, long-distance dispersal, and the extent of clonal propagation, for example, are critically lacking.     

Genetic structure of crown-of-thorns starfish (Acanthaster planci) in Australia

Significant genetic structuring is reported for crown-of-thorns starfish Acanthaster planci (L.) populations within Australia, sampled in 1987 and 1989, with Eastern Australian and Western Australian populations forming different groups. Eastern Australian populations sampled included the three most southerly, on isolated reefs of Elizabeth and Middleton Reefs, and Lord Howe Island. The Lord Howe Island population formed a discrete outlier from all other Australian populations, with average genetic distances (Nei's unbiased D) from other populations of 0.05 compared with 0.01 among all other populations. This is greater than the levels of genetic differentiation described for A. planci from sites at the extremes of the Pacific. The genetic distinction of Western Australia and Lord Howe Island is considered to reflect restricted dispersal as a result of current patterns. The Western Australian populations also showed less genetic diversity than the Great Barrier Reef populations, and this might reflect a founder effect or recent bottleneck in the Western Australian populations.Contribution No. 554 from the Australian Institute of Marine Science     

Genetic population structure in two tropical sponge-dwelling shrimps that differ in dispersal potential

The spatial context in which host races of parasitic animals originate is a central issue in the controversial theory of sympatric speciation. Sponge-dwelling shrimps in the genus Synalpheus provide a good system for evaluating the possibility of resource-associated divergence in sympatry. I used allozyme electrophoresis to assess the genetic population structure of two Caribbean Synalpheus species sampled in 1988 to 1990 at a hierarchy of spatial scales. S. brooksi Coutière is a host-generalist, using several sponge species in an area, and develops directly, with no planktonic larval stage. G-tests and estimates of F _ST revealed highly structured populations in this species, with significant differentiation among samples from individual reefs within a region, and strong divergence among regions (Panama, Belize, Florida). Moreover, samples of S. brooksi taken from the two sponges Spheciospongia vesparium (Lamarck) and Agelas clathrodes (Schmidt) in Panama, and separated by 3 km, showed significant differentiation at both of the loci that were polymorphic in these populations. Genetic distances between these host-associated populations averaged >60% greater than distances between samples from the same host species and were comparable to, or greater tha, those for some inter-regional comparisons. These genetic data corroborate a previous finding of demographic differences between the same populations. The second species, S. pectiniger Coutière, occurs only in Spheciospongia vesparium, and produces swimming larvae. Although allele frequencies in this species differed significantly among the three regions, S. pectiniger showed no differentiation within regions, and significantly lower differentiation (F _ST) among regions than its direct-developing congener. These data suggest that genetic population structure in these two commensal crustaceans is related to dispersal potential, and that restricted dispersal may allow the divergence of host-associated populations on a local scale.     

Genetic structure,lack of sex-biased dispersal and behavioral flexibility in the pair-living fat-tailed dwarf lemur, <Emphasis Type="Italic">Cheirogaleus medius</Emphasis>

Mating system and dispersal patterns influence the spatio-genetic structure within and between populations. Among mammals, monogamy is rare, and its socio-genetic consequences have not been studied in detail before. The goal of our study was to investigate population history, demographic structure, and dispersal patterns in a population of pair-living fat-tailed dwarf lemurs, Cheirogaleus medius, a small, nocturnal primate from western Madagascar, and to infer their underlying behavioral mechanisms. Tissue samples for DNA extraction were obtained from a total of 140 individuals that were captured in two subpopulations about 3 km apart. Analyses of mtDNA variability at the population level revealed very low levels of genetic variability combined with high haplotype diversity, which is indicative of a recent population bottleneck. We found no evidence for spatial clustering of same-sexed individuals with identical haplotypes within each of two subpopulations but significant clustering between them. Thus, a high level of local subpopulation differentiation was observed (F _ST = 0.230). The sexes showed equal variances in the number of individuals representing each haplotype, as well as equal levels of aggregation of identical haplotypes. Hence, both sexes disperse from their natal area, one pattern expected in a pair-living mammal. There is a possibility of behavioral and social flexibility in this species, however, because we documented pronounced differences in density and sex ratio between the two subpopulations, suggesting that single study sites or populations may not be representative of a given local population or even species.     

Cryptic species of Clavelina (Ascidiacea) in two different habitats: harbours and rocky littoral zones in the northwestern Mediterranean

Marinas and harbours provide ideal sites for the study of population genetics of marine invertebrates with restricted dispersal capabilities. They combine a confinement effect, particular ecological conditions (pollution, turbidity), and the possibility of high gene flow through ship-borne propagules, which greatly increases the natural dispersal capability of sexual and asexual propagules in many species with short-lived larvae. We studied the genetic structure of populations of the ascidian Clavelina lepadiformis living inside and outside harbours in the north-western Mediterranean. A 500-bp segment of the cytochrome c oxidase subunit I (COI) mitochondrial gene was sequenced in three populations from inside harbours (interior form) and in three populations from the rocky littoral (exterior form). Two congeneric Mediterranean species, Clavelina sp. and C. dellavallei, were used for comparison. We found that the interior and exterior forms of C. lepadiformis belong to two distinct clades, with a genetic divergence of 5%. Gene-flow values among these forms were insignificant. The lack of gene flow and the genetic divergence suggest that the interior and exterior forms of C. lepadiformis are in fact cryptic species rather than differentiated populations of the same species. Levels of gene flow were higher among interior habitats than among exterior habitats, a pattern likely maintained by genetic exchange through ships. We discuss the possible origins of the present-day distribution of these cryptic species. We contend that the study of species living both inside and outside these particular habitats will reveal more instances of genetic discontinuities allowing local adaptations.     

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     

Co-existing populations of Pacific ocean perch, Sebastes alutus, in Queen Charlotte Sound, British Columbia

Variation at five microsatellite loci (Sal1, Sal2, Sal3, Sal4 and Sal5) was examined in approximately 1300 Pacific ocean perch (Sebastes alutus) sampled from 14 coastal sites in British Columbia, Canada. Mean observed heterozygosities by locus ranged from 71% to 88%, and by sample ranged from 75% to 84%. Theta values ranged from 0 to 0.04 over the five loci, and averaged 0.015. Among Pacific ocean perch samples, Š ranged from 0.001 to 0.056. Canonical discriminate analysis of multilocus genotypes and neighbour-joining analysis of pairwise genetic distances between samples both indicated the presence of three populations, one off the west coast of Vancouver Island (the Vancouver Island population) and two co-existing populations in Queen Charlotte Sound, Dixon Entrance and along the west coast of the Queen Charlotte Islands (the eastern and western QCI populations). Pacific ocean perch of the eastern and western QCI populations were caught in close proximity to each other, but individual samples showed little evidence of admixture. Fall and spring samples collected within geographic areas were genetically similar, indicating seasonally stable population structure. Restricted gene flow between the Vancouver Island and the two more northerly populations may result from limited adult dispersal and larval retention within the California Current and Alaska Gyre, respectively, but the presence of two populations within Queen Charlotte Sound cannot be explained entirely by larval retention hypotheses. The presence of two Pacific ocean perch populations in central British Columbia has implications for fisheries management.     

Genetic population structure and sociogenetic organisation in Formica truncorum (Hymenoptera; Formicidae)

Summary The genetic population structure and the sociogenetic organization of the red wood ant Formica truncorum were compared in two populations with monogynous colonies and two populations with polygynous colonies. The genetic population structure was analysed by measuring allele frequency differences among local subsets of the main study populations. The analysis of sociogenetic organisation included estimates of nestmate queen and nestmate worker relatedness, effective number of queens, effective number of matings per queen, relatedness among male mates of nestmate queens and relatedness between queens and their male mates. The monogynous populations showed no differentiation between subpopulations, whereas there were significant allele frequency differences among the subpopulations in the polygynous population. Workers, queens and males showed the same genetical population structure. The relatedness among nestmate workers and among nestmate queens was identical in the polygynous societies. In three of the four populations there was a significant heterozygote excess among queens. The queens were related to their male mates in the polygynous population analysed, but not in the monogynous ones. The data suggest limited dispersal and partial intranidal mating in the populations with polygynous colonies and outbreeding in the populations having monogynous colonies. Polyandry was common in both population types; about 50% of the females had mated at least twice. The males contributed unequally to the progeny, one male fathering on average 75% of the offspring with double mating and 45–80% with three or more matings. Correspondence to: L. Sundström     

Genetic differentiation in Sargassum polyceratium (Fucales: Phaeophyceae) around the island of Curaçao (Netherlands Antilles)

The seaweed Sargassum polyceratium Montagne inhabits a broad spectrum of subtidal and intertidal habitats. Genetic diversity and spatial genetic structure were examined within and among 12 stands using random amplified polymorphic DNA (RAPD) phenotypes. Data were analyzed using analysis of molecular variance (AMOVA) and Shannon's information measure. In both analyses, 60-75% of the variation occurred within stands and 25-40% between stands. These values are consistent with out-crossing, high-dispersal species. Significant differentiation was found among bays ca. 25 km apart (Shannon's G'_st averaged 0.37 and pairwise AMOVA K_st values averaged 0.272) and among stands 150-200 m apart within bays (AMOVA K_st values averaged 0.149). Effects of shore (windward vs. leeward), depth, and bay on population structure were tested. These analyses revealed that the factor depth is confounded with shore, and that bays show significant differentiation from each other but are not completely isolated from one another. Mantel tests for differentiation-by-distance were significant along both sides of the island but stronger along the windward side. A neighbor-joining analysis of genetic distances among stands showed that the effects of currents around both tips of the island were especially important for shallow populations. For S. polyceratium, depth and bay promote population differentiation along shores, yet dispersal around the tips of the island simultaneously connects these populations to varying degrees. This study highlights the importance of investigating the relative contribution of habitat factorsin relation to island-scale population structure.     

Genetic differentiation in Littorina saxatilis (Gastropoda)

Littorina saxatilis, an ovoviviparous gastropod, displays considerable morphological variability, perhaps due to limited dispersal abilities. Gene and genotype frequencies of 2 polymorphic enzymatic loci were determined electrophoretically for 7 populations of L. saxatilis, to estimate degree of population differentiation. Results show that significant population differentiation may occur over distances of as little as 2 km, while widely separated populations may be nearly identical. An index of probability of genotypic identity calculated for populations of L. saxatilis shows variable and heterogeneous values. Comparison of this index for L. saxatilis with an index for 12 populations of Nassarius obsoletus —which possesses strong dispersal abilities due to long-term veliger larva and high, homogeneous indices of probability of genotypic identity between populations — indicates a strong correlation between population differentiation and dispersal ability of the organism. Possible factors influencing population differentiation in L. saxatilis are discussed.     

Genetic structure of populations of two closely related brittle stars with contrasting sexual and asexual life histories,with observations on the genetic structure of a second asexual species

Ophiocoma pumila Lütken andOphiocomella ophiactoides (H. L. Clark) are morphologically similar brittle stars with contrasting life histories, the former obligately sexual, the latter fissiparous (capable of both sexual reproduction and asexual proliferation by binary fission). Electrophoretic analysis of five polymorphic enzymes was used to assess the genetic consequences of these differing life histories and provide a genetic perspective on the taxonomic relationship between the two species. Genotypic diversity ofOphiocoma pumila collected at Discovery Bay, Jamaica, in 1985 conformed to expectations for a sexually reproducing population. In contrast, genotypic diversity ofOphiocomella ophiactoides at this site was significantly lower than expected for a sexually reproducing population, due largely to the predominance of clonal proliferation over larval recruitment. Large variation in clonal composition over a short (50 m) distance emphasized the very localized scale of clonal mixing in this species. Allozymic data are indicative of a close sibling species relationship betweenOphiocoma pumila andOphiocomella ophiactoides which suggests that the present generic separation of the two species should be re-examined. Electrophoretic analysis was also used to examine the genetic structure of sponge- and alga-dwelling populations of a second fissiparous brittle star,Ophiactis savignyi (Müller & Troschel), which was also collected at Jamaica in 1985. Striking differences in the allelic composition of sponge- and alga-dwellingO. savignyi were observed. Genotypic diversity ofO. savignyi in sponges was very low, each sponge being dominated by a single genotype. Genotypic diversity ofO. savignyi in algae was higher, although still significantly lower than expectations for a sexually reproducing population. In the light of the highly clonal composition of fissiparous brittle-star populations, the adaptive significance of clonal growth may be related to an increase in the overall fitness of dispersed clones (genets), compared to individuals of strictly sexual counterparts, through greater genotypespecific biomass and, hence, fecundity.Contribution No. 476 of the Discovery Bay Marine Laboratory of the University of the West Indies     

Small-scale genetic structure in the sea palm Postelsia palmaeformis Ruprecht (Phaeophyceae)

Documenting the scale of movement among populations is an important challenge for marine ecology. Using nine microsatellite markers, evidence of genetic structure in a marine kelp, the sea palm Postelsia palmaeformis Ruprecht, was examined in the vicinity of Cape Flattery, Washington state, USA (48° 24′ N, 124°44′ W). Genetic clustering analysis implemented without reference to geographic structure strongly suggested that a number of distinct genetic clusters existed among the 245 plants sampled in August in the years 1997–2001. Subsequent analysis showed that clustering was associated with geographically defined populations both among (km scale) and within (m scale) sampling sites. F _st analysis of geographically defined populations revealed significant genetic differentiation among populations of plants as little as 5 m apart, evidence of genetic structuring at even smaller scales, and a sharp increase in F _st across populations separated by up to 23 m. F _st values were also high and approximately unchanging (F _st=0.470) for populations separated by greater distances (up to 11 km), consistent with a scenario of rare dispersal by detached, floating plants carried by variable currents. The results corroborate natural history observations suggesting that P. palmaeformis has extremely short (1–3 m) spore dispersal distances, and indicate that the dynamics of sea palm populations are more affected by local processes than recruitment from distant populations.     

Latitudinal variability in spatial genetic structure in the invasive ascidian, Styela plicata

Increases in temperature can shorten planktonic larval durations, so that higher temperatures may reduce dispersal distances for many marine animals. To test this prediction, we first quantified how minimum time to settlement is shortened at higher temperatures for the ascidian Styela plicata. Second, using latitude as a correlate for ocean temperature and spatial genetic structure as a proxy for dispersal, we tested for a negative correlation between latitude and spatial genetic structure within populations, as measured by anonymous DNA markers. Spatial genetic structure was variable among latitudes, with significant structure at low and intermediate latitudes (high and medium temperatures) and there was no genetic structure within high-latitude (low temperature) populations. In addition, we found consistently high genetic diversity across all Australian populations, showing no evidence for recent local bottlenecks associated S. plicata’s history as an invasive species. There was, however, significant genetic differentiation between all populations indicating limited ongoing gene flow.     

Isolation by distance in the scleractinian coral <Emphasis Type="Italic">Seriatopora hystrix</Emphasis> from the Red Sea

Pelagic dispersal of larvae in sessile marine invertebrates could in principle lead to a homogeneous gene pool over vast distances, yet there is increasing evidence of surprisingly high levels of genetic differentiation on small spatial scale. To evaluate whether larval dispersal is spatially limited and correlated with distance, we conducted a study on the widely distributed, viviparous reef coral Seriatopora hystrix from the Red Sea where we investigated ten populations separated between ~0.150 km and ~610 km. We addressed these questions with newly developed, highly variable microsatellite markers. We detected moderate genetic differentiation among populations based on both F _ST and R _ST (0.089 vs. 0.136, respectively) as well as considerable heterozygote deficits. Mantel tests revealed isolation by distance effects on a small geographic scale (≤20 km), indicating limited dispersal of larvae. Our data did not reveal any evidence against strictly sexual reproduction among the studied populations.     

New expansions in old clades: population genetics and phylogeny of Gnatholepis species (Teleostei: Gobioidei) in the Pacific

Species of the reef goby genus Gnatholepis exhibit enormous geographic ranges with little evidence of population segregation detectable based on mitochondrial DNA. To determine if genetic differentiation is evident with more rapidly evolving markers, seven microsatellite loci were screened from the species Gnatholepis anjerensis and G. scapulostigma and population segregation was tested among fish from across the South Pacific. Both AMOVA and pairwise F _ST analyses showed that, in concordance with previous mitochondrial results, most genetic variance occurs within individual populations, as population differentiation is evident only over the largest distances (>3,700 km). This result is contrasted with previous studies demonstrating that despite their relatively long larval periods, some gobiid fishes exhibit population differentiation on small (<100 km) geographic scales. Coalescence analysis showed that current Pacific populations of these species originated in the Pleistocene, presumably related to sea level fluctuations associated with episodes of glaciation. However, rate analysis based on a phylogeny of Gnatholepis species indicates that the species themselves are much older, consistent with a complex history of rapid, short-term population contractions and expansions, with corresponding rapid dispersal.