Molecular genetic structure suggests limited larval dispersal in grass rockfish,<Emphasis Type="Italic"> Sebastes rastrelliger</Emphasis>

The grass rockfish (Sebastes rastrelliger Jordan and Gilbert, 1880) is a non-migratory, benthic, near shore species distributed along rocky reefs and sea grass beds. It occurs from Baja California, Mexico, to Oregon, USA, spanning the Oregonian and Californian biogeographic provinces. In California this fish receives intense fishing pressure from an expanding and loosely regulated live-fish fishery. It is not known whether or by what mechanism larvae and juveniles are retained locally or dispersed widely during the early life-history phase. Tissue samples of S. rastrelliger were collected between 1996 and 2001 from 405 adult fish at eight sites (42.70°N; 124.50°W to 32.67°N; 117.25°W) spanning the species range. Individuals were surveyed for polymorphism at six microsatellite loci. Allele frequency heterogeneity was not significant among all sites (F_ST=0.001; P=0.18), nor in pairwise comparisons, but a clear correlation of genetic and geographic distance was detected (P=0.019). Fit of genetic and geographic distance was stronger within biogeographic provinces than at a range-wide scale, suggesting that populations north and south of the Point Conception biogeographic boundary are not in equilibrium with respect to migration and genetic drift. Estimates of mean coastal dispersal distances associated with the isolation-by-distance relationship are on the order of 10 km generation^–1. Such limited dispersal in a species with a pelagic early life history suggests active retention mechanisms near the shore. This has important implications for coastal management zones and design of marine reserves.Communicated by J.P. Grassle, New Brunswick     

Hydrology influences population genetic structure and connectivity of the intertidal amphipod Corophium volutator in the northwest Atlantic

The mechanisms driving genetic structure in marine systems are elusive due to the difficulty of identifying temporal and spatial barriers to dispersal. By studying marine invertebrate species with limited dispersal potential, genetic structure can be directly related to physical and biological factors restricting connectivity. In the northwest Atlantic, the benthic brood-rearing amphipod Corophium volutator is distributed across basins with distinct circulation patterns and has the potential to disperse passively during its adult stage. We analyzed spatial genetic variation and migration rates across C. volutator’s North American range with sequence data for mitochondrial DNA and three novel nuclear markers using frequency and coalescent-based methods. We found low genetic differentiation within basins, but strong subdivision within the Bay of Fundy and a striking biogeographic break between the Bay of Fundy and Gulf of Maine, suggesting that genetic drift may act on populations in which connectivity is restricted due to the limitation of passive dispersal by hydrological patterns.     

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     

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.     

Implications of life history for genetic structure and migration rates of southern African coastal invertebrates: planktonic,abbreviated and direct development

The amount of genetic structure in marine invertebrates is often thought to be negatively correlated with larval duration. However, larval retention may increase genetic structure in species with long-lived planktonic larvae, and rafting provides a means of dispersal for species that lack a larval dispersal phase. We compared genetic structure, demographic histories and levels of gene flow of regional lineages (in most cases defined by biogeographic region) of five southern African coastal invertebrates with three main types of larval development: (1) dispersal by long-lived planktonic larvae (mudprawn Upogebia africana and brown mussel Perna perna), (2) abbreviated larval development (crown crab Hymenosoma orbiculare) and (3) direct development (estuarine isopod Exosphaeroma hylecoetes and estuarine cumacean Iphinoe truncata). We hypothesized that H. orbiculare, having abbreviated larval development, would employ a strategy of larval retention, resulting in genetic structure comparable to that of the direct developers rather than the planktonic dispersers. However, regional population structure was significantly lower in all species with planktonic larvae, including H. orbiculare, than in the direct developers. Moreover, nested clade analysis identified demographic histories resulting from low levels of gene flow (isolation by distance and allopatric fragmentation) in the direct developers only, and migration rates were significantly higher in all three species having planktonic larvae than in the direct developers. We conclude that the amount of genetic structure within marine biogeographic regions strongly depends on the presence or absence of free-swimming larvae. Whether such larvae are primarily exported or retained, whether they have long or short larval duration, and whether or not they are capable of active dispersal seems to have little effect on connectivity among 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.     

Regional differentiation and post-glacial expansion of the Atlantic silverside, <Emphasis Type="Italic">Menidia menidia,</Emphasis> an annual fish with high dispersal potential

The coastal marine environment of the Northwest Atlantic contains strong environmental gradients that create distinct marine biogeographic provinces by limiting dispersal, recruitment, and survival. This region has also been subjected to numerous Pleistocene glacial cycles, resulting in repeated extirpations and recolonizations in northern populations of marine organisms. In this study, we examined patterns of genetic structure and historical demography in the Atlantic silverside, Menidia menidia, an annual marine fish with high dispersal potential but with well-documented patterns of clinal phenotypic adaptation along the environmental gradients of the Northwest Atlantic. Contrary to previous studies indicating genetic homogeneity that should preclude regional adaptation, results demonstrate subtle but significant (F _ST = 0.07; P < 0.0001) genetic structure among three phylogeographic regions that partially correspond with biogeographic provinces, suggesting regional limits to gene flow. Tests for non-equilibrium population dynamics and latitudinal patterns in genetic diversity indicate northward population expansion from a single southern refugium following the last glacial maximum, suggesting that phylogeographic and phenotypic patterns have relatively recent origins. The recovery of phylogeographic structure and the partial correspondence of these regions to recognized biogeographic provinces suggest that the environmental gradients that shape biogeographic patterns in the Northwest Atlantic may also limit gene flow in M. menidia, creating phylogeographic structure and contributing to the creation of latitudinal phenotypic clines in this species.     

Connectivity between marine reserves and exploited areas in the philopatric reef fish Chrysoblephus laticeps (Teleostei: Sparidae)

‘No-take’ marine protected areas (MPAs) are successful in protecting populations of many exploited fish species, but it is often unclear whether networks of MPAs are adequately spaced to ensure connectivity among reserves, and whether there is spillover into adjacent exploited areas. Such issues are particularly important in species with low dispersal potential, many of which exist as genetically distinct regional stocks. The roman, Chrysoblephus laticeps, is an overexploited, commercially important sparid endemic to South Africa. Post-recruits display resident behavior and occupy small home ranges, making C. laticeps a suitable model species to study genetic structure in marine teleosts with potentially low dispersal ability. We used multilocus data from two types of highly variable genetic markers (mitochondrial DNA control region and seven microsatellite markers) to clarify patterns of genetic connectivity and population structure in C. laticeps using samples from two MPAs and several moderately or severely exploited regions. Despite using analytical tools that are sensitive to detect even subtle genetic structure, we found that this species exists as a single, well-mixed stock throughout its core distribution. The high levels of connectivity identified among sites support the findings of previous studies that have indicated that inshore MPAs are an adequate tool for managing overexploited temperate reef fishes. Even though dispersal of adult C. laticeps out of MPAs is limited, the fact that the large adults in these reserves produce exponentially more offspring than their smaller counterparts in exploited areas makes MPAs a rich source of recruits. We nonetheless caution against concluding that the lack of structure identified in C. laticeps and several other southern African teleosts can be considered to be representative of marine teleosts in this region in general. Many such species are represented in more than one marine biogeographic province and may be comprised of regionally adapted stocks that require individual management.     

New insight on population genetic connectivity of widespread amphidromous prawn Macrobrachium lar (Fabricius, 1798) (Crustacea: Decapoda: Palaemonidae)

Due to the sparse and unstable nature of insular freshwater habitats, marine larval dispersal of amphidromous species is considered a critical element of population persistence. We assessed population genetic structure of freshwater prawn Macrobrachium lar across its range that encompasses two biogeographic barriers: the vast open ocean separating Western and Central Pacific regions and the Indo-Malay archipelago separating Indian and Pacific oceans. A total of 173 samples collected from 21 islands throughout the Indo-Pacific were sequenced at 16S and 28S rDNA. We observed distinct genetic isolation of populations located at the eastern and southwestern edge of the species range but no evidence of an effect of the Indo-Pacific barrier. Differentiation patterns are consistent with a stepping-stone model of dispersal. Genetic differences of Central Pacific populations may reflect founder events associated with colonization of isolated islands, or be a signature of a past bottleneck after population depletion caused by drastic climatic events.     

Low to moderate levels of genetic differentiation detected across the distribution of the New Zealand abalone, Haliotis iris

Two regions of the mitochondrial genome (cytochrome oxidase I and ATPase 8–ATPase 6) were used to examine the population genetic structure of New Zealand’s endemic abalone (Haliotis iris). Samples were collected from 28 locations around New Zealand between January 2005 and February 2008. At least four phylogeographic breaks were present and occurred across the Chatham rise, in the western Cook Strait region, along the southeast coast of the South Island, and at East Cape in the North Island. Gene flow across the Chatham rise is probably limited due to infrequent dispersal across large geographic distances (~850 km), while factors limiting gene flow around the North and South Islands are less clear, and understanding these may require intense temporal and spatial sampling in complex hydrographic regions. High genetic diversity and weak genetic structure may be a general feature of abalone potentially reflecting large and/or ancient populations.     

Complex genetic patterns and a phylogeographic disjunction among New Zealand mud snails Zeacumantus subcarinatus and Z. lutulentus

The comparative analysis of genetic structure and phylogeography of marine species can reveal the relative importance of common biogeographic barriers and species-specific evolutionary histories. In the present study, mitochondrial COI sequences from a total of 724 individuals collected from 38 localities throughout New Zealand during 2006 and 2007 were used to examine the genetic structure and demographic histories of the intertidal gastropods Zeacumantus subcarinatus and Zeacumantus lutulentus. For both species, results revealed isolation of populations located along the northern South Island and southern and western North Island, and extensive genetic structure throughout the remainder of their ranges. Despite this, long-distance dispersal and secondary admixture of divergent haplotypes was evident, especially for the widespread Z. subcarinatus. These findings reveal the importance of common barriers to gene flow and highlight how the interaction of inherent dispersal limitations of direct-developing marine organisms and periodic long-distance movements can produce complex genetic patterns.     

A spatial gradient in the potential reproductive output of the sea mussel <Emphasis Type="Italic">Mytilus californianus</Emphasis>

Spatial variation in reproductive output from different populations within a region could have important consequences for recruitment, and cascading effects on populations and communities of marine species, but is rarely examined over meso-scales (i.e., tens to hundreds of kilometers). In this study, reproduction in the dominant mid-intertidal mussel, Mytilus californianus, was examined from sites spanning Point Conception, California over a 6-month period (March–August 2000). There was a dramatic geographic pattern in the relationship between size and potential reproductive output that was qualitatively similar across all 6 months sampled. Increases in allocation to reproductive tissue with increasing body size occurred at all sites, but the slope nearly doubled at sites south of Point Conception compared to northern sites. The spatial variation in size-specific reproductive output, coupled with additional spatial gradients in mussel density and size distributions, combined to increase total reproductive output by over eightfold at southern relative to northern sites. This study highlights the need to explicitly examine spatial patterns of reproductive output at these meso-scales, in order to better understand connectivity and source–sink dynamics in marine systems.     

Genetic uniformity of widely separated populations of the coral reef starfishLinckia laevigata from the East Indian and West Pacific Oceans,revealed by allozyme electrophoresis

Gene flow among 18 widely separated populations of the starfishLinckia laevigata was investigated using allozyme electrophoresis at seven polymorphic enzyme loci. Little genetic differentiation was observed among East Indian-West Pacific populations separated by thousands of kilometres. Gene flow was estimated to be of the same order of magnitude as that found in the highly connected Great Barrier Reef region. The absence of genetic structure over such a broad geographic range is consistent with any of three conclusions: (1) there is extensive dispersal among widely separated populations across the range examined, (2) there has been a rapid expansion of the East Indian-West Pacific populations in the recent past, or (3) the loci surveyed are under the influence of balancing selection. The first two conclusions are not totally exclusive, since a recent expansion over several thousand generations would also require rapid dispersal at some stage, although present-day levels of dispersal need not be of the same order. With the available data, it is not possible to distinguish which of these mechanisms is the most likely. The most parsimonious conclusion is that extremely low levels of population differentiation are consistent with the existence of a large, homogeneous, panmictic population, with extensive dispersal occurring throughout the East Indian-West Pacific.     

Extensive gene flow among mytilid (Bathymodiolus thermophilus) populations from hydrothermal vents of the eastern Pacific

Prior studies of the hydrothermal vent mussel Bathymodiolus thermophilus (Bivalvia: Mytilidae), provided conflicting predictions about the dispersal ability and population structure of this highly specialized species. Analyses of morphological features associated with its larval shells revealed a feeding larval stage that might facilitate dispersal between ephemeral vent habitats. In contrast, an allozyme study revealed substantial genetic differentiation between samples taken from populations 2370 km apart on Galápagos Rift (Latitude 0°N) and the East Pacific Rise (13°N). To resolve the discrepancy between these studies, we examined allozyme and mitochondrial (mt) DNA variation in new samples from the same localities plus more recently discovered sites (9° and 11°N) along the East Pacific Rise. Although analysis of 26 enzyme-determining loci revealed relatively low levels of genetic variation within the five populations, no evidence existed for significant barriers to dispersal among populations. We estimated an average effective rate of gege flow (Nm) of 8 migrants per population per generation. Two common mtDNA variants predominated at relatively even frequencies in each population, and similarly provided no evidence for barriers to gene flow or isolation-by-distance across this species' known range. Larvae of this species appear to be capable of dispersing hundreds of kilometers along a continuous ridge system and across gaps separating non-contiguous spreading centers.     

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.     

Mitochondrial DNA analyses of the red rock lobsterJasus edwardsii supports an apparent absence of population subdivision throughout Australasia

Nucleotide sequence polymorphism in the mitochondrial genomes of 132 adult lobsters (Jasus edwardsil) collected from widespread locales across southern Australia and from New Zealand (April 1989 to June 1990) was assayed, using six restriction endonucleases, to test the hypothesis of a lack of genetic subdivision in a marine species with a long-lived planktonic larva. The mean amount of mtDNA diversity among the 132 mitochondrial genomes was 0.77%. Phenetic clustering and gene-diversity analyses, as well as pairwise comparison of the genetics of specimens from each, or grouped, locales did not detect the presence of genetic subdivision across approx 4600 km of Southern Ocean habitats. The inability of this study to detect population subdivision does not preclude fortutitous, active or habitat-specific larval settlement from producing and maintaining hidden groupings. If genetic homogeneity is maintained in this species by larval dispersal in ocean currents flowing to the east, then westerly populations may deserve special conservation status.     

Contrasting population genetic structures of sympatric, mass-spawning Caribbean corals

Coral reef conservation management policy often focuses on larval retention and recruitment of marine fish with scant data available on important, less motile reef-building species such as corals. To evaluate the concept of population connectivity in corals, we tested whether broadcast spawning reproduction per se confers the same degree of dispersal to two sister species, Montastraea annularis (Anthozoa: Scleractinia; Ellis and Solander 1786) and M. faveolata (Ellis and Solander 1786), both dominant taxa in reefs of the northern Caribbean. Genetic analyses of ten nuclear DNA loci (seven microsatellite and three single-copy RFLP) reveal strikingly different patterns of population genetic subdivision for these closely related, sympatric species, in spite of likely identical dispersal abilities. Strong population genetic structure typified the architecture of M. annularis, whereas M. faveolata populations were principally genetically well mixed. A higher level of clonality was observed in M. annularis potentially because of a susceptibility to physical fragmentation. Clonality did not, however, significantly contribute to population genetic structure or low-level Hardy–Weinberg and linkage disequilibria observed in some populations. The lack of consistent association between reproductive mode and dispersal reinforces the perspective that population connectivity is not so much a function of predictable marine population source and sink relationships as is due to a more complex interface of oceanic currents interacting with and amplifying stochastic fluctuations in larval supply and settlement success. Our results support others promoting an overall ecosystem approach in marine protected area design.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Estimating dispersal potential for marine larvae: dynamic models applied to scleractinian corals

Dispersal influences ecological dynamics, evolution, biogeography, and biodiversity conservation, but models of larval dispersal in marine organisms make simplifying assumptions that are likely to approximate poorly the temporal dynamics of larval survival and capacity for settlement. In particular, larval mortality rates are typically assumed to be constant throughout larval life; and all larvae are frequently assumed to acquire and lose competence at the same time. To improve upon these assumptions, we here develop simple models of dispersal potential that incorporate rates of mortality, and acquisition and loss of settlement competence. We fit these models to empirical competence and survival data for five scleractinian coral species, to test the models' ability to characterize empirical survival and competence patterns, and to estimate the dispersal potential implied by those patterns. The models fit the data well, incorporating qualitative features of competence and survival that traditional approaches to modeling dispersal do not, with important implications for dispersal potential. Most notably, there was high within-cohort variation in the duration of the competent period in all species, and this variation increases both self-recruitment and long-distance dispersal compared with models assuming a fixed competent period. These findings help to explain the seeming paradox of high genetic population structure, coupled with large geographic range size, observed in many coral species. More broadly, our approach offers a way to parsimoniously account for variation in competence dynamics in dispersal models, a phenomenon that our results suggest has important effects on patterns of connectivity in marine metapopulations.     

Chaotic genetic patchiness and high relatedness of a poecilogonous polychaete in a heterogeneous estuarine landscape

The genetic structure of benthic marine invertebrates is often described as “chaotic” when genetic structure cannot be explained and barriers to dispersal and gene flow cannot be identified. Here, chaotic patterns of genetic structure for the polychaete Pygospio elegans (Claparède) sampled at 16 locations from the heterogeneous Isefjord–Roskilde Fjord estuary complex in Denmark were found. There was no isolation by distance, and the geography of the estuary complex did not seem to pose a barrier to dispersal and gene flow in this species. We investigated whether characteristics of the environment could be related to the genetic structure and possibly restrict gene flow in this species. Additionally, since P. elegans is poecilogonous, producing larvae with different pelagic developmental periods, we investigated whether observed developmental modes in the samples might clarify the genetic patterns. None of the tested factors explained the population genetic structure. However, a high degree of relatedness among individuals in almost all samples was found. Samples with a larger percentage of young individuals had more related individuals, suggesting that different cohorts could be comprised of individuals with different degrees of relatedness. Relatedness within a site could be increased by limited larval dispersal, collective dispersal of related larvae, sweepstakes reproductive success, or asexual reproduction, but distinguishing between these requires further study. Using a “seascape genetics” approach allowed us to investigate some of the numerous potential factors that could influence population genetic structure in a poecilogonous species.     

Mitochondrial control region variability and global population structure in the swordfish,Xiphias gladius

Little is known about what limits genetic exchange in highly vagile, open ocean vertebrate species, such as the swordfish Xiphias gladius L. Reduced abundance of swordfish in some regions, and increased fishing pressure in others, has raised concerns and fueled interest in a more complete evaluation of the resource. In this study, global population structure in swordfish was assessed by sequencing a 300 base pair segment of the 5 end of mitochondrial DNA control region from 159 swordfish collected in three ocean basins: the Mediterranean, Atlantic and Pacific, over the years 1988 to 1994. Among the 159 individuals, 95 polymorphic sites delineated 121 unique haplotypes, indicating a high level of polymorphism on a global scale. A phylogenetic analysis of the unique DNA haplotypes revealed two divergent clades with differing geographic distributions. Phylogeographic concordance of this pattern with that of two other pelagic fish species suggests a biogeographic explanation for this structure. An analysis of molecular variance (AMOVA) revealed significant geographic partitioning of molecular variation among the three ocean basins, indicating that swordfish populations are structured on a global scale. Estimates of genetic exchange among populations within an ocean basin were high, indicating panmixia within ocean basins. Since the haplotypic diversity exhibited by the swordfish control regions is extremely high, much larger sample sizes may be necessary to detect subdivision within ocean basins.