Lack of population genetic structuring in the marine planktonic diatom Pseudo-nitzschia pungens (Bacillariophyceae) in a heterogeneous area in the Southern Bight of the North Sea

Several marine holoplanktonic organisms show a high degree of geographically structured diversity for which it often remains unclear to what extent this differentiation is due to the presence of cryptic taxa. For the genetically distinct diatom Pseudo-nitzschia pungens var. pungens, we used six microsatellite markers to investigate the spatial and temporal genetic composition in the heterogeneous Southern Bight of the North Sea. Although our sampling area (ca. 100 km) comprised water bodies with different environmental conditions (marine, estuary, saline lake) and different degrees of connectivity (from complete isolation to supposedly free exchange between environments), no evidence of genetic differentiation was found. Expanding our sampling area (ca. 650 km), suggested a homogenous population structure over even larger areas in the North Sea. Our results suggest that the population structure of this diatom is mainly shaped by strong homogenizing effects of gene flow preventing genetic drift, even in water bodies with limited connectivity. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Population structure of short-beaked common dolphins (<Emphasis Type="Italic">Delphinus delphis</Emphasis>) in the North Atlantic Ocean as revealed by mitochondrial and nuclear genetic markers

The understanding of population structure and gene flow of marine pelagic species is paramount to monitoring, management and conservation studies. Such studies are often hampered by the potentially high dispersal behavior of the species, the lack of obvious geographical barriers in the marine environment and the scarce sample availability. Short-beaked common dolphins (Delphinus delphis) are widespread in coastal and open-ocean habitats of the North Atlantic Ocean, nevertheless population structure and migratory patterns are poorly understood. Furthermore, concern has been raised about the status of the species because large numbers of dolphins have been taken incidentally in several fisheries throughout the North Atlantic in the past decades. In the present study, a large number of individual samples were obtained from seasonal and spatial aggregations of common dolphins from western (wNA) and eastern North Atlantic (eNA) regions, mostly using opportunistic sampling (i.e. from incidental entanglement in fishing gear or beach-cast carcasses). Genetic variability was investigated using nuclear (14 microsatellite loci) and mitochondrial (360 bp of the control region) genetic markers. Levels of genetic diversity were relatively high in all sampled areas and no evidence of recent reduction of effective population size (i.e. bottleneck) was detected at the nuclear loci. Significant population structure was detected between the two main regions (wNA and eNA) where it appeared to be more pronounced at mitochondrial (F _ST = 0.018, P < 0.001) than nuclear markers (F _ST = 0.005, P < 0.05), indicating the presence of at least two genetically distinct populations of common dolphins in the North Atlantic Ocean. In contrast, no significant genetic structure was detected between temporal aggregations of dolphins from within the same region, suggesting possible seasonal movement patterns at a regional scale. The observed levels of genetic differentiation between classes of markers are discussed here as a possible consequence of migratory patterns or recent population subdivision. An erratum to this article can be found at     

Population connectivity in the temperate damselfish <Emphasis Type="Italic">Parma microlepis</Emphasis>: analyses of genetic structure across multiple spatial scales

This study investigated the utility of microsatellite markers for providing information on levels of population connectivity for a low dispersing reef fish in New South Wales (NSW), Australia, at scales ≤400 km. It was hypothesized that the temperate damselfish Parma microlepis, which produces benthic eggs and has limited post-settlement dispersal, would exhibit spatial genetic structure and a significant pattern of isolation-by-distance (IBD). A fully nested hierarchical sampling design incorporating three spatial scales (sites, location and regions, separated by 1–2, 10–50 and 70–80 km respectively) was used to determine genetic variability at seven microsatellite loci. Broad-scale genetic homogeneity and lack of IBD was well supported by single and multi-locus analyses. The proportion of the total genetic variation attributable to differences among regions, locations or sites was effectively zero (Φ/R-statistics ≤0.007). The geographic distribution of genetic diversity and levels of polymorphism (H _E 0.21–0.95) indicate high mutation rates, large effective population sizes, and high rates of gene flow. Significant gene flow may be driven by factors influencing pre-settlement dispersal, including the East Australian Current (EAC) and habitat continuity. Genetic connectivity may not reflect demographically important connectivity, but does imply that P. microlepis populations are well connected from an evolutionary perspective. Total observed genetic diversity was accounted for within 1–2 km of reef and could be represented within small Marine Protected Areas. Reef fishes in NSW which have life histories similar to P. microlepis (e.g. pre-settlement durations ≥2 weeks) are also likely to exhibit genetic homogeneity. Genetic markers are, therefore, most likely to provide information on demographically relevant connectivity for species with lower dispersal capabilities, small population sizes, short life spans, and whose habitats are rare, or patchily distributed along-shore. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Patterns of mussel recruitment in southern Africa: a caution about using artificial substrata to approximate natural recruitment

Quantifying sessile marine invertebrate recruitment often requires destructive sampling or extrapolation from artificial substrata, the latter introducing the danger of artifacts. We measured intertidal mussel recruitment into mussel beds and into brushes at three-month intervals for five years across 3,200 km of southern Africa and determined substrata effects on recruitment rate. Recruitment into mussel beds showed a strong, coast-wide gradient, with high recruitment on the West coast, diminishing on the South coast, and increasing slightly on the East coast. At scales of 10 s of km, brushes reflected natural temporal recruitment variability, with a strong significant linear correlation between recruitment into brushes and into mussel beds. However, the relationship became semi-logarithmic when comparing among locations at a scale of 100 s of km. Artificial substrata thus reflect local natural settlement well but may be a poor indicator of it when spatial scales are large, particularly when mussel bed topography is complex, or localities have very different recruitment densities.     

Endemism and dispersal: comparative phylogeography of three surgeonfishes across the Hawaiian Archipelago

To evaluate the hypothesis that a general correlation exists between species range size and dispersal ability, we surveyed mitochondrial cytochrome b sequence variation in three surgeonfish species with vastly different ranges: Ctenochaetus strigosus, Hawaiian endemic, N = 531; Zebrasoma flavescens, North Pacific, N = 560; Acanthurus nigrofuscus, Indo-Pacific, N = 305. Collections were made throughout the 2,500 km expanse of the Hawaiian Archipelago and adjacent Johnston Atoll. Analyses of molecular variance demonstrate that all three species are capable of maintaining population connectivity on a scale of thousands of km (all species global Φ_ST = NS). However, rank order comparison of pairwise Φ_ST results and Exact test P-values revealed modest but significantly different patterns of gene flow among the three species surveyed, with the degree of genetic structure increasing as range size decreases (P = 0.001). These results are consistent with mtDNA surveys of four additional Hawaiian reef fauna in which a wide-spread Indo-Pacific species exhibited genetic homogeneity across the archipelago, while three endemics had significant population subdivision over the same range. Taken together, these seven cases invoke the hypothesis that Hawaii’s endemic reef fishes evolved from species with reduced dispersal ability that, after initial colonization, could not maintain contact with parent populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Molecular insight into the population structure of common and spotted dolphins inhabiting the pelagic waters of the Northeast Atlantic

Several cetacean species exhibit fine-scale population structure despite their high dispersal capacities and the apparent continuity of the marine environment. In dolphins, most studies have focused on coastal areas and continental margins, and they revealed differentiated populations within relatively small geographic areas, sometimes in conjunction with a specialisation for different habitats (ecotypes). We analysed the population genetic structure of short-beaked common dolphins (Delphinus delphis) and Atlantic spotted dolphins (Stenella frontalis) in the Azores and Madeira, the two most isolated archipelagos of the North Atlantic. The archipelago of the Azores is divided into three groups of islands and stands 900 km away from Madeira. It is not known whether individuals migrate between groups of islands and archipelagos, nor whether distinct ecotypes are present. These questions were investigated by genetic analyses of 343 biopsy samples collected on free-ranging dolphins. The analyses consisted in sequencing part of the mitochondrial hyper-variable region, screening up to 14 microsatellite loci, and molecular sexing. Results did not unravel any population structure at the scale of the study area. Lack of differentiation matches expectations for spotted dolphins, which are transient in both archipelagos, but not for common dolphins, which are present year-round in the Azores and potentially resident. Absence of genetic structure over hundreds and even thousands of kilometres implies the existence of gene flow over much larger distances than usually documented in small delphinids, which could be achieved through individual movements. This finding indicates that population structure in oceanic habitat differs from that observed in coastal habitat.     

Comparative phylogeography of marine cladocerans

We examined the population genetics of six species of marine cladocerans, using a ~600 bp fragment of the cytochrome oxidase subunit I gene sequence. Phylogenetic analysis revealed significant intraspecific, semi-allopatric phylogenetic breaks in four out of five species belonging to the Podonidae, supporting an ancient radiation and oceanic expansion for this group. By contrast, Penilia avirostris (Sididae) displayed no phylogeographic structure across a global sampling, suggesting a recent worldwide expansion. Our results also show a transoceanic distribution of identical or very similar haplotypes in several species of marine Cladocera, which may be interpreted as either natural transport or evidence of recent anthropogenic transport. If the latter is the case, marine cladocerans represent one of the first genetically documented cases of exotic or invasive marine zooplankton, likely an underreported group. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Sequence data from this article have been deposited with the EMBL/GenBank Data Libraries under accession nos. EU675871–EU675924.     

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.     

Spatial and temporal genetic homogeneity in the Arctic surfclam (<Emphasis Type="Italic">Mactromeris polynyma</Emphasis>)

Commercially harvested marine bivalve populations show a broad range of population-genetic patterns that may be driven by planktonic larval dispersal (gene flow) or by historical (genetic drift) and ecological processes (selection). We characterized microsatellite genetic variation among populations and year classes of the commercially harvested Arctic surfclam, Mactromeris polynyma, in order to test the relative significance of gene flow and drift on three spatial scales: within commercially harvested populations in the northwest Atlantic; among Atlantic populations; and between the Atlantic and Pacific oceans. We found small nonsignificant genetic subdivision among eight populations from the northwest Atlantic (F _ST = 0.002). All of these Atlantic populations were highly significantly differentiated from a northeast Pacific population (F _ST = 0.087); all populations showed high inbreeding coefficients (F _IS = 0.432). We tested one likely source of heterozygote deficits by aging individual clams and exploring genetic variation among age classes within populations (a temporal Wahlund effect). Populations showed strikingly different patterns of age structure, but we found little differentiation among age classes. In one case, we were able to analyze genetic diversity between age classes older or younger than the advent of intensive commercial harvesting. The results generally suggest spatially broad and temporally persistent genetic homogeneity of these bivalves. We discuss the implications of the results for the biology and management of surfclam populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Temporal variation of recruits as a basis of ephemeral genetic heterogeneity in the western rock lobster Panulirus cygnus

Allozymes were examined in quantitative lunar monthly collections of larval recruits of the western rock lobster Panulirus cygnus George over three recruitment seasons at two sites nearly 350 km apart in Western Australia. At Alkimos, the southern site, recruitment occurs in a relatively narrow peak early in the spring, whereas at the northern Houtman Abrolhos Islands, recruitment extends into the summer months. In the 1995/1996 recruitment season, the frequency of the GPI ^* 100 allele increased from early to late in the season, but the frequencies were indistinguishable at the two sites in each monthly collection. The combination of this temporal variation in allelic frequencies with the contrasting patterns of recruitment at the Abrolhos Islands and Alkimos resulted in genetically different cohorts at the two sites. This pattern was ephemeral, as it was not repeated in the subsequent two years. Thus, ephemeral genetic patchiness in P. cygnus can be generated by the locally-specific genetic mix of recruits obtained from a common larval pool. This mechanism is the probable explanation of previously observed temporal and possible spatial genetic variation in adult P. cygnus, and highlights the importance of studying recruitment in order to understand the genetic structure of marine species. Received: 22 February 1999 / Accepted: 8 June 1999     

Meta-analysis of ecosystem services associated with oyster restoration

Restoration of foundation species promises to reverse environmental degradation and return lost ecosystem services, but a lack of standardized evaluation across projects limits understanding of recovery, especially in marine systems. Oyster reefs are restored to reverse massive global declines and reclaim valuable ecosystem services, but the success of these projects has not been systematically and comprehensively quantified. We synthesized data on ecosystem services associated with oyster restoration from 245 pairs of restored and degraded reefs and 136 pairs of restored and reference reefs across 3500 km of U.S. Gulf of Mexico and Atlantic coastlines. On average, restoration was associated with a 21-fold increase in oyster production (mean log response ratio = 3.08 [95% confidence interval: 2.58–3.58]), 34–97% enhancement of habitat provisioning (mean community abundance = 0.51 [0.41–0.61], mean richness = 0.29 [0.19–0.39], and mean biomass = 0.69 [0.39–0.99]), 54% more nitrogen removal (mean = 0.43 [0.13–0.73]), and 89–95% greater sediment nutrients (mean = 0.67 [0.27–1.07]) and organic matter (mean = 0.64 [0.44–0.84]) relative to degraded habitats. Moreover, restored reefs matched reference reefs for these ecosystem services. Our results support the continued and expanded use of oyster restoration to enhance ecosystem services of degraded coastal systems and match many functions provided by reference reefs.     

Mortality of newly metamorphosed eastern oysters (Crassostrea virginica) in mesohaline Chesapeake Bay

 Stocks of eastern oysters, Crassostrea virginica (Gmelin), in mesohaline Chesapeake Bay, USA, exhibit a high degree of inter-annual and spatial variability in recruitment. We found that cumulative oyster spatfall on off-bottom collector plates, measured throughout the summer in 14 years over a span of three decades, was highly positively correlated (r ² = 0.8) with juvenile oyster recruitment on adjacent oyster bars. Total abundances of juvenile oysters on these bars were, however, generally 99.7% lower than predicted from cumulative seasonal larval settlement on collector plates. We propose that although the number of larvae metamorphosing was the key factor in determining the gross annual pattern of recruitment to these mesohaline oyster bars, the actual magnitude of recruitment was governed by post-settlement processes, such as competition for limited resources and predation. We tested the hypothesis that predation may be partly responsible for high post-settlement juvenile oyster mortality. We performed a series of 3-d field investigations over two summers (1989, 1990) at a mesohaline site, employing cages of various mesh sizes (400, 800, 1500 μm) to protect hatchery-reared spat of 0.5 to 4.0 mm shell height. Mortality rates for spat held for 3 d in the estuary (17.8%) were significantly higher (P = 0.0001) for the smallest spat (0.5 to 2.0 mm) compared with those of 2.01 to 4.0 mm (4.2%). In 1990, but not in 1989, enclosure within 400 and 800 μm mesh cages significantly (P = 0.004) increased survival during 3-d deployments (9.4 and 10.1%, respectively) compared with spat unprotected by mesh cages (21.9%). In a series of laboratory predation studies that used the entire community of invertebrates that could penetrate the cages, microscopic juvenile polyclad flatworms, Stylochus ellipticus, were the only organisms that we observed crawling into living oysters and feeding on oyster tissue. Large flatworms (50 to 200 mm²) are known to be important predators on oysters, but this ability of flatworms that were so small (<ca. 5 mm²) and translucent as to be almost invisible without magnification to feed on immediate post-metamorphic oysters has not been documented previously. Our results suggest that the rate of mortality due to predation in mesohaline Chesapeake Bay is much reduced once spat survive for 2 to 3 weeks post-metamorphosis. Thus, it is likely that predation in the 1 to 2 week period immediately after settlement may be a crucial factor in the structuring of eastern oyster populations. Received: 21 December 1998 / Accepted: 2 December 1999     

Genetics of larvae and spat growth rate in the oyster Crassostrea virginica

The heritability of oyster (Crassostrea virginica) larval growth rate was estimated to be in the range of 0.25 to 0.50 and a significant part of this genetic variation is of the additive type. Larval growth rate and spat growth rate were found to be highly correlated. These results suggest that a selection program for faster growing larvae and spat would be successful.     

Phylogeography and demographic history of <Emphasis Type="Italic">Atherina presbyter</Emphasis> (Pisces: Atherinidae) in the North-eastern Atlantic based on mitochondrial DNA

A fragment of the mitochondrial control region was used to assess phylogeographic patterns and historical demography of the sand-smelt Atherina presbyter in the North-eastern Atlantic, covering its geographical range. A striking result is the highly marked differentiation between the Canary Islands population and western European ones. A genetic structure among European populations of A. presbyter was revealed, with a pattern of isolation-by-distance or a gradient effect at a scale of hundreds kilometres, an uncommon pattern likely related to the biological and life-history traits of the sand-smelt. The northern European populations present a much lower genetic diversity when compared to southern populations, which is consistent with a recent colonization from southern populations. The results showed signs of Pleistocene signatures, with the population age estimates for the European populations being clearly older than the Last Glacial Maximum (18,000 years bp). Nevertheless, paleotemperature reconstructions show that the sand-smelt could not have inhabited the western European shores during the last glacial phase. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of recruitment on genetic patchiness in the urchinEchinometra mathaei in Western Australia

Enzyme polymorphisms in the sea urchinEchinometra mathaei were examined to test the relative influences of population turnover and patchiness in recruitment on genetic heterogeneity. We found that the total variance in allelic frequency among three populations separated by approximately 4 km at Rottnest Island, Western Australia (collected in February 1985) is as large as that among five additional samples collected over a distance of 1 300 km along the Western Australian coast in August 1987. This suggests that the forces causing genetic differentiation act on a local scale and occur in a single generation. A comparison of sites with different histories of recruitment indicates that the observed genetic differences among age groups are the result of prerecruitment effects, and that differences among sites reflect their individual histories of recruitment.     

Genetic structure of the common sole Solea vulgaris at different geographic scales

The genetic structure of the flatfish Solea vulgaris was investigated on several spatial scales and at the temporal level through analysis of electrophoretic variation at 8 to 12 polymorphic enzyme loci. No differentiation was apparent at the temporal scale. Some differentiation was detected at and above the regional scale. Isolation by distance was evidenced by the significant correlation between genetic and geographic distances, and by the consistency of the results of multiple-locus correspondence analysis with geographic sampling patterns. The analysis suggested that the geographic unit of population structure (i.e. a geographical area corresponding to a panmictic or nearly panmictic population) lies within a radius of the order of 100 km. The isolation-by-distance pattern in S. vulgaris contrasted with the known genetic structures of other flatfish species of the northeastern Atlantic and Mediterranean in a way that may be related to the range of their respective temperature tolerances for eggs and larvae.     

Geographical variation in metacercarial infection levels in marine invertebrate hosts: parasite species character versus local factors

Despite the important roles played by parasites in local population dynamics and community structure of marine ecosystems, there is a lack of information on the geographical variation in infection levels displayed by particular host–parasite species combinations. This study examines geographical variation in infection levels by the metacercarial stages of trematode parasites in crustacean and bivalve second intermediate hosts. Analyses were based on a dataset compiled from the literature, consisting of 164 local samples representing 49 host–parasite species pairs for crustaceans, and 338 entries representing 36 host–parasite species pairs for bivalves. The analyses indicate that for all measures of infection levels [prevalence (percentage of individuals infected), intensity (mean no. of metacercariae per infected individual), abundance (mean no. of metacercariae across all individuals in a sample)], there was statistically significant repeatability of infection values within host–parasite species pairs. However, it is only for values of intensity and abundance of infection in crustacean hosts that the repeatability was strong; this suggests that infection levels are specific properties of crustacean–trematode species pairs, showing significant consistency across localities despite spatial variation in abiotic and biotic conditions. Although the magnitude of variation in infection levels within parasite species pairs (measured as coefficients of variation) was independent of scale in crustacean hosts, infection levels in bivalves increased in variability at large (>100 km) spatial scales. These results suggest that there is a considerable geographical consistency in parasite load, especially in crustacean hosts, which should lead to consistent ecological and ecosystem effects of marine trematodes. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Genetic structuring and gene flow in Cerastoderma glaucum (Bivalvia: Cardiidae): evidence from allozyme variation at different geographic scales

Genetic variation at 16 enzyme loci was analysed in 20 Mediterranean and 1 Baltic population of Cerastoderma glaucum. Spatial genetic variation at different geographic scales was investigated. In general, this species was fairly genetically structured (over all loci Š=0.088), according to the fragmentary nature of its habitat. Spatial structuring of genetic diversity was shown to follow different models, depending on the geographic scale considered: a stepping-stone model provided a good fit at a wide scale, with gene flow inversely related to geographic distance, whereas at small scale, genetic relationships among samples could not be interpreted as simply the effect of physical distance among populations. Results are discussed taking into account the intrinsic and extrinsic factors which are most likely to affect the patterns of genetic structuring in coastal marine animals. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00227-001-0753-x     

Current shifts and kin aggregation explain genetic patchiness in fish recruits

The scales of population structure in marine species depend on the degree to which larvae from different populations are mixed in the plankton. There is an intriguing trend in marine population genetic studies of significant genetic structure for larvae, recruits, or populations at fine scales that is unpatterned across space and changes through time. This "chaotic genetic patchiness" suggests that larval pools are not well mixed in the plankton. However, few studies have been able to distinguish among potential causes of spatial and temporal genetic heterogeneity: changes in larval migration patterns, changes in environmental selection, or stochasticity caused by "sweepstakes" reproductive success of spawners creating detectable family structure. Here we use microsatellite markers to show that significant allele frequency shifts occurred sporadically in space and time for cohorts of recruits of Paralabrax clathratus (kelp bass) collected once every two weeks over two years from five sites in the Santa Barbara Channel, California, USA. We found that the pattern of genetic differentiation among cohorts was explained by a combination of (1) family structure in some cohorts, evidenced by half and full siblings, and (2) an indication of changes in larval delivery. It is unlikely but possible that environmental selection also plays a role. Although sampling of potential source populations was incomplete, cohorts arriving during western current flows show most genetic similarity with a population sample collected in the west, and cohorts arriving during current flows from the southeast show similarity with population samples collected in the south and east. Despite the family structure apparent in some cohorts, these "sweepstakes" events occur on too fine a scale to create lasting year class genetic structure. The results corroborate oceanographic models of larval dispersal, which suggest that larval mixing in the plankton is less extensive than previously believed.