Phylogeography and Limited Genetic Connectivity in the Endangered Boring Giant Clam across the Coral Triangle

Abstract: The Coral Triangle is the global center of marine biodiversity; however, its coral reefs are critically threatened. Because of the bipartite life history of many marine species with sedentary adults and dispersive pelagic larvae, designing effective marine protected areas requires an understanding of patterns of larval dispersal and connectivity among geographically discrete populations. We used mtDNA sequence data to examine patterns of genetic connectivity in the boring giant clam (Tridacna crocea) in an effort to guide conservation efforts within the Coral Triangle. We collected an approximately 485 base pair fragment of mtDNA cytochrome c oxidase 1 (CO1) from 414 individuals at 26 sites across Indonesia. Genetic structure was strong between regions (φ_ST=0.549, p < 0.00001) with 3 strongly supported clades: one restricted to western Sumatra, another distributed across central Indonesia, and a third limited to eastern Indonesia and Papua. Even within the single largest clade, small but significant genetic structure was documented (φ_ST=0.069, p < 0.00001), which indicates limited gene flow within and among phylogeographic regions. Significant patterns of isolation by distance indicated an average dispersal distance of only 25–50 km, which is far below dispersal predictions of 406–708 km derived from estimates of passive dispersal over 10 days via surface currents. The strong regional genetic structure we found indicates potent limits to genetic and demographic connectivity for this species throughout the Coral Triangle and provides a regional context for conservation planning. The recovery of 3 distinct evolutionarily significant units within a well‐studied taxonomic group suggests that biodiversity in this region may be significantly underestimated and that Tridacna taxa may be more endangered than currently recognized.     

Life history, ecology and the biogeography of strong genetic breaks among 15 species of Pacific rockfish, Sebastes

Strong genetic change over short spatial scales is surprising among marine species with high dispersal potential. Concordant breaks among several species signals a role for geographic barriers to dispersal. Along the coast of California, such breaks have not been seen across the biogeographic barrier of Point Conception, but other potential geographic boundaries have been surveyed less often. We tested for strong-population structure in 11 species of Sebastes sampled across two regions containing potential dispersal barriers, and conducted a meta-analysis including four additional species. We show two strong breaks north of Monterey Bay, spanning an oceanographic gradient and an upwelling jet. Moderate genetic structure is just as common in the north as it is in the south, across the biogeographic break at Point Conception. Gene flow is generally higher among deep-water species, but these conclusions are confounded by phylogeny. Species in the subgenus Sebastosomus have higher structure than those in the subgenus Pteropodus, despite having larvae with longer pelagic phases. Differences in settlement behavior in the face of ocean currents might help explain these differences. Across similar species across the same coastal environment, we document a wide variety of patterns in gene flow, suggesting that interaction of individual species traits such as settlement behavior with environmental factors such as oceanography can strongly impact population structure.     

Integrating larval connectivity into the marine conservation decision-making process across spatial scales

Larval dispersal connectivity is typically integrated into spatial conservation decisions at regional or national scales, but implementing agencies struggle with translating these methods to local scales. We used larval dispersal connectivity at regional (hundreds of kilometers) and local (tens of kilometers) scales to aid in design of networks of no-take reserves in Southeast Sulawesi, Indonesia. We used Marxan with Connectivity informed by biophysical larval dispersal models and remotely sensed coral reef habitat data to design marine reserve networks for 4 commercially important reef species across the region. We complemented regional spatial prioritization with decision trees that combined network-based connectivity metrics and habitat quality to design reserve boundaries locally. Decision trees were used in consensus-based workshops with stakeholders to qualitatively assess site desirability, and Marxan was used to identify areas for subsequent network expansion. Priority areas for protection and expected benefits differed among species, with little overlap in reserve network solutions. Because reef quality varied considerably across reefs, we suggest reef degradation must inform the interpretation of larval dispersal patterns and the conservation benefits achievable from protecting reefs. Our methods can be readily applied by conservation practitioners, in this region and elsewhere, to integrate connectivity data across multiple spatial scales.     

Oceanic variability and coastal topography shape genetic structure in a long-dispersing sea urchin

Understanding the scale of marine population connectivity is critical for the conservation and sustainable management of marine resources. For many marine species adults are benthic and relatively immobile, so patterns of larval dispersal and recruitment provide the key to understanding marine population connectivity. Contrary to previous expectations, recent studies have often detected unexpectedly low dispersal and fine-scale population structure in the sea, leading to a paradigm shift in how marine systems are viewed. Nonetheless, the link between fine-scale marine population structure and the underlying physical and biological processes has not been made. Here we show that patterns of genetic structure and population connectivity in the broadcast-spawning and long-distance dispersing sea urchin Centrostephanus rodgersii are influenced by physical oceanographic and geographic variables. Despite weak genetic differentiation and no isolation-by-distance over thousands of kilometers among samples from eastern Australia and northern New Zealand, fine-scale genetic structure was associated with sea surface temperature (SST) variability and geography along the southeastern Australian coast. The zone of high SST variability is characterized by periodic shedding of eddies from the East Australian Current, and we suggest that ocean current circulation may, through its influence on larval transport and recruitment, interact with the genetic consequences of large variance in individual reproductive success to generate patterns of fine-scale patchy genetic structure. If proven consistent across species, our findings suggest that the optimal scale for fisheries management and reserve design should vary among localities in relation to regional oceanographic variability and coastal geography.     

Correlation between pelagic larval duration and realised dispersal: long-distance genetic connectivity between northern New Zealand and the Kermadec Islands archipelago

The extent to which marine populations are “open” (panmixia) or “closed” (self-recruitment) remains a matter of much debate, with recent reports of high levels of genetic differentiation and self-recruitment among populations of numerous species separated by short geographic. However, the interpretation of patterns of gene flow (connectivity) is often based on a stepping stone model of dispersal that can genetically homogenise even distant populations and blur genetic patterns that may better reflect realised dispersal. One way in which realised long-distance dispersal can be accurately determined is by examination of gene flow of taxa between isolated archipelagos and a mainland where there is no possible stepping stone dispersal across the open ocean. We investigated the genetic structuring of populations of the intertidal gastropod Nerita melanotragus from the subtropical Kermadec Islands and temperate New Zealand’s North Island (the mainland), separated by 750 km of open ocean and characterised by contrasting environmental conditions. Analyses of seven microsatellite markers revealed an absence of genetic structuring with low F _ST and Jost’s D values (from 0.000 to 0.007 and from 0.000 to 0.015, respectively) over large geographic distances and no evidence of isolation by distance among all populations. These results indicate that the realised dispersal of N. melanotragus is of at least 750 km, this species exhibits a very “open” form of connectivity and its larvae exhibit sufficient phenotypic plasticity to settle successfully in different environmental conditions, ranging from subtropical to cool temperate.     

Multiscale patterns of genetic structure in a marine snail (Solenosteira macrospira) without pelagic dispersal

The northern Gulf of California (NGC) is one of the most dynamic and productive marine ecosystems in the world, yet knowledge about population connectivity and dispersal patterns is lacking for many of its resident species. Using nuclear and mitochondrial markers, we investigated the effects of open water, geographical distance and suitable habitat on patterns of genetic structure of Solenosteira macrospira, a benthic buccinid whelk with direct development. We collected samples in April 2004, 2005 and May 2007 from the upper NGC (31°34.39″N, 114°44.45″W). Phylogenetic analyses, hierarchical analyses of variance and Bayesian assignment tests substantiated a break between the east and west coasts. Genetic distance between population pairs increased with geographical distance, but only when assuming a U-shaped dispersal pathway over the open water of the NGC. Given S. macrospira’s association with rocky intertidal habitats, and its limited dispersal potential, we assumed that the geographical distribution of rocky habitat would play a significant role in genetic differentiation of S. macrospira. Nevertheless, populations separated by sand were more similar than populations separated by rocks. The influence of open water, geographical distance and suitable habitat (rocks vs. sand) also varied significantly across different genetic markers that presumably evolve at different rates. Specifically, the more rapidly evolving nuclear microsatellites suggested that physical transport processes strongly influence genetic differentiation on contemporary time scales, even in a species with direct benthic development. This underscores the strong, and potentially homogenizing, effect of present-day ocean circulation patterns in the NGC.     

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.     

How do dispersal costs and habitat selection influence realized population connectivity?

Despite the importance of dispersal for population connectivity, dispersal is often costly to the individual. A major impediment to understanding connectivity has been a lack of data combining the movement of individuals and their survival to reproduction in the new habitat (realized connectivity). Although mortality often occurs during dispersal (an immediate cost), in many organisms costs are paid after dispersal (deferred costs). It is unclear how such deferred costs influence the mismatch between dispersal and realized connectivity. Through a series of experiments in the field and laboratory, we estimated both direct and indirect deferred costs in a marine bryozoan (Bugula neritina). We then used the empirical data to parameterize a theoretical model in order to formalize predictions about how dispersal costs influence realized connectivity. Individuals were more likely to colonize poor-quality habitat after prolonged dispersal durations. Individuals that colonized poor-quality habitat performed poorly after colonization because of some property of the habitat (an indirect deferred cost) rather than from prolonged dispersal per se (a direct deferred cost). Our theoretical model predicted that indirect deferred costs could result in nonlinear mismatches between spatial patterns of potential and realized connectivity. The deferred costs of dispersal are likely to be crucial for determining how well patterns of dispersal reflect realized connectivity. Ignoring these deferred costs could lead to inaccurate predictions of spatial population dynamics.     

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.     

Mode of reproduction,recruitment, and genetic subdivision in the brooding sponge<Emphasis Type="Italic"> Haliclona</Emphasis> sp.

Sponges display a variety of reproductive strategies that have the potential to influence population genetic structure. Histological examination of ten reproductive individuals of the Western Australian sponge Haliclona sp. showed that this species broods embryonic larvae that are potentially limited in dispersal capabilities. Because sponges have the potential to propagate in a number of modes, allozyme electrophoresis was used to assess the relative importance of asexual and sexual reproduction to recruitment, and to quantify genetic subdivision over different spatial scales. Tissue samples from 227 sponges were collected from reefs within two areas 400 km apart: Hamelin Bay and Rottnest Island. Contrary to expectations for highly clonal populations, genotypic diversity within sites was high, no linkage disequilibrium was found, and there was no evidence of genotypic clustering within reefs. There was no genetic evidence that asexual reproduction is important for the maintenance of populations. Genetic comparisons were consistent with mixing of sexually produced recruits within reefs, on a scale up to a few hundred metres, but significant genetic subdivision between reefs (F_ST=0.069 at Hamelin Bay, 0.130 at Rottnest Island) indicated that water gaps of several hundred metres are effective at preventing dispersal. Subdivision between the two areas, separated by 400 km, was moderately greater (F_ST=0.142) than within, but the same alleles were predominant in the two areas. These genetic patterns are consistent with limited dispersal capabilities of brooded larvae.Communicated by G.F. Humphrey, Sydney     

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.     

Two markers and one history: phylogeography of the edible common sea urchin Paracentrotus lividus in the Lusitanian region

Benthic marine invertebrates with long-lived larvae are believed to have dispersal capabilities that contribute to maintaining genetic uniformity among populations over large geographical scales. However, both hydrological and biological factors may limit the actual dispersal of such larvae. We studied the population genetic structure of the edible common sea urchin Paracentrotus lividus (Lamarck, 1816), to explore its dispersal patterns in the Atlanto-Mediterranean region and, more specifically, to ascertain the role of the Strait of Gibraltar in shaping the genetic structure of this species. For this purpose, we analysed 158 individuals for the mitochondrial 16S rRNA gene and 151 of these for the nuclear single-copy intron adenine nucleotide transporter (ANT) from 16 localities from the Atlantic and Mediterranean basins, spanning over 4,000 km. Mitochondrial 16S rRNA shows higher genetic diversity in the Mediterranean than in the Atlantic and reveals a sharp break between the populations of both basins, probably as a consequence of the barrier imposed by the Almería–Orán hydrological front, situated east of the Strait of Gibraltar. Both markers suggest that a recent population expansion has taken place in both basins, most probably following the Messinian salinity crisis. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effects of habitat fragmentation on the genetic structure and connectivity of the black-lipped pearl oyster Pinctada margaritifera populations in French Polynesia

Habitat destruction leading to increased fragmentation is detrimental to species by reducing population size and genetic diversity and by restraining population connectivity. However, little is known about the effects of naturally fragmented habitats on wild populations, especially when it comes to marine benthic invertebrates with long pelagic larval duration. In this framework, we investigated the connectivity and genetic diversity variation among nine wild populations of the black-lipped pearl oyster, Pinctada margaritifera, throughout French Polynesia using ten microsatellite DNA markers. Despite the naturally fragmented habitat (South Pacific oceanic islands), we found high values of genetic diversity and population admixture, indicating connectivity at small and large spatial scales within sampled sites of the Tuamotu, and between the Society and Tuamotu Archipelagos. In the meantime, habitat geomorphology increased genetic drift in populations occurring in small, closed lagoons. Significant genetic structure not correlated to geographic distance was detected mainly between closed and open lagoons. The Marquesas Islands hosted the most divergent populations, likely a result of vicariance. Our results also highlight that migration patterns among lagoons are not symmetrical. Altogether, the general pattern of gene flow, nonsymmetrical migration rates among populations, absence of isolation by distance and absence of recent extinction events found in our study strongly suggest that P. margaritifera populations of French Polynesia follow an asymmetrical island model of dispersal.     

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.     

Modelling the effect of in-stream and overland dispersal on gene flow in river networks

Modelling gene flow across natural landscapes is a current challenge of population genetics. Models are essential to make clear predictions about conditions that cause genetic differentiation or maintain connectivity between populations. River networks are a special case of landscape matrix. They represent stretches of habitat connected according to a branching pattern where dispersal is usually limited to upstream or downstream movements. Because of their peculiar topology, and the increasing concern about conservation issues in hydrosystems, there has been a recent revival of interest in modelling dispersal in river networks. Network complexity has been shown to influence global population differentiation. However, geometric characteristics are likely to interact with the way individuals move across space. Studies have focused on in-stream movements. None of the work published so far took into consideration the ability of many species to disperse overland between branches of the same network though. We predicted that the relative contribution of these two dispersal modalities (in-stream and overland) would affect the overall genetic structure. We simulated dispersal in synthetic river networks using an individual-based model. We tested the effect of dispersal modalities, i.e. the ratio of overland/in-stream dispersal, and two geometric parameters, bifurcation angle between branches and network complexity. Data revealed that if geometrical parameters affected population differentiation, dispersal parameters had the strongest effect. Interestingly, we observed a quadratic relationship between p the proportion of overland dispersers and population differentiation. We interpret this U-shape pattern as a balance between isolation by distance caused by in-stream movements at low values of p and intense migrant exchanges within the same branching unit at high values of p. Our study is the first attempt to model out-of-network movements. It clearly shows that both geometric and dispersal parameters interact. Both should be taken into consideration in order to refine predictions about dispersal and gene flow in river network.     

Linking life history to landscape for threatened species conservation in a multiuse region

Landscape-scale conservation that considers metapopulation dynamics will be essential for preventing declines of species facing multiple threats to their survival. Toward this end, we developed a novel approach that combines occurrence records, spatial–environmental data, and genetic information to model habitat, connectivity, and patterns of genetic structure and link spatial attributes to underlying ecological mechanisms. Using the threatened northern quoll (Dasyurus hallucatus) as a case study, we applied this approach to address the need for conservation decision-making tools that promote resilient metapopulations of this threatened species in the Pilbara, Western Australia, a multiuse landscape that is a hotspot for biodiversity and mining. Habitat and connectivity were predicted by different landscape characteristics. Whereas habitat suitability was overwhelmingly driven by terrain ruggedness, dispersal was facilitated by proximity to watercourses. Although there is limited evidence for major physical barriers in the Pilbara, areas with high silt and clay content (i.e., alluvial and hardpan plains) showed high resistance to dispersal. Climate subtlety shaped distributions and patterns of genetic turnover, suggesting the potential for local adaptation. By understanding these spatial–environmental associations and linking them to life-history and metapopulation dynamics, we highlight opportunities to provide targeted species management. To support this, we have created habitat, connectivity, and genetic uniqueness maps for conservation decision-making in the region. These tools have the potential to provide a more holistic approach to conservation in multiuse landscapes globally.     

Multi-marker estimate of genetic connectivity of sole (Solea solea) in the North-East Atlantic Ocean

A thorough knowledge on the genetic connectivity of marine populations is important for fisheries management and conservation. Using a dense population sampling design and two types of neutral molecular markers (10 nuclear microsatellite loci and a mtDNA cytochrome b fragment), we inferred the genetic connectivity among the main known spawning grounds of sole (Solea solea L.) in the North-East Atlantic Ocean. The results revealed a clear genetic structure for sole in the North-East Atlantic Ocean with at least three different populations, namely the Kattegat/Skagerrak region, the North Sea and the Bay of Biscay, and with indications for a fourth population, namely the Irish/Celtic Sea. The lack of genetically meaningful differences between biological populations within the southern North Sea is likely due to a large effective population size and sufficient connection (gene flow) between populations. Nevertheless, an isolation-by-distance pattern was found based on microsatellite genotyping, while no such pattern was observed with the cytochrome b marker, indicating an historical pattern prevailing in the latter marker. Our results demonstrate the importance of a combined multi-marker approach to understand the connectivity among marine populations at region scales.     

Dispersal ecology of deadwood organisms and connectivity conservation

Limited knowledge of dispersal for most organisms hampers effective connectivity conservation in fragmented landscapes. In forest ecosystems, deadwood‐dependent organisms (i.e., saproxylics) are negatively affected by forest management and degradation globally. We reviewed empirically established dispersal ecology of saproxylic insects and fungi. We focused on direct studies (e.g., mark‐recapture, radiotelemetry), field experiments, and population genetic analyses. We found 2 somewhat opposite results. Based on direct methods and experiments, dispersal is limited to within a few kilometers, whereas genetic studies showed little genetic structure over tens of kilometers, which indicates long‐distance dispersal. The extent of direct dispersal studies and field experiments was small and thus these studies could not have detected long‐distance dispersal. Particularly for fungi, more studies at management‐relevant scales (1–10 km) are needed. Genetic researchers used outdated markers, investigated few loci, and faced the inherent difficulties of inferring dispersal from genetic population structure. Although there were systematic and species‐specific differences in dispersal ability (fungi are better dispersers than insects), it seems that for both groups colonization and establishment, not dispersal per se, are limiting their occurrence at management‐relevant scales. Because most studies were on forest landscapes in Europe, particularly the boreal region, more data are needed from nonforested landscapes in which fragmentation effects are likely to be more pronounced. Given the potential for long‐distance dispersal and the logical necessity of habitat area being a more fundamental landscape attribute than the spatial arrangement of habitat patches (i.e., connectivity sensu strict), retaining high‐quality deadwood habitat is more important for saproxylic insects and fungi than explicit connectivity conservation in many cases.     

A panmictic fiddler crab from the coast of Brazil? Impact of divergent ocean currents and larval dispersal potential on genetic and morphological variation in Uca maracoani

Marine species tend to exhibit relatively less population structuring than terrestrial species owing to fewer barriers to gene flow and increased connectivity resulting from greater dispersal abilities. Thus, in many cases, life history plays a more important role in phylogeography of marine taxa than do oceanographic features. Littoral species are of particular phylogeographic interest because they possess life histories that have both marine and terrestrial characteristics. This study evaluates the synergistic impact of divergent ocean currents and a high larval dispersal potential on the phylogeography of the fiddler crab, Uca maracoani, distributed along the coast of Brazil. Patterns of genetic variation were assessed with sequence data for a portion of the mitochondrial COI gene and AFLPs. Geometric morphometric techniques were used to evaluate morphological variation. Results revealed a lack of discernible genetic subdivision. However, geometric morphometrics showed statistically significant morphological differentiation. The absence of a clear phylogeographic pattern appears to be determined primarily by life history characteristics permitting a high level of connectivity. One, or a combination of several factors, may explain the incongruity between genetic and morphologic signatures, including phenotypic plasticity, incomplete lineage sorting, or recent and ongoing genetic divergence.