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.     

Genetic connectivity of the ecosystem engineer Perumytilus purpuratus north to the 32°S southeast Pacific ecological discontinuity

Connectivity in benthic marine animals with complex life cycles occurs primarily during the pelagic larval stage and depends deterministically on oceanographic dynamics. The scale of such larval dispersal is highly uncertain due the difficulty of direct measurement and poor knowledge of larval dynamics and ocean flow variability. This study characterizes the pattern of genetic connectivity in the ecosystem engineer Perumytilus purpuratus between latitudes 23°S and 33°S, which includes the ecological discontinuity reported for many taxa north to 32°S at the southeast Pacific. The genetic discontinuity observed in P. purpuratus around 26°S is described herein while that detected at 28°S is in line with the ecological discontinuity (in coverage, recruitment and density) previously reported for this mussel between 28°S and 32°S. Both discontinuities delimitate two major gene pools upon Bayesian inferences on geographical variation of five microsatellite loci. Interestingly, marker Pepu1 was responsible for most variation between pools and was potentially under selection. In fact, inferences excluding Pepu1 produced a single gene pool (k = 1) in central-northern Chile. The IBD connectivity pattern observed among P. purpuratus beds distributed in the interval 23°S–33°S is congruent with processes driven by larval dynamics, and the dominant equatorward Humboldt Current along a coast largely unaffected by ice during the last Pleistocene glaciation. However, the selective scenario unveiled by microsatellite Pepu1 inside the 25°S–28°S ecological discontinuity is consistent with selective processes associated with specific mesoscale properties operating in this area. This study highlights the usefulness of integrating different oceanographic scales, ecological data and population genetics to better understand connectivity of benthic marine species.     

Predicting regional spread of non-native species using oceanographic models: validation and identification of gaps

Predicting spread is a central goal of invasion ecology. Within marine systems, researchers have increasingly made use of oceanographic circulation models to estimate currents and track species dispersal. However, the accuracy of these models for predicting biological patterns, particularly for non-native species, has generally not been validated. Particularly, we wished to examine the ability of models to predict physical and biological processes, which jointly determine the spread of marine larval organisms. We conducted two empirical studies—a recruitment study and a drift card study—along the coast of New England, USA, focusing on two invaders of concern—the European green crab (Carcinus maenas) and the Asian shore crab (Hemigrapsus sanguineus), to explicitly evaluate the ability of oceanographic models to predict patterns of spread. We used data from the large-scale drift card study to validate our ability to capture dispersal patterns driven purely by physical processes. Next, we conducted a recruitment study to evaluate our ability to reproduce patterns of biological dispersal. We were generally capable of reproducing drift cards patterns—suggesting that the physical mechanics in the model were predictive. However, predicted biological patterns were inconsistent—we were able to predict dispersal patterns for H. sanguineus but not for C. maenas. Our results highlight the importance of validating models and suggest that more work is necessary before we can reliably use oceanographic models to predict biological spread of intertidal organisms.     

Larval supply and juvenile recruitment of coral reef fishes to marine reserves and non-reserves of the upper Florida Keys,USA

For marine organisms, decoupling between the planktonic larval stage and the benthic-associated juvenile stage can lead to variable patterns of population replenishment, which have the potential to influence the effectiveness of marine reserves. We measured spatial and temporal variability in larval supply and recruitment of fishes to coral reefs of different protection levels and tested whether protection level influenced the relationship between supply and recruitment. We sampled pre-settlement larvae and newly settled recruits from four reefs (two reserves and two non-reserves) in the Florida Keys National Marine Sanctuary, USA. Replicate point measures of larval supply over 14 months and 17 monthly measurements of recruitment varied significantly among months and sites. Sites with the same protection level had significantly different patterns of larval supply as well as larval and recruit diversity, but recruitment magnitude differed only by protection level, where densities were greater at reserves. Differences in larval supply among sites included two particularly large peaks in larval abundance at one site, possibly associated with the observed passage of small-scale oceanographic features. To examine whether relationships between larval supply and recruitment varied by protection level, we selected one species that was present in both the light trap samples and the monthly recruitment surveys. Recruitment of the bicolor damselfish Stegastes partitus was significantly and positively related to larval supply at three of the four sites thus, protection level did not influence this linkage. Since local variability among sites can lead to spatial differences in population replenishment, characterization of larval supply and recruitment to potential marine reserve sites may help to identify optimal locations in a region and contribute to more effective reserve design.     

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.     

Assessing the current state of ecological connectivity in a large marine protected area system

The establishment of marine protected areas (MPAs) is a critical step in ensuring the continued persistence of marine biodiversity. Although the area protected in MPAs is growing, the movement of individuals (or larvae) among MPAs, termed connectivity, has only recently been included as an objective of many MPAs. As such, assessing connectivity is often neglected or oversimplified in the planning process. For promoting population persistence, it is important to ensure that protected areas in a system are functionally connected through dispersal or adult movement. We devised a multi-species model of larval dispersal for the Australian marine environment to evaluate how much local scale connectivity is protected in MPAs and determine whether the extensive system of MPAs truly functions as a network. We focused on non-migratory species with simplified larval behaviors (i.e., passive larval dispersal) (e.g., no explicit vertical migration) as an illustration. Of all the MPAs analyzed (approximately 2.7 million km²), outside the Great Barrier Reef and Ningaloo Reef, <50% of MPAs (46-80% of total MPA area depending on the species considered) were functionally connected. Our results suggest that Australia's MPA system cannot be referred to as a single network, but rather a collection of numerous smaller networks delineated by natural breaks in the connectivity of reef habitat. Depending on the dispersal capacity of the taxa of interest, there may be between 25 and 47 individual ecological networks distributed across the Australian marine environment. The need to first assess the underlying natural connectivity of a study system prior to implementing new MPAs represents a key research priority for strategically enlarging MPA networks. Our findings highlight the benefits of integrating multi-species connectivity into conservation planning to identify opportunities to better incorporate connectivity into the design of MPA systems and thus to increase their capacity to support long-term, sustainable biodiversity outcomes.     

Fine-scale genetic structure in short-beaked common dolphins (<Emphasis Type="Italic">Delphinus</Emphasis><Emphasis Type="Italic">delphis</Emphasis>) along the East Australian Current

Oceanographic processes play a significant role in shaping the genetic structure of marine populations, but it is less clear whether they affect genetic differentiation of highly mobile vertebrates. We used microsatellite markers and mtDNA control region sequences to investigate the spatial genetic structure of short-beaked common dolphins (Delphinus delphis) in southeastern Australia, a region characterised by complex oceanographic conditions associated with the East Australian Current (EAC). A total of 115 biopsy samples of dolphins were collected from six localities spanning approximately 1,000 km of the New South Wales (NSW) coastline. We found evidence for contrasting genetic diversity and fine-scale genetic structure, characterised by three genetically differentiated populations with varying levels of admixture. Spatial genetic structure was not explained by a model of isolation by distance, instead it coincides with main patterns of oceanographic variation along the EAC. We propose that common dolphins along the EAC may be adapted to three water masses recently characterised in this region.     

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.     

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.     

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.     

Currents connecting communities: nearshore community similarity and ocean circulation

Understanding the mechanisms that create spatial heterogeneity in species distributions is fundamental to ecology. For nearshore marine systems, most species have a pelagic larval stage where dispersal is strongly influenced by patterns of ocean circulation. Concomitantly, nearshore habitats and the local environment are also influenced by ocean circulation. Because of the shared dependence on the seascape, distinguishing the relative importance of the local environment from regional patterns of dispersal for community structure remains a challenge. Here, we quantify the "oceanographic distance" and "oceanographic asymmetry" between nearshore sites using ocean circulation modeling results. These novel metrics quantify spatial separation based on realistic patterns of ocean circulation, and we explore their explanatory power for intertidal and subtidal community similarity in the Southern California Bight. We find that these metrics show significant correspondence with patterns of community similarity and that their combined explanatory power exceeds that of the thermal structure of the domain. Our approach identifies the unique influence of ocean circulation on community structure and provides evidence for oceanographically mediated dispersal limitation in nearshore marine communities.     

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.     

Phylogeography of the Antarctic planktotrophic brittle star Ophionotus victoriae reveals genetic structure inconsistent with early life history

In the marine environment, connectivity is influenced by physical oceanography as well as life history and behavioral traits, which in combination with historical events such as geology, physical oceanography, and climate, determine population structure. The Antarctic brittle star Ophionotus victoriae develops via a feeding planktonic larval stage, and therefore has potential for long-distance dispersal throughout its Antarctic/subantarctic range. To evaluate this hypothesis, phylogeography of this ecologically dominant species was elucidated by sequence analysis of two mtDNA genes from individuals collected throughout the Antarctic Peninsula and from two subantarctic islands. Counter to expectations of genetic homogeneity, mtDNA data revealed substantial levels of genetic differentiation as well as diversity. Although there were some genetically homogeneous populations, such as those throughout Bransfield Strait, we found O. victoriae to have significant population structure throughout much of the Antarctic Peninsula, with evidence of potential cryptic speciation between the western and eastern Antarctic Peninsula. Furthermore, Antarctic Peninsula populations were genetically distinct from subantarctic island populations. The low levels of connectivity implied for O. victoriae contrast with those found for many other Antarctic benthic taxa, and suggest a complex interplay between oceanography, recent climate history, and larval ecology.     

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.     

Microsatellite DNA markers and morphometrics reveal a complex population structure in a merobenthic octopus species (<Emphasis Type="Italic">Octopus maorum</Emphasis>) in south-east Australia and New Zealand

Five polymorphic microsatellite loci were developed and then used to assess the population genetic structure of a commercially harvested merobenthic octopus species (Octopus maorum) in south-east Australian and New Zealand (NZ) waters. Beak and stylet morphometrics were also used to assess population differentiation in conjunction with the genetic data. Genetic variation across all loci and all sampled populations was very high (mean number alleles = 15, mean expected heterozygosity = 0.85). Microsatellites revealed significant genetic structuring (overall F _ST = 0.024, p < 0.001), which did not fit an isolation-by-distance model of population differentiation. Divergence was observed between Australian and NZ populations, between South Australia and north-east Tasmania, and between two relatively proximate Tasmanian sites. South Australian and southern Tasmanian populations were genetically homogeneous, indicating a level of connectivity on a scale of 1,500 km. Morphometric data also indicated significant differences between Australian and NZ populations. The patterns of population structuring identified can be explained largely in relation to regional oceanographic features.     

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.     

Invasion dynamics of the European shore crab,<Emphasis Type="Italic"> Carcinus maenas</Emphasis>, in Australia

In Australia and most other invaded locations, rates of range expansion by the European shore crab, Carcinus maenas, are typically only a few kilometres per year, despite a planktonic duration upwards of 50 days and off-shore larval development. This relatively static distribution is punctuated by rare episodes of long-distance and large-scale spread, some of which appear to be related to unusual oceanographic conditions and some of which are likely to be human assisted. These observations suggest, first, that long planktonic duration and off-shore development in a marine invertebrate does not preclude very localised recruitment, and, second, that this recruitment norm may be punctuated by brief episodes of wide scale mixing of propagules. Punctuated dispersal has previously been suggested to account for large-scale biogeographic patterns of distribution and speciation, but may also have implications for the processes that stabilise structured spatial metapopulations.     

The influence of contrasting life history traits and oceanic processes on genetic structuring of rabbitfish populations Siganus argenteus and Siganus fuscescens along the eastern Philippine coasts

Genetic variability and structuring of rabbitfish populations with contrasting life histories, Siganus argenteus and Siganus fuscescens were determined using allozyme analysis. A total of 13–14 polymorphic loci were examined from samples collected in 2002 and 2003 from eight reefs representing 25 populations north (Kuroshio Current) and south (Mindanao Current) of the bifurcation of the North Equatorial Current along the eastern Philippine coast. S. fuscescens populations (H _OBS = 0.085) showed higher heterozygosity than S. argenteus (H _OBS = 0.053), consistent with predictions of the neutral theory for demersal egg spawners compared to pelagic egg spawners. The generally lower genetic variability of Kuroshio populations may be due to greater environmental disturbance affecting larval mortality and recruitment success. There was no significant overall population genetic structuring for S. argenteus (F _ST = 0.01485, P > 0.05) compared to S. fuscescens (F _ST = 0.03275, P < 0.05). The latter species showed highly significant genetic structuring among Kuroshio and Mindanao Current populations in both 2002 and 2003 (F _CT = 0.08120, P < 0.05; F _CT = 0.07500, P < 0.05, respectively), as well as among populations within regions. This conforms to expectations of correlations between observed population genetic structure and life history features related to dispersal potential and gene flow. However, there were significant temporal (i.e., 2002 vs. 2003 samples) genetic variations for both S. fuscescens (F _CT = 0.08542, P < 0.05) and S. argenteus (F _CT = 0.06330, P < 0.05), which may reflect interannual variability in recruitment success. Differences in population spatial genetic patterns between the two reef fish species suggest that broad scale physical factors (e.g. NEC bifurcation) and regional environmental perturbations (e.g. incidence of typhoons) affect population genetic structure of sympatric congeneric species with different life histories differently. Finer scale ecological processes, which affect larval dispersal and recruitment (e.g., local hydrographic features, distribution of habitats), particularly in the Mindanao Current region, exert more influence on structuring populations of S. fuscescens.     

Geographic structure in Alaskan Pacific ocean perch (<Emphasis Type="Italic">Sebastes alutus</Emphasis>) indicates limited lifetime dispersal

Prevailing oceanographic processes, pelagic larvae, adult mobility, and large populations of many marine species often leads to the assumption of wide-ranging populations. Applying this assumption to more localized populations can lead to inappropriate conservation measures. The Pacific ocean perch (Sebastes alutus, POP) is economically and ecologically valuable, but little is known about its population structure and life history in Alaskan waters. Fourteen microsatellite loci were used to characterize geographic structure and connectivity of POP collections (1999–2005) sampled along the continental shelf break from Dixon Entrance to the Bering Sea. Despite opportunities for dispersal, there was significant, geographically related genetic structure (F _ST = 0.0123, P < 10⁻⁵). Adults appear to belong to neighborhoods at geographic scales less than 400 km, and possibly as small as 70 km, which indicates limited dispersal throughout their lives. The population structure observed has a finer geographic scale than current management, which suggests that measures for POP fisheries conservation should be revisited.