Genetic and morphological divergence in the purple sea urchin Paracentrotus lividus (Echinodermata, Echinoidea) across the African Mediterranean coast

The present investigation focuses on population structure analysis of the purple sea urchin Paracentrotus lividus across the African Mediterranean coast, with the main aim of assessing the influence of the Siculo-Tunisian Strait on gene flow disruption in this highly dispersive echinoid species. For this purpose, patterns of morphological and genetic variation were assessed among its populations from the western and eastern Mediterranean coasts. A total of 302 specimens from seven Tunisian sites were collected and examined for morphometric variability at twelve morphometric traits. Concordant results, inferred from CDA (canonical discriminant analyses), pairwise NPMANOVA (non parametric multivariate analysis of variance) comparisons and MDS (multidimensional scaling) plot, unveiled significant inter-population differences in the measured traits among the studied populations. Furthermore, the combined use of the one way ANOSIM (analysis of similarities) and the Discriminant/Hotelling analysis allowed unravelling two morphologically differentiated groups assigned to both western and eastern Mediterranean basins. The SIMPER (similarity percentages) routine analysis showed that total dry weight, test diameter and spine length were major contributors to the morphometric separation between locations and between groups. Pattern of phenotypic divergence discerned in P. lividus across the Siculo-Tunisian Strait is interestingly in congruence with that inferred from the genetic investigation of the purple sea urchin populations from the same region based on the analysis of the mtDNA COI (cytochrome oxidase I) gene in 314 specimens from nineteen locations covering a wider geographic transect, streching westward to the Algerian coast and eastward to the Libyan littoral. The specific haplotypic composition characterizing each Mediterranean basin, as inferred from the minimum spanning network, confirmed the geographic partioning of genetic variation, as revealed by F-statistics and AMOVA (analysis of molecular variance) analyses, yielding significant genetic differentiation between eastern and western Mediterranean populations. The newly detected phylogeographic patterns, observed for the first time in P. lividus throughout the explored distribution range, suggest the involvement of different biotic and abiotic processes in shaping such variation, and provide evidence that a large and geographically exhaustive dataset is necessary to unveil phylogeographic structure within widespread marine species, previously cathegorized as panmictic in part of their distribution range.     

West versus East Mediterranean Sea: origin and genetic differentiation of the sea cucumber Holothuria polii

We studied the genetic structure of the sea cucumber Holothuria (Roweothuria) polii (Delle Chiaje 1823) by analysing the mitochondrial DNA variation in two fragments of cytochrome oxidase I (COI) and 16S genes. Individuals were collected in seven locations along the Mediterranean Sea, which cover a wide range of the species distribution. We found high haplotype diversity for COI and moderate diversity for 16S, and low nucleotide diversity for both genes. Our results for the COI gene showed many recent and exclusive haplotypes with few mutational changes, suggesting recent or ongoing population expansion. The Western and Eastern Mediterranean populations exhibited slight but significant genetic differentiation (COI gene) with higher genetic diversity in the East. The most ancient haplotype was not present in the westernmost sampling location (SE Spain). The oldest expansion time was observed in Turkey, corresponding to mid‐Pleistocene. Turkey had also the highest genetic diversity (number of total and exclusive haplotypes, polymorphisms, haplotype and nucleotide diversity). This suggests that this region could be the origin of the subsequent colonizations through the Mediterranean Sea, a hypothesis that should be assessed with nuclear markers in future research.     

Lack of mitochondrial genetic structure in the endangered giant clam populations of Tridacna maxima (Bivalvia: Cardiidae: Tridacninae) across the Saudi Arabian coast

The present investigation focuses on population genetic structure analysis of the endangered giant clam species Tridacna maxima across part of the Red Sea,with the main aim of assessing the influence of postulated potential barriers to gene flow(i.e.,particular oceanographic features and marked environmental heterogeneity)on genetic connectivity among populations of this poorly dispersive bivalve species.For this purpose,a total of 44 specimens of T.maxima were collected from five sampling locations along the Saudi Arabian coast and examined for genetic variability at the considerably variable mitochondrial gene cytochrome c oxidase I(COI).Our results revealed lack of population subdivision and phylogeographic structure across the surveyed geographic spectrum,suggesting that neither the short pelagic larval dispersal nor the various postulated barriers to gene flow in the Red Sea can trigger the onset of marked genetic differentiation in T.maxima.Furthermore,the discerned shallow COI haplotype genealogy(exhibiting high haplotype diversity and low nucleotide diversity),associated with recent demographic and spatial expansion events,can be considered as residual effect of a recent evolutionary history of the species in the Red Sea.     

Significant population genetic structuring of the holoplanktic scyphozoan Pelagia noctiluca in the Atlantic Ocean

Pelagia noctiluca is thought to have a global distribution, yet our understanding of genetic connectivity across the range of this problem animal is poor. Here, we investigate the genetic structure of populations off southern Africa using mitochondrial COI and nuclear ITS1 and ITS2 genes, and compare the results to recent work conducted in the North-East Atlantic. Analyses showed significant differentiation between the southern and northern Atlantic population groups (COI Φ _st = 0.72, ITS2 Φ _st = 0.23, p < 0.001), which suggests historical rather than contemporary gene flow. Southern African samples showed high haplotypic (h = 1) and low nucleotide (π = 0.008) diversity, similar to those from Europe. Phylogenetic analyses suggest South African samples to have diverged earlier than those from the northern Atlantic.     

Genetic variability of the Lessepsian migrant mussel Brachidontes pharaonis (Bivalvia: Mytilidae) in Tunisia

The present study used two mitochondrial markers (16S rRNA and COI) to assess the genetic diversity of a newly founded Lessepsian migrant mussel, Brachidontes pharaonis, in Tunisian waters. The species appears to be restricted to only one population in Rades Harbour, in the northern part of the country. Phylogenetic analyses revealed the monophyly of B. pharaonis in Tunisia. Both molecular markers revealed high genetic variability of the B. pharaonis population. Haplotype networks and demographic analyses confirmed the recent expansion events within this population. Multiple human-mediated introduction events involving several founder populations and intensive population growth rates are probably the main causes of the high polymorphism observed within this invasive mollusc.     

Phylogeography of Ophryotrocha labronica (Polychaeta,Dorvilleidae) along the Italian coasts

Species of the genus Ophryotrocha are a well‐studied group of organisms but, despite the relatively large body of biological studies, little is known about their intra‐specific patterns of genetic diversity. In the present study, we analysed the patterns of genetic variation in samples of Ophryotrocha labronica (Polychaeta, Dorvilleidae) collected along the Italian coasts within three regions with different thermal regimes: the Northern Adriatic Sea (NAS), the Ligurian Sea (LS), and the South/Southeast Sicilian Sea (SS). A partial sequence of the cytochrome c oxidase subunit I (COI) gene was used as a genetic marker. An analysis of molecular variance (AMOVA) showed significant genetic differentiation between the NAS and the other regions. Conversely, little or no genetic structuring was found between the LS and the SS or amongst locations within a given region. A Bayesian phylogenetic tree and a median‐joining network provided evidence for the occurrence of two highly divergent genetic lineages characterized by a high average sequence divergence (17.2%, Kimura two‐parameter distance). The spatial patterns of genetic variation found in O. labronica may reflect the signature of past expansion events of the two genetic lineages. Although the high sequence divergence suggested that cryptic speciation within O. labronica may have occurred, other traits such as the absence of reproductive isolation, pattern of phenotypic variation and habitat specificity prompted us to regard the two groups as distinct COI lineages of O. labronica.     

Fine‐scale phylogeography reveals cryptic biodiversity in Pederson's cleaner shrimp,Ancylomenes pedersoni (Crustacea: Caridea: Palaemonidae), along the Florida Reef Tract

Populations of continuously distributed marine organisms that disperse via pelagic larvae are often assumed to exist in a state of genetic panmixia because of their potential ability for long‐distance dispersal. However, obligate symbionts may have more restricted gene flow due to recruitment limitations, making host specificity an important potential driver of biodiversity. To explore the tension between broad dispersal ability and limited recruitment potential, we used sequences of mtDNA cytochrome c oxidase subunit I (COI) to assess the fine‐scale phylogeography of the cleaner shrimp, Ancylomenes pedersoni, an obligate symbiont of sea anemones, along 300 km of the Florida Reef Tract (FRT). The results indicate high genetic diversity within A. pedersoni populations along the FRT. Pair‐wise ?_ST values indicate significant genetic structuring between northern (Fort Lauderdale and Upper Keys) and southwestern (Lower Keys) populations, with significant isolation by distance. Two divergent COI haplotype lineages were detected through statistical parsimony analysis: one evenly distributed across the entire FRT (Clade 1) and one found primarily in two sample localities in the Lower Keys (Clade 2). Phylogenetic analyses using 16S‐rDNA indicates that this genetic diversity is of paraphyletic origin, and possibly the result of multiple colonization events. These results reveal a complex demographic and evolutionary history for A. pedersoni populations and provide the first evidence of highly divergent intra‐specific lineages independently colonizing the FRT. Because the FRT is a highly impacted coral reef system, understanding phylogeographic patterns along it has value beyond documentation of the factors that generate genetic diversity in tropical reef systems: these data are critical for creating scientifically based management strategies.     

Patterns of genetic and morphometric diversity in the marbled crab (Pachygrapsus marmoratus, Brachyura, Grapsidae) populations across the Tunisian coast

The present study reports on population structure analysis of the marbled crabPachygrapsus marmoratus (Fabricius, 1787) from the Tunisian coast, an appropriate location to study biogeographical process...     

High gene flow promotes the genetic homogeneity of the fish goby Pomatoschistus marmoratus (Risso, 1810) from Mar Menor coastal lagoon and adjacent marine waters (Spain)

The extreme environmental variability of coastal lagoons suggests that physical and ecological factors could contribute to the genetic divergence among populations occurring in lagoon and open‐coast environments. In this study we analysed the genetic variability of lagoon and marine samples of the sand goby, Pomatoschistus marmoratus (Risso, 1810) (Pisces: Gobiidae), on the SW Spain coast. A fragment of mitochondrial DNA control region (570 bp) was sequenced for 196 individuals collected in five localities: Lo Pagan, Los Urrutias and Playa Honda (Mar Menor coastal lagoon), and Veneziola and Mazarrón (Mediterranean Sea). The total haplotype diversity was h = 0.9424 ± 0.0229, and the total nucleotide diversity was π = 0.0108 ± 0.0058. Among‐sample genetic differentiation was not significant and small‐scale patterns in the distribution of haplotypes were not apparent. Gene flow and dispersal‐related life history traits may account for low genetic structure at a small spatial scale. The high genetic diversity found in P. marmoratus increases its potential to adapt to changing conditions of the Mar Menor coastal lagoon.     

黄东海奇异指纹蛤Acila mirabilis和指纹蛤Acila divaricata群体遗传多样性及进化研究

指纹蛤属Acila贝类是我国黄东海重要的底栖生物,研究其遗传多样性及进化对了解这一海区环境的变化及与生物的关系具有重要作用。本研究通过线粒体COI基因标记,分析了黄东海常见的两个指纹蛤属物种—奇异指纹蛤Acila mirabilis和指纹蛤Acila divaricata的分化情况,结果表明这两个种的分化形成时间分别在3.71和4.27百万年前,处于上新世时期,我们推测冰期时海平面下降引起的物种栖息地的缩减以及黄海和东海环境条件的不同是导致物种分化的重要原因。通过群体遗传多样性分析,我们发现分布于黄海的4个奇异指纹蛤群体中,3500-10群体的遗传多样性水平最高,且群体遗传多样性自冷水团中心内侧至外侧呈递减趋势,推测这可能与这一群体对冷水团有较好的适应性有关。分布于东海的3个指纹蛤群体均检测到两个单倍型类群ZA和ZB,两类群的分化时间大约在64万年前,发生于更新世中期,我们认为冰期时海平面升降引起的群体地理隔离与二次接触是导致指纹蛤两个单倍型类群形成的主要原因。     

The genetic structure of the exotic ascidian Styela plicata (Tunicata) from Italian ports,with a re‐appraisal of its worldwide genetic pattern

The pleated ascidian Styela plicata (Lesueur, 1823) is a solitary species commonly found in ports and marinas around the world. It has been recorded in the Mediterranean region since the mid‐19th century. In the present work, the species’ genetic diversity was analysed, employing a 613‐bp portion of the mitochondrial cytochrome c oxidase subunit I (COI) gene from 149 individuals collected in 14 ports along Italian coasts at spatial scales ranging from 1 to approximately 2200 km. Haplotype and nucleotide diversity values were h = 0–0.933 (total h = 0.789) and π = 0–0.145 (total π = 0.0094), respectively. A general southward trend of increasing within‐population genetic diversity was observed. Analysis of molecular variance revealed significant genetic structuring but no significant differences were detected among basins, and no isolation by distance was found. Our data were integrated with the COI sequences available from previous studies and re‐analysed in order to investigate the possible routes of introduction of this ascidian into the Mediterranean Sea. The presence of the two COI haplogroups detected in previous molecular investigations on S. plicata at intercontinental spatial scale was confirmed in the Mediterranean Sea. The results revealed multiple introductions of S. plicata, although some locations appear to have experienced rapid expansion from few founding individuals with reduced genetic diversity. However, continuous introductions would confound the pattern deriving from single founder events and make it difficult to estimate the time needed for gene diffusion into established populations. This mixing of effects creates difficulties in understanding the past and current dynamics of this introduction, and managing this alien invasive ascidian whose genetic structure is continuously shuffled by vessel‐mediated transport.     

A pilot study of genetic differentiation between two phenotypes of a Mediterranean population of the bivalve Cerastoderma glaucum and genetic discrimination with other Cerastoderma glaucum and Cerastoderma edule populations outside the Mediterranean

The internal transcribed spacer 1 (ITS1) region of 29 individuals of Cerastoderma glaucum (from the Mediterranean Sea, the Atlantic and the North Sea) and 18 individuals of Cerastoderma edule (from the Atlantic and the North Sea) and the cytochrome c oxidase subunit I (COI) region of 43 individuals of C. glaucum (from the Mediterranean Sea, the Atlantic and the North Sea) and 9 individuals of C. edule (from the Atlantic and the North Sea) were PCR amplified and sequenced. The ITS1 sequences led us to describe 27 genetic haplotypes, while 21 genetic haplotypes were determined via the COI sequences. The ITS1 and the COI sequences revealed an important genetic variability within the Mediterranean population of C. glaucum, which contained two different phenotypes: orange foot and yellow foot. A genetic differentiation between the two phenotypes was revealed by the nucleotide diversity index (π) and is strengthened by the AMOVA analysis. This result leads us to suspect the presence of two different groups in the same population of C. glaucum. Nevertheless to be confirmed this hypothesis requires further studies using more locations and a larger number of sample sizes.     

中国近海外来囊藻(Colpomenia peregrina)种群遗传多样性研究

外来囊藻(Colpomeniaperegrina)是一种在潮间带分布广泛的褐藻,它能附着在牡蛎壳上进行跨海区扩散,因而是研究生物多样性变动和生物地理格局形成的良好模型。外来囊藻在中国沿海虽有分布,但关于其种群遗传多样性和分化研究的报道尚为空白。本研究采集了中国近海13个外来囊藻种群样本。通过扩增301条线粒体cox3序列,发现了26个单倍型;通过重建单倍型网络图和基于最大似然法与贝叶斯法重建系统进化树,发现外来囊藻种内遗传多样性较高, 13个种群划分为3个遗传世系,其中浙江南麂列岛的7个种群组成遗传世系A,辽宁和山东的样本则分化为遗传世系B和C。中国近海外来囊藻的这种种群遗传结构可能源于更新世末次盛冰期黄渤海、东海边缘海的环境变化和海平面的大幅下降。遗传世系A具有的较高单倍型多样性,其原因可能是南麂列岛独特的地理位置和潮间带环境的共同作用。     

中国沿海不同地区泥蚶(Tegillarca granosa)的遗传多样性分析

以采自海南海口、广西防城港、广西北海、广东湛江、福建漳州、山东荣成、山东威海7个地理群体62个泥蚶个体为材料,获取592bp线粒体COI基因片段序列,并进行遗传多样性及分化分析。多态性遗传参数统计显示:62个个体共检测出103个多态性位点,定义了26个单倍型;总群体单倍型多样性指数为0.834,核苷酸多样性指数为0.01665,平均核苷酸差异数为9.85772。7个群体均显示出较丰富的遗传多样性,群体内遗传距离及群体内遗传多样性参数显示中国沿海泥蚶遗传多样性由北向南呈升高趋势,而群体内遗传分化系数也呈现升高的趋势。基于26个单倍型COI序列构建的NJ树和UPGMA树以及基于群体间遗传距离构建的UPGMA树显示,荣成群体和威海群体亲缘关系较近,聚成一小支,而后与漳州群体相聚,然而南方类群并没有聚为独立的一支。     

Genetic characterisation of oyster populations along the north-eastern coast of Tunisia

The taxonomy of oysters has been traditionally based on characteristics of the shell. More recently, the analysis of protein and DNA polymorphism has provided a means to overcome difficulties in distinguishing the different species of oysters based solely on shell morphology. In order to identify oysters of the Tunisian north-east coast, we sequenced a 16S rRNA mitochondrial fragment from 68 oysters sampled from the Bizert Lagoon and the Gulf of Hammamet in northern Tunisia. Comparison of oyster 16S rRNA sequences available in GenBank showed the presence of both Ostreola stentina and Crassostrea gigas in our samples, which could not be detected on the basis of shell morphology only. These data confirmed that C. gigas, a non-native species, is now naturalised in the Bizert Lagoon. Furthermore, significant levels of genetic divergence among the 16S rRNA haplotypes from O. stentina populations have been observed. Specifically, the haplotypes found in the Bizert Lagoon are closer to those previously detected from Morocco and Portugal, whereas those in the Gulf of Hammamet are closer to the haplotypes from the south of Tunisia, with a divergence ranging from 2.1% to 2.7% between the northern and eastern Tunisian haplotypes. The possible impact of the Siculo-Tunisian Strait on the phylogeography of O. stentina is discussed.     

10个菲律宾蛤仔野生群体的遗传多样性研究

对我国沿海地区10个菲律宾蛤仔野生群体线粒体16S r RNA和COI基因部分序列进行测序,分别得到了长度为473bp和632bp的片段。结果表明,16S r RNA 193条序列A+T平均含量为66.6%,共检测到21个变异位点,193个个体具有22种单倍型;COI基因183条序列A+T平均含量为64.8%,共检测到126个变异位点,183个个体具有67种单倍型。基于群体间遗传距离利用Mega5.1软件构建10个群体的NJ树,聚类结果表明,大连群体和荣成群体聚为一支,其余8个群体聚为一支。AMOVA分析表明,大连群体和荣成群体间分化不显著,而荣成、大连群体与其余8个群体间的分化达到极显著水平(P0.01),说明我国沿海的菲律宾蛤仔野生群体存在一定的遗传分化。     

基于COI基因的中国沿海青蚶野生群体遗传结构及种群动态研究

本研究利用形态学方法和基于线粒体细胞色素氧化酶亚基Ⅰ(COⅠ)的遗传学方法分析了中国沿海青蚶(Barbatia virescens)6个地理群体的形态差异、遗传多样性、遗传结构及其种群历史动态。单因素方差分析(ANOVA)和Tukey检验表明,青蚶不同地理种群间表现出显著形态差异(P0.05)。经PCR扩增测序获得长度为587bp的COⅠ基因片段,112个个体共检测到18个多态性位点、17个单倍型,每个群体均有特有单倍型。青蚶群体的遗传多样性水平较低,总群体的平均单倍型多样度为0.5472,平均核苷酸多样度为0.000974。AMOVA分析表明,群体内个体间的遗传分化是青蚶群体遗传变异的主要原因,占87.40%。阳江群体与其他群体之间存在显著的低程度的遗传分化,而其他群体间的遗传分化不显著。单倍型网络关系图呈典型的星状拓扑结构,没有表现出显著的地理谱系结构。单倍型邻接树结果也没有明显分支,未呈现出地域性差异。中性检验和核苷酸不配对分析结果揭示青蚶种群历史上经历了群体扩事件,扩张时间约为26万年前。研究结果为青蚶资源的保护和可持续利用提供了基础资料。     

基于线粒体COI 基因的毛蚶群体遗传多样性

采用PCR技术,扩增了大连、乳山、烟台、舟山4个毛蚶(Scapharca subcrenata)地理群体共38个个体的线粒体COI基因部分序列,并分析了4个毛蚶群体的遗传多样性和系统发育关系。研究结果显示:38个毛蚶COI部分序列经处理得到长度均为625bp的基因片段,共分为30种单倍型;基于COI部分序列的分析结果,毛蚶4个地理群体总的变异位点为301个,多样性指数Pi为0.15048,平均核苷酸差异数为92.242,单倍型多样性指数S为241。聚类分析显示毛蚶大连群体、乳山群体和烟台群体具有高度的遗传多样性,3个群体交叉聚在一起,没有明显的群体分化;舟山群体单独聚为一支,与其他3个群体分化明显。研究表明,线粒体COI基因不能单独做为毛蚶大连、乳山和烟台群体的遗传标记,但可以作为毛蚶舟山群体的有效群体遗传标记,为线粒体COI基因在群体遗传学的应用提供了基础资料。     

Temperature effects on asexual reproduction of the scyphozoan Aurelia aurita s.l.: differences between exotic (Baltic and Red seas) and native (Mediterranean Sea) populations

Massive occurrences of jellyfish can cause direct impacts on the economy, especially on tourism and commercial fisheries. Translocation of jellyfish species by humans has caused damaging blooms in new habitats. Aurelia aurita s.l. has been introduced in many locations around the world. To test the potential success of Au. aurita s.l. in various habitats, scyphistomae from different climatic locations (Mediterranean, Red and Baltic Seas) were cultured individually for 201 days at three temperatures (14, 21 and 28 °C) with the same salinity, food and light. We tested the null hypotheses that there were no differences in survival or asexual reproduction (budding and strobilation) amongst populations [native (Mediterranean) and exotic (Red and Baltic)]. Survival of the three scyphistoma populations did not differ significantly across temperatures; however, the Red Sea group had lower survival at all temperatures than did the other populations. Most individuals strobilated at 14 °C. Red Sea scyphistomae strobilated more quickly than Baltic and Mediterranean Sea scyphistomae and produced the fewest ephyrae, whereas Baltic Sea scyphistomae produced the most. Our results indicate that Au. aurita from the Baltic or Red Seas introduced into the Northwest Mediterranean Sea would potentially persist and successfully asexually reproduce there. A new invader could even have greater asexual production than the local Au. aurita s.l. Establishment of the invaders could increase genetic variation of subsequent generations and increase their adaptability to environmental changes. Our results suggest that introduction of exotic Au. aurita s.l. populations could increase jellyfish blooms in the Mediterranean Sea.     

Circulation and heat budget of the northern Adriatic Sea (Italy) due to a Bora event in January 2001: a numerical model study

The Princeton Ocean Model was implemented to investigate the response of northern Adriatic Sea during the Bora event in January 2001 when strong wind and surface cooling was reported. The model has been run with realistic wind stress, surface heat flux and river runoffs forcings continuously from 1 January 1999 to 31 January 2001. The wind stress and surface heat flux was computed by the bulk parameterization, using the European Centre for Medium Range Weather Forecast analysis fields and the Comprehensive Ocean Atmosphere Data Set cloud data. All the freshwater sources along the Adriatic coastlines were represented by point or line source functions. Open boundary conditions in the Ionian Sea along a latitudinal boundary were nested within a large scale model of the Mediterranean Sea. The numerical study found that, before the Bora event of 13–17 January 2001, the water column of the northern Adriatic Sea was stratified by salinity, and the temperature was already cooler at the surface and over the shallower shelf region. The pre-Bora circulation of the northern Adriatic Sea was relatively weak and baroclinic with maximum surface currents occurred near the Italian coast. During the Bora event, the water column was well mixed in the most of coastal region of the northern Adriatic Sea. The atmospheric cooling produced colder water over the northern and western Adriatic Coast. The circulation of the northern Adriatic Sea was barotropic and dominantly wind driven, with maximum current speed of about 1 m s⁻¹. The numerical study also demonstrated that the Bora event decreased the heat content of the water column with an area averaged value of 205 W m⁻² over the shallow northern shelf. It was concluded that the heat budget of the northern Adriatic Sea during the Bora event was a balance between the surface heat loss, horizontal net heat inflow and resulting heat content decrease. The horizontal advection played a particularly important role in controlling the water temperature change over the shallower northern shelf.