Population biology of two helminth parasites of flatfishes from the Atlantic coast of Morocco

On the Atlantic Moroccan coast, two species of pleuronectiform fish coexist, wedge sole ( Citharus linguatula ) a temperate species, and spotted flounder ( Dicohgoglossa cuneata ) a tropical one. These two species are the definitive host for Bothriocephalus andresi and Acanthocephaloïdes propinquus , respectively. A negative binomial fits the dispersion of B. andresi in the population of C. linguatula . The fish become parasitized between their first and third year, while still immature, with the highest abundances in the third year. After the fish become ichthyopagous, the level of infection drops drastically. We postulate that copepods are the sole source of infection. The presence of B. andresi in both Atlantic and Mediterranean populations of C. linguatula shows that the parasite follows the host during its migration along the Mediterranean coasts.
The prevalence of A. propinquus remains above 50% as a consequence of the large numbers of amphipods included in the diet of D. cuneata , regardless of the age of the fish or the season. The amphipods, Phtisica marina and Pariambus typicus are an important food and represent potential intermediate hosts for A. propinquus . The population dynamics of A. propinquus on the Atlantic Moroccan coasts are similar to those observed in parasite populations of Gobius niger in the Gulf of Lion (French Mediterranean). The observation of B. andresi and A. propinquus in both Moroccan Atlantic and Mediterranean fishes highlights the problem of the evolution of these parasite populations during the colonization of the Mediterranean by the hosts.     

Genetic isolation between Atlantic and Mediterranean albacore populations inferred from mitochondrial and nuclear DNA markers

Genetic population structure of Atlantic and Mediterranean albacore Thunnus alalunga was investigated using nucleotide sequence variations of the glucose‐6‐phosphate dehydrogenase gene intron ( G6PD ) and the mitochondrial DNA (mtDNA) D‐loop region ( Dloop ). Restriction analysis using Ase I digestion detected two major restriction types ( A and B ) at the Dloop locus with strong frequency differences between Atlantic and Mediterranean samples. Thirty‐six individuals of 100 Mediterranean albacore were of the B type whereas no B type individuals were found in the Atlantic samples ( n  = 102). Phylogenetic analysis using nucleotide sequence data of the Dloop locus indicated that the B type lineage recently arose from the ancestral A lineage in the Mediterranean Sea and has not dispersed into the Atlantic Ocean. The frequencies of two alleles ( L and S ) at the G6PD locus were significantly different between the samples from the Atlantic ( L  = 0·495) and the Mediterranean ( L  = 0·725), but no significant heterogeneity was observed between mtDNA‐ A and ‐ B types of the Mediterranean sample. These molecular data indicate that gene flow between the Atlantic and Mediterranean albacore populations have been considerably restricted and strongly suggest these populations should continue to be treated as two distinct management units.     

Phylogeographical lineages in brown trout (Salmo trutta): investigating microgeographical differentiation between native populations from Northern Spain

SUMMARY 1. The large microgeographical differentiation revealed by allozyme studies in brown trout ( Salmo trutta) populations is one of the most striking features of this species. Additionally, allozymes showed great genetic differences between Atlantic and Mediterranean populations on a macrogeographical scale.
2. This study was carried out in order to assess whether the great differences observed between Atlantic and Mediterranean populations persisted where the two are geographically close (the 'microgeographical scale'). Sixteen populations of brown trout, S. trutta , were screened for genetic variation at 25 allozyme loci. The sampling sites, which occupied a relatively small geographical area, were distributed across Cantabrian (Atlantic) and Mediterranean drainages in Northern Spain.
3. The neighbour-joining tree, inferred from Nei's genetic distance, showed that brown trout populations clustered into two different groups. These groups corresponded to the Cantabrian and the Mediterranean groups of populations, although no clear geographical pattern emerged within each of the groups. This geographical pattern is basically caused by significant differences in the frequency distribution of the CK-A1 * locus, with a higher frequency of * 115 in Cantabrian samples (0.586 ± 0.091) while allele * 100 was more frequent in Mediterranean samples (0.931 ± 0.038). In addition, this study revealed alleles exclusive to the Mediterranean and Cantabrian populations, agreeing with previous findings.
4. Genetic differentiation between Cantabrian and Mediterranean regions (14.19%) was similar to that estimated in Spain at a larger scale (13%), showing that most of the differences between the regions can be observed even in a small geographical area.     

The influence of the parasite invasion on antioxidant enzyme activity in the liver and muscles of a host, the Black Sea flounder Psetta maxima maeotica

The influence of the intestine cestode Botriocephalus gregarius on the antioxidant enzyme system in the liver and muscle of its host, the Black Sea flounder Psetta maxima maeotica, was studied. The significant increase of superoxidedismutase activity in examined tissues and changes of catalase and peroxidase activities were detected. High correlation (r > or = 0.6) between enzyme activities and the parasite number was noted. The possibility to use antioxidant enzyme activity as biomarkers of the parasite invasion in fish is discussed.     

The poor health of deep-water species in the context of fishing activity and a warming climate: will populations of Molva species rebuild or collapse?

Many deep-water fish populations, being K-selected species, have little resilience to overexploitation and may be at serious risk of depletion as a consequence. Sea warming represents an additional threat. In this study, the condition, or health, of several populations of common ling (Molva molva), blue ling (Molva dypterygia) and Mediterranean or Spanish ling (Molva macrophthalma) inhabiting different areas in the North Atlantic and the Mediterranean was evaluated, to shed light on the challenges these deep-water species are facing in the context of fishing activity and a warming climate. The data on the condition of Molva populations which are analysed here have been complemented with data on abundance and, for the southernmost species (Mediterranean ling), with two other health indicators (parasitism and hepato-somatic index). Despite some exceptions (e.g., common ling in Icelandic waters), this study shows that the condition of many populations of Molva species in the northeastern Atlantic and the Mediterranean Sea has worsened, a trend which, in recent decades, has usually been found to be accompanied by a decline in their abundance. In addition, the poor health status of most populations of common ling, blue ling and Mediterranean ling considered in this analysis points to a lower sustainability of these populations in the future. Overall, the health status and abundance of Molva populations in the northeastern Atlantic and the Mediterranean suggest that only some populations located in the North Atlantic may be able to rebuild, whereas the populations in southern North Atlantic and the Mediterranean, which are probably most at risk from sea warming, are facing serious difficulties in doing so. In the context of fisheries and global warming, this study's results strongly indicate that management bodies need to consider the health status of many of the populations of Molva species, particularly in southern European waters, before implementing their decisions.     

Comparative ionic flux and gill mucous cell histochemistry: effects of salinity and disease status in Atlantic salmon (Salmo salar L.)

Two experiments were conducted to assess the physiological effects of freshwater exposure and amoebic gill disease (AGD) in marine Atlantic salmon (Salmo salar L.). The first experiment monitored marine salmon during a 3 h freshwater exposure, the standard treatment for AGD in Tasmania. The second experiment described the gill mucous cell histochemistry for freshwater adapted and seawater acclimated fish (AGD affected and unaffected) for possible correlations to ionoregulation. When exposed to freshwater, marine Atlantic salmon experienced a minor ionoregulatory dysfunction represented by a net efflux of Cl(-) ions at 3 h. AGD affected fish experienced the net efflux of Cl(-) ions 1 h sooner, and had a significantly greater net efflux of total ammonia. Changes to gill mucous cell populations corresponded to differing salinity and the presence of AGD. In AGD affected fish, these populations significantly differed between lesion and non-lesion associated areas of the gill filament. Our results have shown changes in the ionoregulatory capacity of Atlantic salmon due to freshwater exposure and AGD. Gill mucous cell histochemistry indicates the potential importance of the mucous layer in ionoregulation and disease. In comparison to previous studies on rainbow trout, these results suggest that Atlantic salmon have a greater short-term ionoregulatory capacity.     

Living together but remaining apart: Atlantic and Mediterranean loggerhead sea turtles (Caretta caretta) in shared feeding grounds

Juvenile loggerhead sea turtles (Caretta caretta) from Atlantic nesting populations migrate into the western Mediterranean, where they share feeding grounds with turtles originating in the Mediterranean. In this scenario, male-mediated gene flow may lead to the homogenization of these distant populations. To test this hypothesis, we genotyped 7 microsatellites from 56 Atlantic individuals sampled from feeding grounds in the western Mediterranean and then compared the observed allele frequencies with published data of 112 individuals from Mediterranean nesting beaches. Mediterranean populations were found to be genetically differentiated from the Atlantic stock reaching the western Mediterranean (F(st) = 0.029, P < 0.001); therefore, the possible mating events between Atlantic and Mediterranean individuals are not sufficient to homogenize these 2 areas. The differentiation observed between these 2 areas demonstrates that microsatellites are sufficiently powerful for mixed stock analysis and that individual assignment (IA) tests can be performed in combination with mitochondrial DNA (mtDNA) analysis. In a set of 197 individuals sampled in western Mediterranean feeding grounds, 87% were robustly assigned to Atlantic or Mediterranean groups with the combined marker, as compared with only 52% with mtDNA alone. These findings provide a new approach for tracking the movements of these oceanic migrants and have strong implications for the conservation of the species.     

Genetic divergence in the Atlantic-Mediterranean Montagu's blenny, Coryphoblennius galerita (Linnaeus 1758) revealed by molecular and morphological characters

Coryphoblennius galerita is a small intertidal fish with a wide distribution and limited dispersal ability, occurring in the northeastern Atlantic and Mediterranean. In this study, we examined Atlantic and Mediterranean populations of C. galerita to assess levels of genetic divergence across populations and to elucidate historical and contemporary factors underlying the distribution of the genetic variability. We analyse three mitochondrial and one nuclear marker and 18 morphological measurements. The combined dataset clearly supports the existence of two groups of C. galerita: one in the Mediterranean and another in the northeastern Atlantic. The latter group is subdivided in two subgroups: Azores and the remaining northeastern Atlantic locations. Divergence between the Atlantic and the Mediterranean can be the result of historical isolation between the populations of the two basins during the Pleistocene glaciations. Present-day barriers such as the Gibraltar Strait or the 'Almeria-Oran jet' are also suggested as responsible for this isolation. Our results show no signs of local extinctions during the Pleistocene glaciations, namely at the Azores, and contrast with the biogeographical pattern that has been observed for Atlantic-Mediterranean warm-water species, in which two groups of populations exist, one including the Mediterranean and the Atlantic coast of western Europe, and another encompassing the western tropical coast of Africa and the Atlantic islands of the Azores, Madeira and Canaries. Species like C. galerita that tolerate cooler waters, may have persisted during the Pleistocene glaciations in moderately affected locations, thus being able to accumulate genetic differences in the more isolated locations such as the Azores and the Mediterranean. This study is one of the first to combine morphological and molecular markers (mitochondrial and nuclear) with variable rates of molecular evolution to the study of the relationships of the Atlantic and Mediterranean populations of a cool-water species.     

Distribution of Anisakis larvae, identified by genetic markers, and their use for stock characterization of demersal and pelagic fish from European waters: an update

In the present paper, recent results obtained on the use of different distributions observed in larval species of Anisakis, genetically identified by means of allozyme markers, for stock characterization of demersal (Merluccius merluccius), small (Trachurus trachurus) and large pelagic (Xiphias gladius) finfish species in European waters, are reviewed and discussed. Several species of Anisakis were identified in the three fish hosts: A. simplex (s.s.), A. physeteris, A. typica, A. ziphidarum, A. pegreffii, A. brevispiculata and A. paggiae. Canonical discriminant analysis performed on all the samples of the three fish species collected in areas comprising their geographical range, according to the different species of Anisakis identified, showed distinct fish populations in European waters. In all the three fish hosts, the pattern of distribution of Anisakis larvae allowed discrimination of Mediterranean stocks from Atlantic stocks. In the case of swordfish, the possible existence of a southern Atlantic stock separated from a northern one is also suggested. Congruence and discordance with the population genetic data inferred from allozyme markers on the same samples of the three fish species are also discussed.     

Electrophoretic studies on the Anisakis simplex complex (Ascaridida:Anisakidae) from the Mediterranean and North-East Atlantic

Full-size image

and 1986. Electrophoretic studies on the Anisakis simplex complex (Ascaridida: Anisakidae) from the Mediterranean and North-East Atlantic. International Journal for Parasitology 16: 633–640. The genetic variation of the sibling species Anisakis simplex A and A. simplex B was investigated by electrophoretic analysis of 22 gene-enzyme systems. The two species are reproductively isolated and no gene flow takes place between them. Three loci, Sod, Adk-2 and Lap-1, show distinct alleles in A. simplex A and A. simplex B, allowing their reliable identification both at the larval and adult stages. A fourth locus, Got, appears to be diagnostic at the 95% level. The value of Nei's genetic distance found between A. simplex A and A. simplex B is 0.28. Parameters of genetic variability (He, P, A) are given for both species. The geographic distribution of A. simplex A and A. simplex B appears to be mainly Mediterranean for the former, and mainly North Atlantic for the latter. Several paratenic hosts (fish and squid) and one cetacean definitive host are identified for each of the two species. The names A. pegreffii and A. simplex are tentatively proposed for A. simplex A and A. simplex B respectively.     

Common dolphinfish,Coryphaena hippurus,spawn at their easternmost Atlantic range adjacent to the Mediterranean Sea

This study provides sound evidence of the spawning activity of common dolphinfish (Coryphaena hippurus) in the Gulf of Cadiz (southwest Spain) during summer months. Along with waters off Côte d'Ivoire, this area adjacent to the Mediterranean Sea is the easternmost Atlantic spawning ground documented for the species. All the fish analysed (5 males and 15 females) were at spawning stage, except the smallest female (61.5 cm in fork length), which might not have yet reached sexual maturation. The oceanographic conditions during the samplings were similar to those occurring during the peak spawning season in the Gulf of Mexico. The study results are consistent with recent genetic studies indicating differentiation between Mediterranean and Atlantic dolphinfish populations, with the Strait of Gibraltar representing a boundary between two distinct reproductive units. The present results may contribute to improve the understanding of the dynamics of dolphinfish population and the management of eastern Atlantic and Mediterranean stocks.     

Genetic identity of YOY bluefin tuna from the eastern and western Atlantic spawning areas

We used 320 young-of-the-year (YOY) specimens of the highly migratory and overfished Atlantic bluefin tuna, Thunnus thynnus, Linnaeus 1758, to evaluate the hypothesis that Atlantic bluefin tuna comprises 2 stocks with spawning grounds in the Gulf of Mexico and in the Mediterranean Sea. Significant genetic differentiation at 8 nuclear microsatellite loci (F(ST) = 0.0059, P = 0.0005) and at the mitochondrial control region (Phi(ST) = 0.0129, P = 0.0139) was detected among YOY Atlantic bluefin tuna captured on spawning grounds in the Gulf of Mexico (n = 40) versus the western (n = 255) and eastern (n = 25) basins of the Mediterranean Sea. The genetic divergence among spawning populations, combined with the extensive trans-Atlantic movements reported for juvenile and adult Atlantic bluefin tuna, indicates a high degree of spawning site fidelity. Recognition of genetically distinct populations necessitates independent management of Atlantic bluefin tuna on spawning grounds and warrants evaluation of the level of mixing of populations on feeding grounds. The genetic pattern might not have been detected unless juvenile specimens or actively spawning adults had been sampled.     

Genetic evidence for three species within Pseudoterranova decipiens (Nematoda, Ascaridida, Ascaridoidea) in the North Atlantic and Norwegian and Barents Seas

Genetic variation of 1017 specimens of codworm, Pseudoterranova decipiens, collected from fish and seals at 23 sampling locations in the North Atlantic and Norwegian and Barents Seas, was analysed on the basis of 16 enzyme loci. Three reproductively isolated species, provisionally designated P. decipiens A, B and C, were detected, showing distinct alleles at the following loci: Mdh-1, 6Pgdh, Np, Pgm, Est-2 (between species A and B); Mdh-3, 6Pgdh, Np, Sod-1, Adk, Pgm, Est-2, Mpi (between A and C); Mdh-1, Mdh-3, Sod-1, Adk, Pgm, Est-2, Mpi (between B and C). One F1 hybrid was observed between P. decipiens A and B, but this apparently does not lead to any gene exchange between the two species, which do not show any evidence of introgression. No hybrids or introgressed individuals were observed between P. decipiens C and either A or B. Genetic distances among conspecific populations were low (average Nei's D 0.001-0.005), even though they were collected thousands of kilometres apart, indicating high levels of gene flow within each of the three species. The values of Nei's index D were 0.44 between P. decipiens A and B, 0.57 between B and C, and 0.79 between A and C. Estimated evolutionary divergence times, using Nei's formula, range from 2 to 4 million years. Differences between P. decipiens A, B and C were also found with respect to genetic variability, morphology, geographical distribution and hosts. Mean heterozygosity values of 0.08, 0.05 and 0.02 were obtained for P. decipiens A, B and C, respectively. Preliminary morphological examination of adult males, previously identified by multilocus electrophoresis, revealed differences in the relative size and pattern of caudal papillae. P. decipiens B is widespread in the study area, whereas P. decipiens A was found only in the North-East Atlantic and Norwegian Sea. In this area P. decipiens A is most common in the grey seal, Halichoerus grypus, while the common seal, Phoca vitulina, is the main host for P. decipiens B. In Canadian Atlantic waters, where P. decipiens A is apparently absent, P. decipiens B infects both grey and common seals; a few specimens were also found in the hooded seal, Cystophora cristata. The only definitive host so far identified for P. decipiens C is the bearded seal, Erignathus barbatus; P. decipiens C appears to be widespread, occurring in both the North-West Atlantic and Barents Sea.     

Historical population dynamics and demography of the Eastern Atlantic pomacentrid Chromis limbata (Valenciennes, 1833)

Recent studies have focused on the relationship between the marine fauna of the Eastern Atlantic and the Mediterranean Sea, but within the Atlantic, little is known about genetic relationships between populations of the Macaronesian islands. In this study, we tested whether the paleo-climatology and paleo-oceanography of the region could predict the genetic relationships among three Eastern Atlantic populations (Azores, Madeira, and Canaries) of a damselfish, Chromis limbata, and compared our results with its Mediterranean and adjacent Atlantic sister species, Chromis chromis. We combined phylogeographic and coalescent approaches using the fast evolving mitochondrial control region gene. No population structure was found for the three archipelagos. The coalescence time estimated for C. limbata (0.857-1.17 Mya) was much greater than that estimated for C. chromis. We propose that this difference reflects differences in glaciating extents in the Northeastern Atlantic and the Mediterranean. Diversity indexes (Hd and genetic distances) together with historical demographic parameters of C. limbata (Theta and g) revealed a more stable population history when compared to C. chromis. Our results suggest that the Macaronesian populations of C. limbata have probably been less affected by the last glaciation than the Mediterranean populations of C. chromis. Migration across the three archipelagos was estimated and a prevailing northwest trend was detected. This result supports the idea of a colonization of the Azores by warm water fish from Madeira or the westernmost Canary islands which acted as major glacial refugia for the tropical and subtropical marine fauna during the glaciations.     

Population genetic structure of North Atlantic, Mediterranean Sea and Sea of Cortez fin whales, Balaenoptera physalus (Linnaeus 1758): analysis of mitochondrial and nuclear loci

Samples were collected from 407 fin whales, Balaenoptera physalus , at four North Atlantic and one Mediterranean Sea summer feeding area as well as the Sea of Cortez in the Pacific Ocean. For each sample, the sex, the sequence of the first 288 nucleotides of the mitochondrial (mt) control region and the genotype at six microsatellite loci were determined. A significant degree of divergence was detected at all nuclear and mt loci between North Atlantic/Mediterranean Sea and the Sea of Cortez. However, the divergence time estimated from the mt sequences was substantially lower than the time elapsed since the rise of the Panama Isthmus, suggesting occasional gene flow between the North Pacific and North Atlantic ocean after the separation of the two oceans. Within the North Atlantic and Mediterranean Sea, significant levels of heterogeneity were observed in the mtDNA between the Mediterranean Sea, the eastern (Spain) and the western (the Gulf of Maine and the Gulf of St Lawrence) North Atlantic. Samples collected off West Greenland and Iceland could not be unequivocally assigned to either of the two areas. The homogeneity tests performed using the nuclear data revealed significant levels of divergence only between the Mediterranean Sea and the Gulf of St Lawrence or West Greenland. In conclusion, our results suggest the existence of several recently diverged populations in the North Atlantic and Mediterranean Sea, possibly with some limited gene flow between adjacent populations, a population structure which is consistent with earlier population models proposed by Kellogg, Ingebrigtsen, and Sergeant.     

Maintenance of genetic differentiation across a transition zone in the sea: discordance between nuclear and cytoplasmic markers

To investigate the origin and maintenance of the genetic discontinuity between Atlantic and Mediterranean populations of the common sea bass (Dicentrarchus labrax) we analysed the genetic variation at a fragment of mitochondrial cytochrome b sequence for 18 population samples. The result were also compared with new or previously published microsatellite data. Seven mitochondrial haplotypes and an average nucleotidic divergence of 0.02 between Atlantic and Mediterranean populations that matches a Pleistocene allopatric isolation were found. The frequency variation at the cytochrome b locus was many times greater between Atlantic and Mediterranean populations (theta(C) = 0.67) than at microsatellite loci (theta(N)= 0.02). The examination of the different evolutionary forces at play suggests that a sex-biased hybrid breakdown is a likely explanation for part of the observed discrepancy between mitochondrial and nuclear loci. In addition, an analysis is made of the correlation between microsatellite loci points towards the possible existence of a hybrid zone in samples from the Alboran Sea.     

Bothriocephalus scorpii (Müller) (Cestoda: Pseudophyllidea) in turbot Scophthalmus maximus (L.) and brill S. rhombus (L.) from the southern North Sea

Turbot (11–70 cm long) and brill (21–70 cm long) were collected in the summer of 1968 north of the Brown Bank, Texel Hole and were infected by Bothriocephalus scorpii. Infections occurred even in the first year of life of the fish. In turbot, with increasing length, the average weight of tapeworm rose from 0–6 to 14-9 g/flsh. Brill were less commonly infected and then only slightly. The intermediate host of the tapeworm seems to be Gobius species, upon which juvenile turbot and brill prey most.     

Relationships of wild Brassica species with chromosome number 2n = 18, based on RFLP studies.

An array of 10 wild Brassica species with chromosome number 2n = 18 represented by 34 populations was analyzed for genome similarity using genomic and cDNA clones. Species studied included B. bourgeaui (Webb) O. Kuntze, B. cretica Lam., B. hilarionis G.E. Post, B. incana Ten., B. insularis Moris, B. macrocarpa (Guss.) Caruel, B. montana Pourret, B. oleracea L., B. rupestris Rafin., and B. villosa Biv. The RFLP data were used to calculate similarities between populations that were subsequently treated in a cluster analysis. Most populations of a species were grouped together and were separate from populations of other species. The previously identified B. incana - B. rupestris - B. villosa complex was verified, and genetic similarity between the species B. montana and B. oleracea was evident. An interesting association between B. insularis and B. macrocarpa was observed. The UPGMA analysis showed that the species tended to cluster according to geographic region: B. cretica and B. hilarionis comprise a cluster that could be called Eastern Mediterranean; B. oleracea, B. bourgeaui, and B. montana define an Atlantic - Western Mediterranean cluster; B. incana, B. rupestris, and B. villosa form an Italian group; and a B. insularis - B. macrocarpa association may be called Central Mediterranean.     

The occurrence of two opecoeliid digeneans in Mullus barbatus and M. surmuletus from the Spanish south-eastern Mediterranean

The infection by Opecoeloides furcatus and Poracanthium furcatum (Opecoeliidae) was studied in 121 Mullus barbatus and 113 M. surmuletus collected from the Spanish south-eastern Mediterranean. The prevalence of infection was most frequent in M. surmuletus with values of 81.42% for O. furcatus and 38.05% for P. furcatum. In M. barbatus the prevalences of O. furcatus and P. furcatum were 54.54% and 14.88% respectively. Statistically significant differences were found between the infection of the two hosts with P. furcatum. No significant differences in worm burdens could be attributable to host size or to seasonal changes, although a lower infection of M. barbatus by O. furcatus occurred in the autumn. Furthermore, the electrophoretic mobility of the enzyme malic dehydrogenase (MDH) was also studied and both digeneans presented different patterns, corresponding in both cases to homozygotic genotypes.     

Genetic differentiation among populations of the salt marsh beetle Pogonus littoralis (Coleoptera: Carabidae): a comparison between Atlantic and Mediterranean populations

Genetic differentiation was studied among populations of the salt marsh beetle Pogonus littoralis (Coleoptera: Carabidae), comparing Atlantic and Mediterranean populations over a large part of its range. The genetic structure of this highly mobile beetle was investigated by studying allozyme polymorphism at nine enzyme loci in 13 populations. Mediterranean P. littoralis were highly significantly differentiated from Atlantic populations. Moreover, more isolated Atlantic populations showed increased differentiation and decreased genetic diversity compared to less fragmented Mediterranean populations.