Mitochondrial phylogeography of the killifish Aphanius fasciatus (Teleostei, Cyprinodontidae) reveals highly divergent Mediterranean populations

We studied the genetic structure of the Mediterranean killifish Aphanius fasciatus. Analysis of the sequence variation in a 372-bp portion of the mitochondrial control region in 623 fish from 27 sampling sites along the species’ distributional range (Tyrrhenian coast, Sardinia, Sicily, Adriatic coast, Malta, Tunisia, and Greece) yielded 120 distinct haplotypes. Most of the haplotypes are unique, and only 15 % are shared among different populations. The high F _ST value (=0.80) suggests a strong population genetic structuring. The phylogenetic analysis based on Bayesian inference, maximum likelihood and maximum parsimony, and the median-joining network show a sharp separation of the Southeastern Sicilian populations (belonging to the Hyblean region) and of the fluvial Tunisian population of Rio Melah from the others. The Adriatic, the Eastern Sicilian, and the Greek populations are well differentiated, while the group of populations from the Central-Western Mediterranean does not show a clear pattern of differentiation. Our findings indicate that the current genetic structuring of A. fasciatus reflects historical geographical patterns occurring within the Mediterranean basin from the Late Miocene to the Pleistocene. The presence of divergent evolutionary entities in the Hyblean region and the Tunisian Rio Melah supports their inclusion as target areas for the conservation of A. fasciatus.     

Phylogeography and genetic structure of the Mediterranean killifish <Emphasis Type="Italic">Aphanius fasciatus</Emphasis> (Cyprinodontidae)

The purpose of this study was to determine the phylogeographic structure of the brackish-hypersaline cyprinodont fish Aphanius fasciatus (Valenciennes, 1821), using sequencing and RFLP analysis of a 1,330 bp mitochondrial DNA segment containing part of the 16S rRNA gene as well as the genes for tRNA-Leu, NADH subunit 1 and tRNA-Ile. Individuals were collected from 13 different sites in Greece and Turkey, while seven published A. fasciatus sequences were also included to cover the area of distribution of the species. Pairwise sequence divergence values ranged from 0 to 4.51%. Congruent phylogenies were recovered with maximum likelihood, maximum parsimony and neighbour-joining methods. All analyses revealed two main groups. The first group consists of populations from almost all localities that drain into the Aegean Sea. The second group comprises the remaining population samples, which in some cases seem to consist of population-specific subgroups. Our results show that vicariant events have predominantly affected the evolution of A. fasciatus, with the Messinian salinity crisis having shaped the present genetic structure of its populations. Additionally, the life-history traits of the species, which determine a low potential for dispersal, coupled with the typical fragmentation of brackish-hypersaline water habitats have led to a high degree of isolation of A. fasciatus populations, even at restricted spatial scales. Analysis of the partitioning of the total amount of polymorphism with analyses of molecular variance (AMOVA) gave a value of F _ST = 84.6%. Potential conservation policies concerning A. fasciatus should also consider the low-genetic variability in the majority of its populations and the presence of fixed haplotypes in some of them.     

An osteological study of the dugong Dugong dugon (Sirenia) from India

An osteological study of the dugong (sea cow) Dugong dugon (Müller) from India, based on complete skeletons of an adult male and a juvenile female indicated certain morphological variations with age. It has also revealed a close osteological similarity between the dugongs from India and the Red Sea. Based on a statistical analysis, regression equations for certain typical skull measurements have been calculated; these equations, characteristic of the dugong from India, could be used for comparing dugongs of different regions.     

Biology of Serranicotyle labracis,ectoparasite of Dicentrarchus labrax (Teleostei): contribution to the study of its populations

Serranicotyle labracis (Van Beneden and Hesse, 1863) Maillard, Euzet and Silan, 1988 (synonym: Microcotyle labracis Van Beneden and Hesse, 1863) is a monogenean Polyopisthocotylea and a gill parasite of the bass Dicentrarchus labrax (Linné, 1758). Analysis of a western Mediterranean population of this parasite (Gulf of Lions, Languedoc, France) clarified several aspects of its biology. Its location in the gill biotope (Silan et al. 1987) was not fundamentally different from that found by Oliver (1977) and Winch (1983), but its distribution in host populations differed from that reported in the earlier studies; it displayed a negative binomial distribution in the study zone. At a finer level of observation, the behaviour of the hosts, as a function of degree of maturity and sex, appeared to determine more than any other factor the extent of infestation by the parasite. Previous suggestions in the literature to account for these differences in population dynamics are discussed.     

Subitaneous and diapause eggs in Mediterranean populations ofPontella mediterranea (Copepoda: Calanoida): A morphological study

Transmission electron microscope studies on the morphology of subitaneous and diapause eggs in the neustonic copepodPontella mediterranea, collected in coastal waters of the Gulf of Naples from May to December 1986, revealed striking structural differences between the two egg types and the existence of a third egg type that was morphologically similar to a diapause egg but which hatched within 2 to 3 d. In smooth, subitaneous eggs the plasma membrane was surrounded by a thin outer chorion as compared to the thick and highly complex four-layer structure enveloping subitaneous eggs with short spines, and diapause eggs. This coat may be assembled just prior to or after deposition since the morphology of developing oocytes was similar in females laying either subitaneous or diapause eggs. The only other major structural difference between the two egg types was the presence of numerous, disc-like bodies in the ooplasm of diapause eggs. Development was arrested for at least 3 mo, prior to the initiation of the first cleavage, suggesting that these eggs were kept dormant by some endogenous and, possibly, structural mechanism preceeding the termination of diapause.Dedicated to B. Scotto di Carlo, teacher and friend, who died tragically at sea during the course of a cruise on December 15, 1988     

Social organization and sexual pattern in the Mediterranean parrotfish Sparisoma cretense (Teleostei: Scaridae)

We examined the social organization and reproductive pattern of a population of Sparisoma cretense L. at Lampedusa Island (Italy). During the breeding season (July to September) individuals occur either in territorial or in non-territorial groups, which quantitatively differ with respect to use of space, sex-ratio and sexual activity. Territorial groups consist of one male plus one to three females, whereas in non-territorial groups as many as 54 fish share a common area. Among non-territorial groups, variations, in both sex-ratio and behavior, suggest they could represent either feeding aggregations or reproductive aggregations. Spawning, always in pairs, occurs daily in a short period of time before dusk and has been observed only in territorial groups. Group spawning has never been observed, but another alternative mating tactic, streaking on pair spawning, was recorded. In the non-breeding season fish do not aggregate in organized social units. Histological examination of gonads showed that adults are larger than 12 cm total length. Females have an asynchronous ovary, typical of species spawning several times during the breeding season. Males show secondary testes, indicating that their gonads develop ovaries as juveniles, which are later redifferentiated into functional testes. Histological and demographic data seem to indicate that, as in other species of this genus, prematurational sex-change occurs. The sexual pattern appears to be essentially gonochoristic, but the potential for sex-change is not excluded. Received: 5 January 1999 / Accepted: 22 June 1999     

Structural, morphological and genetic variability in Halophila stipulacea (Hydrocharitaceae) populations in the western Mediterranean

Halophila stipulacea (Forssk.) Ascher. is a dioecious seagrass that colonized the Mediterranean basin probably following the opening of the Suez Canal (1869). Natural meadows have been reported since the end of the last century on the eastern side of the basin and only recently along the northern coast of Sicily. In the present study we examined the morphological and genetic variability of two natural meadows located along the Sicilian coast (Vulcano Island and Oliveri-Tindari coastal lakes). In order to determine morphological and genetic polymorphism, samples were collected at different depths (5, 15 and 25 m depth at the Vulcano site) and positions (edge vs mid-bed) within the meadows. Statistically significant differences in phenotypic features were found between the factors “depth” and “position” within the same meadow and between the two localities. Genetic diversity was assessed using randomly amplified polymorphic DNA and found to be high. Deep and shallow stands of the Vulcano Island meadow clustered in different positions in the UPGMA tree. The shallow Vulcano stand was found to be closer to the shallow Oliveri-Tindari meadow than to the deeper stand from Vulcano. Mantel's test did not allow rejection of the null hypothesis of independence of morphological and molecular distance matrices. We conclude that (i) H. stipulacea shows high morphological and genetic polymorphism, (ii) environmental and/or ecological barriers exist between different depths, and (iii) trends of morphological and genetic variability may be influenced by different environmental and/or ecological factors. Received: 15 November 1998 / Accepted: 21 May 1999     

Recent strandings of the giant jellyfish Rhizostoma luteum Quoy and Gaimard, 1827 (Cnidaria: Scyphozoa: Rhizostomeae) on the Atlantic and Mediterranean coasts

We present reports of sightings of living and stranded specimens of Rhizostoma luteum on the Atlantic coast of Morocco and along the south shore of the Iberian Peninsula in June–July 2012 and in January–February 2013. During summer 2012 and following the dominant currents, the jellyfish first appeared in the Gulf of Cadiz west of the Strait of Gibraltar. Subsequently, seven additional sightings were reported east of the Strait, in the Alboran Sea. In winter 2013, another event of stranded individuals of this species occurred in the Gulf of Cadiz. A phylogenetic analysis performed on the mitochondrial cytochrome c oxidase I (COI) gene sequence in specimens from both stranding events confirmed the morphological classification, ratifying that R. luteum differs from Rhizostoma octopus and Rhizostoma pulmo. This study records the presence of this species for the first time in 60 years.     

The role of sexual harassment in cave and surface dwelling populations of the Atlantic molly, Poecilia mexicana (Poeciliidae, Teleostei)

Sexual coercion is a common behaviour for males trying to compensate for being inferior in male competition and/or female choice. We measured the cost of male sexual harassment for females as reduced feeding time in three populations of the Atlantic molly (Poecilia mexicana). Of these populations, one originated from a typical river habitat, another from a lightless cave chamber and the third one from a small, milky creek outside the cave. We gave hungry females an opportunity to feed in the presence of either another female or a male. We found the following: (1) male sexual harassment represents a cost in the river dwelling population and both small and large males significantly reduce female feeding time; and (2) sexual harassment was not detected in the other two populations (cave and cave entrance). In the cave molly, small males are at a disadvantage in female choice, but predation by an aquatic heteropteran selects against large male body size.     

Genetic protein variation in red mullet (Mullus barbatus) and striped red mullet (M. surmuletus) populations from the Mediterranean Sea

Starch-gel electrophoresis of allozymes was used to differentiate the two red mullet species (Mullus barbatus L. and M. surmuletus L.) in the Mediterranean Sea and, further, to investigate the genetic stock structure of M. barbatus in the eastern Mediterranean area. Twenty putative enzyme-coding loci were examined in eight M. barbatus samples caught in the Aegean and Ionian Seas (Greece) and in the Gulf of Lion (France), and two M. surmuletus samples caught in the Aegean and Gulf of Lion. A high degree of genetic polymorphism was found in both species. Species-specific electrophoretic patterns were found in PGI* and PGM*. Estimates of variance of allele frequencies among samples (F _ST) and ² analyses both revealed significant differences (P < 0.05) among the M. barbatus samples. Most of the genetic variation was among samples regardless of region. The mean value of Nei's genetic distance between the two species was 0.329. Genetic distance among M. barbatus samples was low (maximum Nei's D = 0.012), with the sample from Platania differing most from other M. barbatus samples. This is probably be due to founder effects existing at this area. These results suggest that allozyme analysis may provide important information on the genetic structure of the red mullet to ensure sustainable management of this species. Received: 7 May 1997 / Accepted: 13 October 1997     

Personality trait differences between mainland and island populations in the common frog (Rana temporaria)

Understanding and predicting species range expansions is an important challenge in modern ecology because of rapidly changing environments. Recent studies have revealed that consistent within-species variation in behavior (i.e., animal personality) can be imperative for dispersal success, a key process in range expansion. Here we investigate how habitat isolation can mediate differentiation of personality traits between recently founded island populations and the main population. We performed laboratory studies of boldness and exploration across life stages (tadpoles and froglets) using four isolated island populations and four mainland populations of the common frog (Rana temporaria). Both tadpoles and froglets from isolated populations were bolder and more exploratory than conspecifics from the mainland. Although the pattern can be influenced by possible differences in predation pressure, we suggest that this behavioral differentiation might be the result of a disperser-dependent founder effect brought on by an isolation-driven environmental filtering of animal personalities. These findings can have important implications for both species persistence in the face of climate change (i.e., range expansions) and ecological invasions as well as for explaining rapid speciation in isolated patches.     

Genetic structure of populations of two species of Chthamalus (Crustacea: Cirripedia) in the north-east Atlantic and Mediterranean

Protein electrophoresis on starch gels was used to investigate population genetic structure of the barnacles Chthamalus montagui Southward and C. stellatus (Poli) over their north-east Atlantic and Mediterranean ranges. In each species, a single locus exhibited marked differentiation of allele frequencies between Atlantic and Mediterranean localities; in C. stellatus, genetic differentiation between the two basins had not previously been noted. In both species, mean heterozygosity per locus appeared higher in the Mediterranean samples than in the Atlantic, and Mediterranean populations had more alleles at the loci studied. Possible explanations for the differentiation between the Atlantic and the Mediterranean populations are discussed. Received: 30 May 1996 / Accepted: 17 September 1996     

Genetic relationships between marine and marginal-marine populations of Cerithium species from the Mediterranean Sea

Results of isozyme electrophoresis were used to explore the genetic relationships between several Mediterranean morphs of Cerithium (Gastropoda: Prosobranchia), for which taxonomy is currently uncertain because of high intraspecific variability and low interspecific differentiation. The large species, classically known as C. vulgatum Bruguière, 1789 was identified at four sites (two in the French Mediterranean and two in southern Spain). Two different larval types were found in the French sites, but poecilogony could not be demonstrated. Individuals collected from harbours were not genetically distinct from open-sea populations of classic C. vulgatum. However, a population in the Embiez lagoon (French Mediterranean) which morphologically resembles C. vulgatum did display distinct genetic traits, supporting its status as a separate species. Of the small Cerithium species usually known as C. rupestre, two sympatric species (C.“rupestre” Risso, 1826 and C. lividulum Risso, 1826) were distinguished. Genotype frequencies within the analysed populations revealed much heterozygote deficiency. F _ST values (fixation index measuring the effects of population subdivision) suggest a higher genetic differentiation for C. lividulum populations than for C. vulgatum populations. We assume that a high larval dispersal capability (via planktotrophy) allows a high gene flow between populations of C. vulgatum. Received: 24 November 1998 / Accepted: 24 September 1999     

Allozyme and mitochondrial DNA variation in orange roughy,Hoplostethus atlanticus (Teleostei: Trachichthyidae): little differentiation between Australian and North Atlantic populations

Allozyme and mitochondrial DNA (mtDNA) genetic variation was compared in orange roughy (Hoplostethus atlanticus Collett) collected from waters off southern Australia and from waters about 22 000 km away in the North Atlantic west of Scotland. Samples were screened for 11 polymorphic allozyme loci and with 9 restriction enzymes. Significant heterogeneity between the two areas was detected for three allozyme loci (ADA ^*, CK ^* and GPI-1 ^*), and the overall G _ST (gene-diversity statistic) value of 1% was small but significant. Significant mtDNA haplotype heterogeneity was observed after ²- of haplotype frequencies but not after a G _ST analysis. Nucleotide sequence-diversity analysis showed very low net divergence (0.0023%) between the two samples. The Australian orange roughy had a lower allozyme heterozygosity and a lower mitochondrial DNA nucleon diversity than the North Atlantic sample. The very limited, although significant, allozyme and mitochondrial DNA heterogeneity between these areas suggests that there is some gene flow between these two populations. The species appears to be widespread, with its presence reported from the southern Pacific, southern Indian, and northern and southern Atlantic Oceans, and it is likely that gene flow between the antipodes is mediated by stepping-stone exchange between adjacent populations rather than by direct migration.     

Demography of central and marginal populations of monkeyflowers (Mimulus cardinalis and M. lewisii)

Every species occupies a limited geographic area, but how spatiotemporal environmental variation affects individual and population fitness to create range limits is not well understood. Because range boundaries arise where, on average, populations are more likely to go extinct than to persist, range limits are an inherently population-level problem for which a demographic framework is useful. In this study, I compare demographic parameters and population dynamics between central and marginal populations of monkeyflowers, Mimulus cardinalis and M. lewisii, along an elevation gradient spanning both species' ranges. Central and marginal populations of both species differed in survival and fecundity. For M. lewisii, these components of fitness were higher in central than in marginal populations, but for M. cardinalis the converse was true. To assess spatiotemporal variation in population dynamics, I used transition matrix models to estimate asymptotic population growth rates (lambda) and found that population growth rates of M. lewisii were highest at the range center and reduced at the range margin. Population growth rates of M. cardinalis were highest at the range margin and greatly reduced at the range center. Life table response analysis decomposed spatiotemporal variation in lambda into contributions from each transition between life stages, finding that transitions from large nonreproductive and reproductive plants to the seed class and stasis in the reproductive class made the largest contributions to spatial differences in lambda. These transitions had only low to moderate sensitivities, indicating that differences in projected population growth rates resulted mainly from observed differences in transition matrix parameters and their underlying vital rates.     

Phylogenetic relationships of coral-associated gobies (Teleostei,Gobiidae) from the Red Sea based on mitochondrial DNA data

Bryaninops, Gobiodon, Paragobiodon and Pleurosicya are the most abundant genera of coral-associated gobies. These genera are adapted to live among coral, while other small reef gobies (e.g., the genus Eviota) show no obligate association with this living substrate. Thirteen coral-associated species and two Eviota species were sampled from different regions of the Red Sea, along with four populations/species of Gobiodon from the Indian and western Pacific Oceans. A molecular phylogenetic analysis was performed using partial sequences of 12S rRNA, 16S rRNA and cytochrome b mitochondrial genes, 1,199 base pairs in total. Several clades were consistently resolved in neighbor joining-, maximum parsimony-, maximum likelihood and Bayesian analyses. While each of the four genera Gobiodon, Paragobiodon, Bryaninops and Pleurosicya proved to be monophyletic, their relative position in the phylogeny did not support an emergence of coral-associated gobiids as a monophyletic assemblage. Instead, two separate monophyletic sub-groups were discovered, the first comprising Gobiodon and Paragobiodon, and the second Bryaninops and Pleurosicya. Our molecular phylogenetic examinations also revealed one unassigned species of Gobiodon from the Maldives as a distinct species and confirmed three putative and yet unassigned species from the Red Sea. Moreover, the uniformly black colored species of Gobiodon are not monophyletic but have evolved independently within two distinct species groups. Genetic distances were large in particular within Pleurosicya and Eviota. Estimated divergence times suggest that coral-associated gobies have diversified in parallel to their preferred host corals. In particular, divergence times of Gobiodon species closely match those estimated for their typical host coral genus Acropora.     

Demographic parameters of two populations of red coral (Corallium rubrum L. 1758) in the North Western Mediterranean

The demographic and reproductive structure of populations represents the main data set needed for conservation and management plans. Mediterranean red coral Corallium rubrum has been exploited for 2,000 years, but only recently management plans have been solicited by the international community. We examined and compared the demographic features of two red coral shallow populations located in distinct geographic locations: Portofino (Italy) and Cap de Creus (Spain). Adults and juveniles density, growth rates, population size and age structure, fecundity and fertility were examined. Juveniles were the dominant class (33 %) in both populations. The analysis of the gamete content of 653 colonies revealed that the populations have balanced sex ratios and similar fertility and polyp fecundity. The average annual growth rate, determined on 119 colonies by annual growth rings count, was similar in both populations (0.24 mm year^?1), decreasing with colony age. Maximum life span of 99 % of the colonies was 60 and 40 years at Portofino and Cap de Creus, respectively. Minimum harvestable size (7 mm basal diameter) was reached in 30–35 years, and the percentage of colonies above it was 6.7 % at Portofino and 2.1 % at Cap de Creus, where juvenile and adult colony densities were significantly lower and the percentage of commercial-sized colonies reduced by 25 % in just a few years. Notwithstanding similar growth and fecundity, the two populations showed different densities and size/age structures suggesting local factors, together with different fishing pressures, have to be taken into account in the management plans for this species.     

Contrasting life histories and secondary production of populations of Munnopsurus atlanticus (Isopoda: Asellota) from two bathyal areas of the NE Atlantic and the NW Mediterranean

The isopod Munnopsurus atlanticus occupies bathyal depths in both the Bay of Biscay (NE Atlantic; between 383 and 1022 m) and in the Catalan Sea (Northwestern Mediterranean; between 389 and 1859 m). The species was dominant in both assemblages, reaching bathymetric peaks of abundance on the upper part of the continental slope (400 m depth) in the Bay of Biscay and at ˜600 m in the Catalan Sea. Both the Atlantic and the Mediterranean populations are bivoltines. Demographic analysis of the Bay of Biscay population revealed the production of two generations per year with different potential longevity (5 mo for G₁ and 11 mo for G₂). The mean cohort-production interval (CPI) was estimated at 8 mo, and results of the demographic analysis were also used to estimate production for the Catalan Sea populations. Mean annual density (D) and biomass (B) were higher in the Bay of Biscay (D = 356.7 individuals 100 m⁻²; B = 0.803 mg dry wt m⁻² yr⁻¹) than in the Mediterranean (D = 16.3 individuals 100 m⁻²; B = 0.078 mg dry wt m⁻² yr⁻¹). Also, mean annual production was an order of magnitude higher in the Atlantic (between 4.063 and 4.812 mg dry wt 100 m⁻² yr⁻¹ depending on the method used) than in the Catalan Sea (between 0.346 and 0.519 mg dry wt 100 m⁻² yr⁻¹). M. atlanticus feeds on a wide variety of benthic and pelagic food sources. In both study areas, phytodetritus was not important in the diet of M. atlanticus. In contrast, gut-content data suggested an indirect coupling with phytoplankton production in both areas via foraminiferans. The life history and the recorded production are considered in respect to both the dynamics and levels of primary production and the total mass flux in the respective study areas. Differences in the secondary production of both populations seemed to be more consistently explained by differences in total mass flux than by differences in the primary production levels; this is also consistent with the variety of food sources exploited by M. atlanticus. Received: 22 February 1999 / Accepted: 3 February 2000