Rates of molecular evolution in nuclear genes of east Mediterranean scorpions

Scorpions of the genus Mesobuthus represent a useful terrestrial model system for studying molecular evolution. They are distributed on several Aegean islands and the adjacent mainland, they are believed to have low rates of dispersal, and evolutionary divergence dates of taxa are available based on biogeographic events that separated islands from each other and the mainland. Here, we present data on polymorphism and synonymous (Ks) and non-synonymous (Ka) substitution rates for nine nuclear protein-coding genes of two east Mediterranean scorpion species, Mesobuthus gibbosus and M. cyprius (Buthidae). Levels of polymorphism tend to be lower in populations from islands (mean nucleotide diversity pi = 0.0071 +/- 0.0028) than in mainland populations (mean pi = 0.0201 +/- 0.0085). By using linear regression of genetic divergence versus isolation time, we estimate Ks to be 3.17 +/- 1.54 per (site x 10(9) years), and Ka to be 0.39 +/- 0.94 per (site x 10(9) years). These estimates for both Ks and Ka are considerably lower than for many other invertebrates, such as Drosophila, and may be attributed to scorpions' mammal-like generation times (approximately 2 years) and low metabolic rates. Phylogenetic analysis using maximum likelihood revealed a phylogeny that is congruent with that expected based on biogeographic events and in which divergences at synonymous sites are proportional to the dates that the taxa are believed to have split. Tests of equality of branch lengths for the Cyprus and Crete lineages revealed that Ks-estimates are about the same in both lineages, as expected from the biogeographic events that separated the islands, but Ka was increased in the Cyprus lineage compared to the Cretan lineage.     

Messinian vegetation maps of the Mediterranean region using models and interpolated pollen data

This study proposes to compare the outputs from the CARAIB vegetation model forced by results from the LMD General Circulation Model with interpolated pollen data (Kriging method) from the Mediterranean region during the Messinian. The vegetation maps that have been obtained represent distinct phases of the salinity crisis: before the crisis and during the marginal evaporitic phase (interpolated map), and during the complete desiccation phase (simulated map). However, they are comparable in terms of vegetation density and agree on a strong contrast between the Northern (forest vegetation) and Southern (open vegetation) Mediterranean regions. Main differences concern the type of forests in the northern Mediterranean region, which are explained by discrepancies between precipitation amount predicted by the model and that calculated by a transfer function using pollen records. The interpolation method has been successfully tested in France using interpolated current pollen records by comparison with the present-day potential vegetation map. The resulting Messinian map is useful to validate or improve model simulation which does not take into account the depth of the Mediterranean Basin when it dried up. The Southern Mediterranean landscapes were open, with a steppe-like vegetation to the West and a savannah-like vegetation to the East. Forests prevailed to the North, organized in a mosaic system mainly controlled by relief. Such a contrast provides some explanation of the large number of deep fluvial canyons cut on the Northern margin at opposed to the South during the Mediterranean desiccation.     

The coastal plant communities of Juniperus macrocarpa in the Mediterranean region

Juniperus macrocarpa is a threatened taxon growing on coastal areas of the Mediterranean region. A remarkable part of its communities are included in Juniperion turbinatae, considered as priority habitats for conservation in the Habitats Directive 92/43 EEC. We have summarized the current status of this type of vegetation, including the described associations of this alliance in order to analyse the variability of the syntaxa, based on its floristic composition and geographic distribution. The available relevés were arranged in a synthetic table and subjected to a detendred correspondence analysis. Two new suballiances (Juniperenion turbinatae and Asparago orientalis-Juniperenion macrocarpae) have been proposed. A typological classification of these communities with ecological and biogeographical diagnoses is given.     

Seasonal variability of copepod abundance in the Balearic region (Western Mediterranean) as an indicator of basin scale hydrological changes

Spatial and temporal changes of the copepod community have been investigated and related to the environmental variability of the Balearic Sea (Western Mediterranean). The period studied spans from 1994 to 1999 during which we analyzed the abundance and structure copepod variability over a cross-shore transect. Results showed a close link between hydrological changes and the variations of copepod abundance. The synchronous variability of copepods and hydrography indicated the rapid response of this zooplankton group to the inflow of cold and warm water masses coming through the study area. Cluster analysis revealed four main copepod assemblages that distinguished the coastal from the oceanic species and those species with different water masses preference. The copepod assemblage composed of Calanus helgolandicus, Clausocalanus arcuicornis, C. pergens, C. paululus, Calocalanus tenuis and Pleuromamma gracilis was associated with cool salty waters, whereas the assemblage formed by Temora stylifera, C. pavo, C. styliremis, Centropages bradyi and Acartia danae was related to warmer less saline Mediterranean waters. Moreover, it is suggested that changes in sea water temperature and salinity are linked to large-scale changes likely occurring at a basin scale, which is reflected in the Western Mediterranean mesoscale hydrographic changes. Therefore, it is stressed that changes in the Balearic copepod community can be used as potential tracers of the western Mediterranean water masses. Guest editors: S. Souissi & G. A. Boxshall Copepoda in the Mediterranean: Papers from the 9th International Conference on Copepoda, Hammamet, Tunisia     

Biogeographic patterns in Mediterranean and Macaronesian species of Saxifraga (Saxifragaceae) inferred from phylogenetic analyses of ITS sequences

A biogeographic study of Saxifraga section Saxifraga was performed based on phylogenetic analyses of ITS (internal transcribed spacer) sequences of nuclear ribosomal DNA. ITS sequences from 21 species and 31 populations were examined to identify colonization patterns for the two species of Saxifraga occurring in Macaronesia and for S. globulifera in the west Mediterranean basin. Phylogenetic analysis of the sequence data yield a single most parsimonious tree with many of the major clades well supported by bootstrap and decay values. The ITS tree provided resolution at specific and populational levels that points to two biogeographic patterns within the genus. In contrast to the molecular evidence provided by other authors for a Mediterranean origin of several Macaronesian genera of angiosperms, our results indicate that the Madeiran archipelago was colonized a single time by a species of Saxifraga originating from the Eurosiberian region. On the other hand, the molecular evidence also suggests that populations of S. globulifera from North Africa have been isolated for a long time from populations occurring in the Iberian Peninsula, and that the endemic S. reuteriana has evolved from the Iberian populations of S. globulifera. The Mediterranean Sea has probably been an effective isolating barrier for some plant groups that occur in Europe and North Africa.     

The distribution patterns of reptiles in the Riff region, northern Morocco

A biogeographical classification of reptiles in the Riff region (northern Morocco, Africa) was carried out to look for shared distribution patterns, here termed chorotypes. Baroni‐Urbani & Buser's similarity index was applied to the presence/absence data of reptiles in 10×10 km UTM (Universal Transverse Mercator) squares, and then UPGMA (Unweighted Pair‐Group Method using arithmetic average) was used to classify the species. A probabilistic method was employed to assess the statistical significance of the groups obtained. An ordination method, the Canonical Correspondence Analysis, was also used to study the distribution of the reptiles within a continuous framework.
A gradual longitudinal replacement of reptile species was found throughout the Riff, with no sharp discontinuities for the distributions of most of the species. This may be due to the biogeographical northward movement of the Saharan boundaries, which have not yet reached biogeographical equilibrium. Thus, Saharan reptiles enter the Riff region from the east, through the lower basin of the River Moulouya.
Seven reptile chorotypes were identified in the Riff, and these comprise Mediterranean species and others endemic to the Maghreb (the region that spans most of North‐western Africa, excluding the Sahara). These chorotypes have a western distribution, and are segregated from one another according to altitude. Historical and ecological processes can account for the distributions shared by these species, which have inhabited the Riff for longer than eastern reptiles.     

Thaliacean distribution and abundance in the northern part of the Levantine Sea (Crete and Cyprus) during the eastern Mediterranean climatic transient, and a comparison with the western Mediterranean basin

First results are presented on the composition, abundance and vertical distribution of the thaliacean fauna in the Levantine basin obtained from stratified tows at three deep-sea sites in 1993: SE off Crete, and SW and NE off Cyprus. Samples with a 10 m²-MOCNESS (mesh size 1.67 mm) were poor in species and specimens as compared to samples with a 1 m²-double-MOCNESS (0.333 mm). Of the 12 species identified, six species belonged to the most abundant Doliolida, predominated by the phorozooids of Doliolum nationalis, five species belonged to the Salpida and one to the Pyrosomatida. Thaliaceans, most abundant by species and numbers SE off Crete, comprised ≤0.2% of the local mesozooplankton standing stocks. Presumably, they did not contribute substantially to the vertical flux generating the locally increased biomass and activity of the microbial benthos at the 4,300 m deep Cretean site. Most doliolids and salps were collected from the epipelagic and upper mesopelagic layers, and appeared to be most abundant close to the deep maximum of chlorophyll-a. Samples below 150 m were rarely rich in specimens, although two species performed diel migrations from the mesopelagic zone into the surface layers where some vertical segregation was evidenced. The aspect of niche separation is discussed.     

Micro-scale distribution of some arthropods inhabiting a Mediterranean sandy beach in relation to environmental parameters

A study on the spatial distribution of two tenebrionids and an isopod species was conducted at a small extent. Two species of the genus Phaleria were chosen because of their sympatric distribution and apparently similar adaptations to the beach environment. These scavenger species were studied in association to Tylos europaeus. Distributions, both perpendicular and parallel to the shoreline, were studied simultaneously with a grid of 20 × 20 pitfall traps arranged at a distance of 1 m from each other. Faunal samples were collected at two different hours of the night and core samples of sand were taken next to each pitfall trap for successive laboratory analysis. An index of dispersion was applied to test for aggregation of all species and of environmental parameters on the total, across- and long-shore. Kolmogorov–Smirnov two-sample test was used to compare the distributions between species. Regression analyse were applied to evaluate relationships between environmental parameters and species distributions. The study showed that the distribution of Phaleria species and T. europaeus was scale-dependent. The species and the values of the environmental parameters showed aggregated distributions both across- and long-shore. Aggregations of each Phaleria species differed according to their resting and foraging phases. For all species, sand moisture and salinity were the most important parameters explaining distribution. Granulometric parameters were correlated with the choice of the resting areas of both Phaleria species, whereas the spatial distribution of T. europaeus was correlated to the distribution of the organic matter.     

Biogeographical network analysis of plant species distribution in the Mediterranean region

The delimitation of bioregions helps to understand historical and ecological drivers of species distribution. In this work, we performed a network analysis of the spatial distribution patterns of plants in south of France (Languedoc‐Roussillon and Provence‐Alpes‐Côte d'Azur) to analyze the biogeographical structure of the French Mediterranean flora at different scales. We used a network approach to identify and characterize biogeographical regions, based on a large database containing 2.5 million of geolocalized plant records corresponding to more than 3,500 plant species. This methodology is performed following five steps, from the biogeographical bipartite network construction to the identification of biogeographical regions under the form of spatial network communities, the analysis of their interactions, and the identification of clusters of plant species based on the species contribution to the biogeographical regions. First, we identified two sub‐networks that distinguish Mediterranean and temperate biota. Then, we separated eight statistically significant bioregions that present a complex spatial structure. Some of them are spatially well delimited and match with particular geological entities. On the other hand, fuzzy transitions arise between adjacent bioregions that share a common geological setting, but are spread along a climatic gradient. The proposed network approach illustrates the biogeographical structure of the flora in southern France and provides precise insights into the relationships between bioregions. This approach sheds light on ecological drivers shaping the distribution of Mediterranean biota: The interplay between a climatic gradient and geological substrate shapes biodiversity patterns. Finally, this work exemplifies why fragmented distributions are common in the Mediterranean region, isolating groups of species that share a similar eco‐evolutionary history.     

Profile of cancer in the Eastern Mediterranean region: The need for action

BackgroundMany countries in the Eastern Mediterranean region (EMR) are undergoing marked demographic and socioeconomic transitions that are increasing the cancer burden in region. We sought to examine the national cancer incidence and mortality profiles as a support to regional cancer control planning in the EMR.MethodsGLOBOCAN 2012 data were used to estimate cancer incidence and mortality by country, cancer type, sex and age in 22 EMR countries. We calculated age-standardized incidence and mortality rates (per 100,000) using direct method of standardization.ResultsThe cancer incidence and mortality rates vary considerably between countries in the EMR. Incidence rates were highest in Lebanon (204 and 193 per 100,000 in males and females, respectively). Mortality rates were highest in Lebanon (119) and Egypt (121) among males and in Somalia (117) among females. The profile of common cancers differs substantially by sex. For females, breast cancer is the most common cancer in all 22 countries, followed by cervical cancer, which ranks high only in the lower-income countries in the region. For males, lung, prostate, and colorectal cancer in combination represent almost 30% of the cancer burden in countries that have attained very high levels of human development.ConclusionsThe most common cancers are largely amenable to preventive strategies by primary and/or secondary prevention, hence a need for effective interventions tackling lifestyle risk factors and infections. The high mortality observed from breast and cervical cancer highlights the need to break the stigmas and improve awareness surrounding these cancers.     

Refinements of calcareous nannofossil biostratigraphy at the Miocene/Pliocene Boundary in the Mediterranean region

The high-resolution quantitative study of the calcareous nannofossil assemblages in two Mediterranean deep-sea successions (ODP Sites 969B and 975B) encompassing the Miocene/Pliocene boundary allows the recognition of a set of reliable bioevents useful to detecting the base of the Zanclean stage. The results have been successfully compared with two on-land sections (Cava Serredi, Tuscany, and Montepetra borehole, Marche Region, Central Italy). This study confirms that Ceratolithus acutus and Triquetrorhabdulus rugosus, the markers traditionally used for identifying the Miocene/Pliocene Boundary are very rare in the Mediterranean area and cannot be used for biostratigraphic correlation. Conversely, the absence interval (paracme) of Reticulofenestra pseudoumbilicus and the distribution range of a new species belonging to the Reticulofenestra genus (Reticulofenestra zancleana nov. sp.) show high biostratigraphical potential. The position of the new biohorizons has been compared to those of the planktonic foraminifers events, and correlated to the CaCO₃ cycles reconstructed for the two sections. On the basis of these new nannofossil bioevents, Rio et al.’s (1990) MNN12 biozone is subdivided into three subzones, thus improving the biostratigraphic resolution of the Early Pliocene.     

The fish assemblage on a coralligenous shallow shelf off the Mediterranean coast of northern Israel

The fish assemblage on the shallow coralligenous shelf (16-30m depth) off Haifa, Israel was sampled using trammel nets throughout a period of 1 year. Complementary data were obtained via underwater censuses of fish on an artificial reef established later in the sampling area. Fortythree species of fish were sampled by trammel nets, 79% of which were observed also during the underwater censuses. Although fish of Red Sea origin constituted only 11.6% of the species composition in the net samples, they contributed 46.2% of the fish abundance and 40.6% of the biomass in these samples. This was supported by the finding that species of Red Sea origin contributed 64% of the abundance of large fish counted on the artificial reef. Siganus luridas, S. rivulatus and Sargocentron rubrum are the main contributors in number and biomass among fish of Red Sea origin. It is suggested that the biogenic rocky bottom of this area contains several components which are similar to biogenic habitats favoured by these benthic species in the Red Sea; this may explain the high abundance of these species in the studied area.     

Structure of ground-foraging ant assemblages in relation to land-use change in the northwestern Mediterranean region

The abandonment by humans of marginal and less productive zones signifies an important change in land use in North Mediterranean agroecosystems. Human perturbations have led to a highly diversified landscape, with a mosaic made up of patches of land at different stages of succession, from cultivated fields to closed forest. Our aim here is to characterize ant assemblages and their functional groups in response to these land-use changes. This progressive abandonment results in an initial increase in ant richness and abundance, which can reach high levels if the succession proceeds as far as woodland. In terms of the ant functional groups, this land-use change implies: (1) the appearance of Subordinate Camponotini; (2) an increase in Generalized Myrmicinae, Cryptics and Cold-climate specialists in terms of ant species richness, overall abundance and, for Generalized Myrmicinae and Cryptics, an increase in abundance percentage; (3) a decrease in percentage abundance of Opportunists; (4) a progressive decrease in species richness as well as overall and percentage abundance of Hot-climate Specialists throughout the transformation from crops to woodlands; and (5) an initial increase of Dominant Dolichoderinae followed by a decrease in ant species richness, overall abundance and percentage abundance. Using the ant functional group approach for the clearly separate stages of the regeneration process is a promising method for comparing responses of ant communities to human land use.     

Biogeography of the marine birds of a confined sea, the Mediterranean

Aim The Mediterranean sea is a winter productive oligotrophic basin where Atlantic water replaces water lost through evaporation, this influx being a major source of productivity and fertility. The long coastlines and the large number of islands cause high oceanographic heterogeneity. Moreover, during its geological history, it has dried out several times. So we describe the consequences of these particular features on species richness, distribution, and breeding ecology of marine birds. Location The Mediterranean sea (including the Black Sea and the Sea of Azov) communicates with the Atlantic Ocean only through a 14 km wide channel (Straits of Gibraltar), and since 1869, with the Red Sea through the Suez Canal. Methods The Mediterranean was subdivided into different areas, according to physical oceanographic entities and productivity, linked to numbers and distribution of both breeding and wintering marine birds (defined as species strongly dependent on marine resources, breeding only on islands and/or the coastline). Results The total marine bird biomass, and species diversity, are lower in the Mediterranean than in the near Atlantic. The eastern Mediterranean, with lowest primary productivity, contains fewer marine bird taxa than the more productive western part. Taxa which mainly occur in the western and southern parts of the Mediterranean migrate through the Straits of Gibraltar to winter in the southern Atlantic, while those inhabiting the northern and eastern parts are sedentary, as a result of differences in species composition. Northern coastal basin communities (i.e. the Tyrrhenian and the Balearic Seas), are composed of less pelagic, and earlier breeding species, that rear chicks during the productive season. These latter taxa are actually the most typical Mediterranean taxa, in terms of endemism. Main conclusions The Mediterranean marine bird community is not tropical, but rather, shows the highest affinity with the Atlantic temperate community. Its level of endemism is however high and comparable to other confined basins such as the Red Sea.     

Mediterranean island biogeography: Analysis of fern species distribution in the system of islets around Sicily

Abstract

The aim of this article was to test the way in which geographical factors influence island floras in the Mediterranean basin, using ferns as target organisms, and the islands surrounding Sicily as location. A matrix with presence/absence data concerning fern taxa in the 16 islands studied was compiled. Cluster analysis, principal co-ordinates analysis (PCoA), principal components analysis (PCA) and a Bayesian analysis were performed. For each island, the total number of fern taxa was regressed against three factors: island area, island elevation and isolation. All the analyses pointed to affinities between islands according to their different geological composition, independently from their geographic position. A clear positive island species/area relationship (ISAR) was shown only for the volcanic islands. The island species/(area×elevation) relationship (ISAER), on the contrary, was unsatisfactory. The main features of interest are the following: (1) the clear division of the islands into two groups, volcanic vs. sedimentary; (2) the floristic richness of the volcanic compared to sedimentary islands and (3) the uniqueness of the pteridophyte flora of Pantelleria. This seems to demonstrate that the lower number of taxa in the islands farthest away from the “mainland” (Sicily, Tunisia) is not due to isolation, but due to another factor, probably habitat availability.     

Climate change and fitness components of a migratory bird breeding in the Mediterranean region

The increase in spring temperatures in temperate regions over the last two decades has led to an advancing spring phenology, and most resident birds have responded to it by advancing their onset of breeding. The pied flycatcher (Ficedula hypoleuca) is a long‐distance migrant bird with a relatively late onset of breeding with respect to both resident birds and spring phenology in Europe. In the present correlational study, we show that some fitness components of pied flycatchers are suffering from climate change in two of the southernmost European breeding populations. In both montane study areas, temperature during May increased between 1980 and 2000 and an advancement of oak leafing was detected by using the normalized difference vegetation index (NDVI) to assess tree phenology. This might result in an advancement of the peak in availability of caterpillars, the main prey during the nestling stage. Over the past 18 yr, the time of egg laying and clutch size of pied flycatchers were not affected by the increase in spring temperatures in these Mediterranean populations. However, this increase seems to have an adverse effect on the reproductive output of pied flycatchers over the same period. Our data suggest that the mismatch between the timing of peak food supply and nestling demand caused by recent climate change might result in a reduction of parental energy expenditure that is reflected in a reduction of nestling growth and survival of fledged young in our study populations. The data seem to indicate that the breeding season has not shifted and it is the environment that has shifted away from the timing of the pied flycatcher breeding season. Mediterranean pied flycatchers were not able to advance their onset of breeding, probably, because they are constrained by their late arrival date and their restricted high altitude breeding habitat selection near the southern border of their range.     

Association between Opuntia species invasion and changes in land-cover in the Mediterranean region

In Mediterranean regions, biological invasions pose a major threat to the conservation of native species and the integrity of ecosystems. In addition, changes in land‐cover are a widespread phenomenon in Mediterranean regions, where an increase in urban areas and major changes from agricultural abandonment to shrub encroachment and afforestation are occurring. However, the link between biological invasions and changes in land‐cover has scarcely been analyzed. We conducted a regional survey of the distribution of the two alien prickly‐pear cacti Opuntia maxima and O. stricta in Cap de Creus (Catalonia, Spain) and related patterns of invasion to spatially explicit data on land‐cover/change from 1973 to 1993 to test the hypotheses that the two Opuntia species invade areas that have experienced large land‐cover transformations. We found that Opuntia invasion is particularly high in shrublands and woodlands located near urban areas. O. maxima are over‐represented in the shrublands and O. stricta in the woodlands that were former crops. Crop coverage has dropped by 71% in this 20‐year period. This study highlights the role of past land‐cover in understanding the present distribution of plant invasions.     

Reevaluating sponge diversity and distribution in the Mediterranean Sea

The aim of this paper was (1) to update sponge diversity and distribution in the Mediterranean and (2) to re-examine faunal relationships among the Mediterranean areas on the basis of their sponge fauna. The Mediterranean demosponge faunal list was updated to 629 species by taking into consideration recent data from previously poorly studied areas. The species lists of 14 Mediterranean areas were compared on the basis of their sponge species richness, species composition, and taxonomic relatedness of species using multivariate analyses and diversity measures, such as PD, Delta+, and Lambda+. The 14 Mediterranean areas examined for their diversity and affinities were assembled into four major zoogeographic groups: the northwestern, northeastern, the central zone, and southeastern areas. Richest in species numbers were the areas belonging to the two northern groups. The species richness comparisons and similarity analyses performed at the generic level showed that it can be safely used as a surrogate for sponge species diversity in the Mediterranean. The results of this study showed that the simple traditional division of the Mediterranean Sea into a western, central, and eastern basin cannot reliably describe the distribution of sponges in the area. Thus, the W to E faunal decline previously presented for several faunal groups shifts to a general NNW-SSE pattern when one examines separately the northern and the southern parts of the traditional basins. This gradient seems to be in agreement with differences in key environmental variables, such as latitude, salinity, temperature, and water circulation, besides the typically examined distance from Gibraltar. Handling editor: T. P. Crowe     

Identification of deep-water pink shrimp abundance distribution patterns and nursery grounds in the eastern Mediterranean by means of generalized additive modelling

Generalized Additive Modelling (GAM) techniques were used to model the time series of abundance data of deep-water pink shrimp, collected during the MEDITS bottom trawl surveys carried out in the Greek seas from 1996 to 2006, as functions of the sampling location (longitude–latitude), depth and year. The life stages of the species were taken into account. All variables were highly significant, although latitude and depth had always the highest explanatory power. The total abundance was higher between 100 and 400 m of depth, with juveniles and adults being more abundant in the 100–200 m and 200–400 m depth strata, respectively. GAM prediction maps showed high abundance concentration areas for all life stages mainly in the Aegean Sea. The most important nursery ground identified was located in the Saronikos Gulf and a secondary in the Thracian Sea. Concerning the concentration areas of the adult specimens, they are located mainly in the Saronikos Gulf, the Thracian Sea, the Thermaikos Gulf, the Cretan Sea and the eastern part of the Aegean. Guest Editor: V. D. Valavanis Essential Fish Habitat Mapping in the Mediterranean