Habitat niche breadth predicts invasiveness in solitary ascidians

A major focus of invasion biology is understanding the traits associated with introduction success. Most studies assess these traits in the invaded region, while only few compare nonindigenous species to the pool of potential invaders in their native region. We focused on the niche breadth hypothesis, commonly evoked but seldom tested, which states that generalist species are more likely to become introduced as they are capable of thriving under a wide set of conditions. Based on the massive introduction of tropical species into the Mediterranean via the Suez Canal (Lessepsian migration), we defined ascidians in the Red Sea as the pool of potential invaders. We constructed unique settlement plates, each representing six different niches, to assess ascidian niche breadth, and deployed them in similar habitats in the native and invaded regions. For each species found on plates, we evaluated its abundance, relative abundance across successional stages, and niche breadth, and then compared (1) species in the Red Sea known to have been introduced into the Mediterranean (Lessepsian species) and those not known from the Mediterranean (non‐Lessepsian); and (2) nonindigenous and indigenous species in the Mediterranean. Lessepsian species identified on plates in the Red Sea demonstrated wider niche breadth than non‐Lessepsian species, supporting the niche breadth hypothesis within the native region. No differences were found between Lessepsian and non‐Lessepsian species in species abundance and successional stages. In the Mediterranean, nonindigenous species numerically dominated the settlement plates. This precluded robust comparisons of niche breadth between nonindigenous and indigenous species in the invaded region. In conclusion, using Red Sea ascidians as the pool of potential invaders, we found clear evidence supporting the niche breadth hypothesis in the native region. We suggest that such patterns may often be obscured when conducting trait‐based studies in the invaded regions alone. Our findings indicate that quantifying the niche breadth of species in their native regions will improve estimates of invasiveness potential.     

The rDNA ITS Region in the Lessepsian Marine Angiosperm <Emphasis Type="BoldItalic">Halophila stipulacea</Emphasis> (Forssk.) Aschers. (Hydrocharitaceae): Intragenomic Variability and Putative Pseudogenic Sequences

Halophila stipulacea is a dioecious marine angiosperm, widely distributed along the western coasts of the Indian Ocean and the Red Sea. This species is thought to be a Lessepsian immigrant that entered the Mediterranean Sea from the Red Sea after the opening of the Suez Canal (1869). Previous studies have revealed both high phenotypic and genetic variability in Halophila stipulacea populations from the western Mediterranean basin. In order to test the hypothesis of a Lessepsian introduction, we compare genetic polymorphism between putative native (Red Sea) and introduced (Mediterranean) populations through rDNA ITS region (ITS1-5.8S-ITS2) sequence analysis. A high degree of intraindividual variability of ITS sequences was found. Most of the intragenomic polymorphism was due to pseudogenic sequences, present in almost all individuals. Features of ITS functional sequences and pseudogenes are described. Possible causes for the lack of homogenization of ITS paralogues within individuals are discussed.     

Diet composition of the Lessepsian bluespotted cornetfish Fistularia commersonii in the eastern Mediterranean

Adaptability is an important factor in defining success or failure of an introduced species. The bluespotted cornetfish, Fistularia commersonii , invaded the Mediterranean from the Red Sea and spread quickly, displaying an explosive growth pattern in the eastern and central Mediterranean. Stomach contents of the Lessepsian piscivore were studied from off the Lebanese coast in order to describe aspects of its feeding ecology in the new environment. Cornetfish of different sizes (25–112 cm L_t) and from various habitats and depths (1–40 m) were sampled over an 18-month period (May 2003 to November 2004). Only 29% of the individuals contained identifiable prey items. The diet of the invasive cornetfish comprised a large variety of prey (41 taxa), Spicara smaris and Boops boops being the most important. Comparisons among prey habitats showed that the invasive cornetfish fed preferably on schooling species living in the water column. A large diversity of prey species characterized autumn and winter seasons while S. smaris and B. boops dominated in the spring and summer. Diet did not vary significantly with predator size except for S. smaris , which was a more important prey to middle size-classes cornetfish. The invasion of F. commersonii might have an important effect on structure and population dynamics of native communities, however it seems premature to predict the potential impact on eastern Mediterranean ecosystems or fisheries.     

Molecular analysis of the recently described lizardfish Saurida lessepsianus (Synodontidae) from the Red Sea and the Mediterranean,with remarks on its phylogeny and genetic bottleneck effect

The opening of the Suez Canal in 1869 led to a massive influx of Red Sea species that invaded the Mediterranean; this was termed ‘Lessepsian migration'. Among these species was a species of lizardfish, identified by some authors as Saurida undosquamis and by others as S. macrolepis. Recently, the Red Sea and the Mediterranean populations were described according to external characteristics as a unique taxon, Saurida lessepsianus. Our molecular study confirms this finding and determines that all previous records of S. undosquamis and S. macrolepis in the Red Sea and the Mediterranean are misidentifications of S. lessepsianus. The Mediterranean population of S. lessepsianus exhibits a lower genetic variability than that of the Red Sea population, suggesting a bottleneck effect.     

Molecular Evidence for Lessepsian Invasion of Soritids (Larger Symbiont Bearing Benthic Foraminifera)

The Mediterranean Sea is considered as one of the hotspots of marine bioinvasions, largely due to the influx of tropical species migrating through the Suez Canal, so-called Lessepsian migrants. Several cases of Lessepsian migration have been documented recently, however, little is known about the ecological characteristics of the migrating species and their aptitude to colonize the new areas. This study focused on Red Sea soritids, larger symbiont-bearing benthic foraminifera (LBF) that are indicative of tropical and subtropical environments and were recently found in the Israeli coast of the Eastern Mediterranean. We combined molecular phylogenetic analyses of soritids and their algal symbionts as well as network analysis of Sorites orbiculus Forskål to compare populations from the Gulf of Elat (northern Red Sea) and from a known hotspot in Shikmona (northern Israel) that consists of a single population of S. orbiculus. Our phylogenetic analyses show that all specimens found in Shikmona are genetically identical to a population of S. orbiculus living on a similar shallow water pebbles habitat in the Gulf of Elat. Our analyses also show that the symbionts found in Shikmona and Elat soritids belong to the Symbiodinium clade F5, which is common in the Red Sea and also present in the Indian Ocean and Caribbean Sea. Our study therefore provides the first genetic and ecological evidences that indicate that modern population of soritids found on the Mediterranean coast of Israel is probably Lessepsian, and is less likely the descendant of a native ancient Mediterranean species.     

The littoral fish community of the Lebanese rocky coast (eastern Mediterranean Sea) with emphasis on Red Sea immigrants

The organisation of the coastal rocky fish community of Lebanon was investigated for the first time, using visual censuses. A total of 62 fish species were recorded between the surface and 32 m depth, 8 species being Red Sea migrants. Species richness and fish abundance were positively correlated first with substrate complexity and second with depth. The trophic structure of the community was dominated by diurnal zooplanktivores (Chromis chromis, Oblada melanura and Spicara smaris) and mesocarnivores 1 (Coris julis and Thalassoma pavo), and did not vary significantly with depth. The Lessepsian migrants represented 13% of the species richness and 19% of the total abundance of individuals. They dominated among herbivores, nocturnal zooplanktivores and macrocarnivores. The Lessepsian Siganus luridus and S. rivulatus (Siganidae), Pempheris vanicolensis (Pempheridae) and Sargocentron rubrum (Holocentridae) were now among the most common fish species on the Lebanese rocky coast. Lessepsian fish species displayed either similar ecological niches in the Red Sea and in the Levantine basin (P. vanicolensis, S. rubrum) or an enlargement of their depth distribution towards deeper waters in their new environment (S. luridus, S. rivulatus). This study will serve as a baseline for future studies as eastern Mediterranean communities are subjected to a steady increase in Red Sea migrant species.     

Patterns of ecological specialization among microbial populations in the Red Sea and diverse oligotrophic marine environments

Large swaths of the nutrient‐poor surface ocean are dominated numerically by cyanobacteria (Prochlorococcus), cyanobacterial viruses (cyanophage), and alphaproteobacteria (SAR11). How these groups thrive in the diverse physicochemical environments of different oceanic regions remains poorly understood. Comparative metagenomics can reveal adaptive responses linked to ecosystem‐specific selective pressures. The Red Sea is well‐suited for studying adaptation of pelagic‐microbes, with salinities, temperatures, and light levels at the extreme end for the surface ocean, and low nutrient concentrations, yet no metagenomic studies have been done there. The Red Sea (high salinity, high light, low N and P) compares favorably with the Mediterranean Sea (high salinity, low P), Sargasso Sea (low P), and North Pacific Subtropical Gyre (high light, low N). We quantified the relative abundance of genetic functions among Prochlorococcus, cyanophage, and SAR11 from these four regions. Gene frequencies indicate selection for phosphorus acquisition (Mediterranean/Sargasso), DNA repair and high‐light responses (Red Sea/Pacific Prochlorococcus), and osmolyte C1 oxidation (Red Sea/Mediterranean SAR11). The unexpected connection between salinity‐dependent osmolyte production and SAR11 C1 metabolism represents a potentially major coevolutionary adaptation and biogeochemical flux. Among Prochlorococcus and cyanophage, genes enriched in specific environments had ecotype distributions similar to nonenriched genes, suggesting that inter‐ecotype gene transfer is not a major source of environment‐specific adaptation. Clustering of metagenomes using gene frequencies shows similarities in populations (Red Sea with Pacific, Mediterranean with Sargasso) that belie their geographic distances. Taken together, the genetic functions enriched in specific environments indicate competitive strategies for maintaining carrying capacity in the face of physical stressors and low nutrient availability.     

Lessepsian invasion without bottleneck: example of two rabbitfish species (Siganus rivulatus and Siganus luridus)

Since the opening of the Suez Canal in the 19th century, a growing number of teleosts species have become permanently established in the eastern Mediterranean. As environmental parameters in the Mediterranean are very different from those in the Red Sea, there is an opportunity to evaluate the gene flow associated with the acclimatisation of these species to their new environment. Initial colonisation may have been restricted to individuals with marginal genotypes compared to populations of the Red Sea, eventually better adapted to the Mediterranean environment. Alternatively, colonisation may have occurred without major genetic differentiation, due to a large flux of immigrants and a continuous gene flow from the populations of origin. Using different molecular genetic techniques (mitochondrial DNA, Exon-Primed Intron-Crossing PCR amplification (EPIC) and Inter-Simple Sequence Repeats (ISSRs), patterns of genetic differentiation on both sides of the Suez isthmus were tested on two Lessepsian rabbitfish species (Siganus rivulatus and Siganus luridus). The absence of genetic differentiation between Mediterranean and Red Sea populations in both species showed that a great number of migrants participated to the colonisation of the Mediterranean, excluding any bottleneck event. This colonisation success suggests large eco-physiological plasticity in S. rivulatus and S. luridus that allowed them to thrive in the new environment.     

Lessepsian fish migration: genetic bottlenecks and parasitological evidence

As a rule, non‐indigenous species (NIS) populations derived from biological invasion events represent a subset of the genetic diversity of the source population. In biological invasions, host–parasite interactions play an important role, and parasitological data for NIS populations can provide useful information such as their area of origin, mechanism of invasion and prospects of success in the new habitat. When both genetic and parasitological data are available, and they suggest the same scenario, the history of an invasion can be inferred with no discrepancy, but when data cannot be reconciled an alternative model should be considered. In this study a comparison of genetic and parasitological data for the Lessepsian migrant the bluespotted cornetfish, Fistularia commersonii, in the Mediterranean Sea presents the opportunity to evaluate the compatibility of information of this nature, and to propose possible invasion scenarios consistent with evidence provided by both criteria.     

Lessepsian migrant Fistularia commersonii from the Rhodes marine area

The bluespotted cornetfish Fistularia commersonii which originates from the Red Sea, appeared in the marine area of Rhodes Island (south-east Aegean Sea). This is a new record of the species based on 37 specimens caught during summer-winter 2001. The species appears to be spreading rapidly west and easily becoming established in Mediterranean coastal habitats.     

No reduction in neutral variability of mitochondrial and nuclear genes for a Lessepsian migrant, Upeneus moluccensis

Using nuclear (introns) and mitochondrial markers, two populations of the Lessepsian fish Upeneus moluccensis sampled from its original (Red Sea) and new habitat (Mediterranean Sea) were compared. No significant genetic difference was observed hence the passage through the Suez Canal does not seem to have affected the genetic diversity at the presumably neutral loci studied.     

First record of the bluespotted cornetfish from the Mediterranean Sea

Three specimens of the Indo-Pacific Bluespotted cornetfish Fistularia commersonii are recorded for the first time from the Mediterranean. The presence of this species in the Mediterranean is due to migration from the Red Sea via the Suez Canal.     

Genetic bottlenecks and successful biological invasions: the case of a recent Lessepsian migrant

Our current understanding of the mechanisms that lead to successful biological invasions is limited. Although adaptations play a central role in biological invasions, genetic studies have so far failed to produce a unified theory. The bluespotted cornetfish, a recent Red Sea invader in the Mediterranean Sea via the Suez Canal, provides an ideal case study for research in the mechanisms of invasive genetics. In this study, we show that the invading bluespotted cornetfish underwent a severe population bottleneck that reduced the genetic diversity of this immigrant to only two mitochondrial haplotypes. Although loss of genetic diversity is considered detrimental to the need to adapt to new environments, bluespotted cornetfish experienced an unprecedented success and rapid spread across the Mediterranean.     

Westward range expansion of the Lessepsian migrant Fistularia commersonii (Fistulariidae) in the Mediterranean Sea, with notes on its parasites

One adult female bluespotted cornetfish Fistularia commersonii was captured near the coast of Arbatax (Sardinia, Italy) in late October 2005. Macroscopic and microscopic analysis of the gonad revealed a postspawning ovary. Several body parts of the damselfish Chromis chromis were found in its stomach contents. Different developmental stages of 58 metazoan parasites belonging to Arthropoda, Nematoda and Platyhelminthes were detected in its mouth and digestive tract. The lepocreadiid Allolepidapedon fistulariae was reported for the first time from the Mediterranean and, with the exception of this species, all the parasites found were reported for the first time in this host. This new record of F. commersonii may confirm the rapid range expansion of this Lessepsian migrant in the western Mediterranean Sea, as well as the presence of its endoparasite A. fistulariae .     

Trypauchen vagina (Bloch & Schneider, 1801) a new established species in the Southwestern Atlantic

Trypauchen vagina (Bloch & Schneider, 1801) is a goby that lives burrowed into the substrata feeding on small invertebrates. It is native to the Indo-pacific region, ranging from Kuwait to China. Recently, this fish has been reported outside the original range of distribution, being found in the Mediterranean Sea, and more recently in the northeastern Brazilian coast. The Mediterranean reports are usually associated with Lessepsian migration, while the reports from Brazil are possibly related to ballast water transportation. In the present work, we provide eight new records from southeastern Brazilian coast, all made in São Paulo state, far from the first record. These additional records raised concern since the presence of an alien species could implicate in environmental and economic losses. Thus, we decide to model the environmental suitability for this goby in the Brazilian coast, specially focusing on major ports, usually places with high ballast water propagule pressure. In addition, an analysis of the suitability in the Red Sea was also made, to verify the hypothesis of Lessepsian migration. The results revealed that temperature and primary productivity are among the most important parameters for the presence of T. vagina, also indicating a high environmental suitability for this species in the Red Sea and Brazilian coast, especially in southeastern region, where the new records were made. Due to the number of collected individuals, it is hard to affirm that this taxon presents a self-sustaining population in Brazilian waters, but the several registers, in different locations and different life stages point to an establishment of the species in this new region. This population status allied with a high environmental suitability is alarming and should motivate new studies concerning T. vagina in Brazilian waters.     

Trait‐based life strategies,ecological niches,and niche overlap in the nekton of the data‐poor Mediterranean Sea

Biological traits can determine species ecological niches and define species responses to environmental variation. Species have a specific functional position in the biological community, resulting in interactions like interspecific competition. In this study, we used biological traits in order to define the life strategies of 205 nektonic species of the Mediterranean Sea. Furthermore, traits related to resource use were analyzed to determine the level of trait and niche overlap and their relationship to life strategies. Focusing on habitats of importance (Posidonia beds, coralligène formations, and lagoons), we investigated strategies and niches of the species present there. Finally, we examined the life strategy of Lessepsian species and investigated the niche overlap between them and indigenous species. Archetypal analysis indicated the existence of three life histories corresponding to strategies already documented for fish (equilibrium, periodic, and opportunistic), with some species also placed in intermediate positions. Niche overlap was evaluated by multiple correspondence analysis and the generation of a single distance metric between all species pairs. This identified species occupying relatively empty (underexploited) ecological niches, like the Lessepsian species Siganus luridus and S. rivulatus, a finding that can also be associated with their establishment in the Mediterranean. Most Lessepsian species were associated with the opportunistic life history strategy, again an important aspect related to their establishment. Also, we documented that most species occurring in important habitats have a relatively high overlap of niches. No significant differences were found in the life strategies across Mediterranean habitats; however, variation in niche overlap and traits related to habitat use was detected. The findings can be useful to determine theoretical competition between species and to identify empty ecological niches. Fisheries science can also benefit from comprehending the dynamics of competing stocks or predict the responses of data‐poor stocks to anthropogenic stressors from known examples of species with shared life strategies.     

Marine molluscs from the Iztuzu beach near Dalyan (Mediterranean coast of Turkey)

Abstract

A total of l84 species of marine molluscs has been identified from shells collected on the beach of Iztuzu near Dalyan (Mu?la province, Turkey). Of these, 16 species of gastropod and 4 species of bivalvia are new For the extreme Eastern Mediterranean. Only three species of Lessepsian migrants (from the Red Sea via the Suez canal) have been found.     

The largescaled terapon Terapon theraps: a new Indo‐Pacific fish in the Mediterranean Sea

A specimen of the largescaled terapon Terapon theraps was recorded for the first time from the Mediterranean Sea. The finding of T. theraps in the northernmost part of the Mediterranean Sea raises the number of Lessepsian migrant species to at least 68 and in the Adriatic Sea to at least 11.     

Genetics of the early stages of invasion of the Lessepsian rabbitfish Siganus luridus

Information on the initial stages of dispersal and settlement are of great interest in understanding the dynamics of biological invasions and in designing management responses. A newly settled population of the Lessepsian rabbitfish migrant Siganus luridus, that arrived in Linosa Island (Sicily Strait) in 2000, offered a unique opportunity to examine the genetic variability of the early phase of invasion and the starting point to test genetic variation within and between colonist and source populations.Demographics and dynamic aspects of S. luridus in the Mediterranean were evaluated by using phylogeographic and demographic (coalescent) methods based on DNA sequences of the mitochondrial control region. Sequences from 95 S. luridus, 25 Siganus rivulatus, and one of Siganus (Lo) vulpinus and S. doliatus were used. Samples were collected in one locality in the Red Sea (Eilat) and three localities in the Mediterranean (Israel, Greece and Linosa, Italy). Data showed (for the first time in a Lessepsian migrant) a lowering of the genetic diversity of the invading population (Mediterranean) (haplotype diversity 0.879, nucleotide diversity 0.592) compared to the parental one (Red Sea) (haplotype diversity 0.978, nucleotide diversity 0.958).Within the Mediterranean populations, there was no pattern of regional separation and mitochondrial diversity appeared to be preserved during the Linosa colonization, with no traces of founder events. Such evidence agrees with the idea that Lessepsian migration involves many individuals from its earliest stages.