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.     

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.     

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 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.     

Asterocherids (Copepoda; Siphonostomatoida) associated with invertebrates from California Reefs: Abrolhos (Brazil)

Four new asterocherid siphonostomatoid copepods are described, and several new geographical and host records are provided from California Reefs off the coast of Brazil. Acontiophorus monanchorae sp. nov. and Scottocheres youngi sp. nov. were both found associated with the sponge Monanchora sp. Asterocheres aplysinus sp. nov. and Asterocheres spongus sp. nov. were each associated with three genera of sponges. Asterocheres crenulatus previously recorded from the region, was associated with the sponge Xestospongia sp. Asterocheres paraboecki was recorded for the first time in the region. Asterocheres spinopaulus, previously reported from sponges, was found associated with bryozoans and cnidarians. Cletopontius titanus and Orecturus bahiensis were again recorded from the region and from sponges. Scottocheres laubieri, previously known from the Mediterranean Sea, was recorded for the first time in Brazilian waters, associated with the sponge Aplysina sp.     

First record of the blackfin coral goby,Paragobiodon lacunicolus (Kendall and Goldsborough, ), from the Red Sea

Three specimens of the blackfin coral goby, Paragobiodon lacunicolus, were collected in 2010 and 2011 at different locations of the eastern Red Sea along the coast of Saudi Arabia. Reexamination of 14 museum specimens from Eritrea also confirmed the presence of the species for the western Red Sea. This represents the first published report of P. lacunicolus from the Red Sea.     

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.     

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.     

Comparative genetics of scyphozoan species reveals the geological history and contemporary processes of the Mediterranean Sea

Jellyfish are useful genetic indicators for aquatic ecosystems as they have limited mobility and are highly exposed to the water column. By using comparative genomics and the molecular clock (timetree) of Rhizostoma pulmo, we revealed a divergence point between the East and West Mediterranean Sea (MS) populations that occurred 4.59 million years ago (mya). It is suggested that the two distinct ecological environments we know today were formed at this time. We propose that before this divergence, the highly mixed Atlantic and Mediterranean waters led to the wide dispersal of different species including R. pulmo. At 4.59 mya, the Western and Eastern MS were formed, indicating the possibility of a dramatic environmental event. For the first time, we find that for the jellyfish we examined, the division of the MS in east and west is not at the Straits of Sicily as generally thought, but significantly to the east. Using genomics of the Aurelia species, we examined contemporary anthropogenic impacts with a focus on migration of scyphozoa across the Suez Canal (Lessepsian migration). Aurelia sp. is among the few scyphozoa we find in both the MS and the Red Sea, but our DNA analysis revealed that the Red Sea Aurelia sp. did not migrate or mix with MS species. Phyllorhiza punctata results showed that this species was only recently introduced to the MS as a result of anthropogenic transportation activity, such as ballast water discharge, and revealed a migration vector from Australia to the MS. Our findings demonstrate that jellyfish genomes can be used as a phylogeographic molecular tool to trace past events across large temporal scales and reveal invasive species introduction due to human activity.     

Genomic signatures of rapid adaptive evolution in the bluespotted cornetfish,a Mediterranean Lessepsian invader

Biological invasions are increasingly creating ecological and economical problems both on land and in aquatic environments. For over a century, the Mediterranean Sea has steadily been invaded by Indian Ocean/Red Sea species (called Lessepsian invaders) via the Suez Canal, with a current estimate of ~450 species. The bluespotted cornetfish, Fistularia commersonii, considered a ‘Lessepsian sprinter’, entered the Mediterranean in 2000 and by 2007 had spread through the entire basin from Israel to Spain. The situation is unique and interesting both because of its unprecedented rapidity and by the fact that it took this species c. 130 years to immigrate into the Mediterranean. Using genome scans, with restriction site‐associated DNA (RAD) sequencing, we evaluated neutral and selected genomic regions for Mediterranean vs. Red Sea cornetfish individuals. We found that few fixed neutral changes were detectable among populations. However, almost half of the genes associated with the 47 outlier loci (potentially under selection) were related to disease resistance and osmoregulation. Due to the short time elapsed from the beginning of the invasion to our sampling, we interpret these changes as signatures of rapid adaptation that may be explained by several mechanisms including preadaptation and strong local selection. Such genomic regions are therefore good candidates to further study their role in invasion success.     

Molecular Phylogeny and Ecology of Textularia agglutinans d’Orbigny from the Mediterranean Coast of Israel: A Case of a Successful New Incumbent

Textularia agglutinans d’Orbigny is a non-symbiont bearing and comparatively large benthic foraminiferal species with a widespread distribution across all oceans. In recent years, its populations have considerably expanded along the Israeli Mediterranean coast of the eastern Levantine basin. Despite its exceptionally widespread occurrence, no molecular data have yet been obtained. This study provides the first ribosomal DNA sequences of T. agglutinans complemented with morphological and ecological characterization, which are based on material collected during environmental monitoring of the hard bottom habitats along the Israeli Mediterranean coast, and from the Gulf of Elat (northern Red Sea). Our phylogenetic analyses reveal that all specimens from both provinces belong to the same genetic population, regardless their morphological variability. These results indicate that modern population of T. agglutinans found on the Mediterranean coast of Israel is probably Lessepsian. Our study also reveals that T. agglutinans has an epiphytic life mode, which probably enabled its successful colonization of the hard bottom habitats, at the Mediterranean coast of Israel, which consist of a diverse community of macroalgae. Our study further indicates that the species does not tolerate high SST (> 35°C), which will probably prevent its future expansion in the easternmost Mediterranean in light of the expected rise in temperatures.     

Recent records of the Nile Soft-shelled Turtle,Tríonyx triunguis,and of the Euphrates Soft-shelled Turtle,Tríonyx euphratìcus,in the Middle East

Abstract

Thriving populations of the Spiny Oyster Spondylus spinosus Schreibers, 1793, an Indo-Pacific bivalve, are reported from off the Mediterranean coast of Israel. The taxonomy and nomenclature of this new Lessepsian migrant are briefly discussed. The polymorphism of the Mediterranean specimens is discussed. Their habitat is described, and the epibionts and predators in the Mediterranean are enumerated.     

A new parasitic copepod (Cyclopoida: Bomolochidae) from a ponyfish (Leiognathidae) caught in Egyptian Mediterranean waters,with a review of hosts and key to species of Nothobomolochus

A new bomolochid copepod belonging to the genus Nothobomolochus Vervoort, 1962 is described from a Red Sea fish species, a ponyfish of the family Leiognathidae that has become established in the Eastern Mediterranean. The new species, N. leiognathicola n. sp., is based on material obtained from the gill chamber of the Red Sea immigrant ponyfish Leiognathus klunzingeri (Steindachner), caught in Egyptian waters off the Alexandria coast at Abuqir. A second new species, N. monodi n. sp., is established to accommodate some material previously described as N. denticulatus (Bassett-Smith, 1898), from the host Hemiramphus far Forsskål. A review of host records reveals that Nothobomolochus species utilise hosts representing five different orders, but are most commonly found on beloniform, clupeiform and perciform fishes. A newly constructed key to the 37 valid species of Nothobomolochus is presented.     

On the record of the Lessepsian migrant Fistularia commersonii (Rüppell, 1835) from the Adriatic Sea

On 7 November and 15 December 2006, two specimens of the Lessepsian migrant Fistularia commersonni were caught in trammel nets off the coastal waters of Tricase Porto (southwestern Adriatic, Italy) and Sveti Andrija (southeastern Adriatic, Croatia), respectively. These represent the first records of this species in the Adriatic Sea.     

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.     

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.