Opisthobranch molluscs of the Sudanese Red Sea

Eighteen opisthobranch species are described and illustrated from the Sudanese Red Sea, including seven new species, one new subspecies and three species for which this is the first Red Sea record. Eleven bullomorphs, four pleurobranchomorphs, four sacoglossans and four phyllidiid mudibranchs are now known from the Red Sea.
There appears to be a high percentage of endemic species in the Red Sea. This is what one might expect, in view of the conditions of near isolation, low temperatures and high salinity that prevailed in this sea 13000 23000 years ago.
Of the groups discussed in this paper, one Red Sea species has invaded the Mediterranean Sea, and one Mediterranean species has invaded the Red Sea, since the opening of the Suez Canal in 1867.
Many Red Sea species are very colourful, in comparison to Mediterranean ones. This could perhaps be because Red Sea waters are cleaner, thereby favouring the evolution of greater precision in colour markings.     

Habitat utilization by an invasive herbivorous fish (<Emphasis Type="Italic">Siganus rivulatus</Emphasis>) in its native and invaded range

Movement is essential for understanding the distribution and abundance of animals. While it has been suggested that invasion success can be facilitated by species’ ability to adapt to novel environments, direct comparisons of movement patterns between native and invaded ranges of animals in their natural habitat are rare. The rivulated rabbitfish Siganus rivulatus was introduced from the Red Sea into the Mediterranean, where it is now found in extremely high abundances, and has overgrazed the coastal marine ecosystem in many locations. Through a continuous acoustic tracking system, we found that the movement of S. rivulatus individuals at a Mediterranean site differed substantially from those at a Red Sea site, with individuals in the Mediterranean having larger overall home ranges and lower site fidelity. However, no variation between sites was found in daily home range sizes. Results show that at the Mediterranean site S. rivulatus individuals have a larger spatial footprint, which may contribute to their impact and ability to expand their distribution. This study demonstrates a potential shift in individual movement of a marine invasive species between its native and invaded range, and highlights the role of movement in understanding biological invasions.     

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.     

Ecological traits and environmental affinity explain Red Sea fish introduction into the Mediterranean

Alien species are considered one of the prime threats to biodiversity, driving major changes in ecosystem structure and function. Identifying the traits associated with alien introduction has been largely restricted to comparing indigenous and alien species or comparing alien species that differ in abundance or impact. However, a more complete understanding may emerge when the entire pool of potential alien species is used as a control, information that is rarely available. In the eastern Mediterranean, the marine environment is undergoing an unparalleled species composition transformation, as a flood of aliens have entered from the Red Sea following the opening of the Suez Canal in 1869. In this study, we compile data on species traits, geographical distribution, and environmental affinity of the entire pool of reef‐associated fish species in the Red Sea and more generally across the Indo‐Pacific. We use this extensive data to identify the prime characteristics separating Red Sea species that have become alien in the Mediterranean from those that have not. We find that alien species occupy a larger range of environments in their native ranges, explaining their ability to colonize the seasonal Mediterranean. Red Sea species that naturally experience high maximum temperatures in their native range have a high probability of becoming alien. Thus, contrary to predictions of an accelerating number of aliens following increased water temperatures, hotter summers in this region may prevent the establishment of many alien species. We further find that ecological trait diversity of alien species is substantially more evenly spaced and more divergent than random samples from the pool of Red Sea species, pointing at additional processes, such as competition, promoting ecological diversity among alien species. We use these results to provide a first quantitative ranking of the potential of Red Sea species to become established in the eastern Mediterranean.     

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.     

Spirorbinae (Polychaeta:Serpulidae) from the Red Sea, including descriptions of a new genus and four new species

A new genus, a new subgenus, four new species and four previously described species are recorded from the Red Sea. The new genus Anomalorbis has four and a half thoracic segments. Re-discovery of the sinistral variety of Janua (Dexiospira) steueri confirms Sterzinger's observation that this species may coil in either direction. Janua (Fauveldora) anticorrugata sp. nov. is also sinistral and a shallow water species. It therefore appears that the Red Sea may have been the centre of origin for sinistral mutants of the Janua complex. Other of the new forms live in fairly deep water. The assemblage encountered shows closest affinity with the Indo-Pacific, but suggests a high degree of endemism, which may be related to the high temperatures prevailing at all depths in the Red Sea.     

Apogon smithi (Kotthaus, 1970) (Teleostei: Apogonidae), a Red Sea cardinalfish colonizing the Mediterranean Sea

Smith's cardinalfish Apogon smithi is recorded for the first time in the Mediterranean Sea. Six specimens were caught by commercial trawler north of Jaffa, Israel, at depths of 40 m and the relatively high number of specimens suggests that A. smithi has established a sustainable population. This is the third Indo-Pacific apogonid species documented in the Mediterranean Sea, and is assumed to be a migrant originating from the Red Sea.     

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.     

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.     

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.     

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.     

Red to Mediterranean Sea bioinvasion: natural drift through the Suez Canal,or anthropogenic transport?

The biota of the eastern basin of the Mediterranean Sea has experienced dramatic changes in the last decades, in part as a result of the massive invasion of Red Sea species. The mechanism generally hypothesized for the 'Red-to-Med' invasion is that of natural dispersal through the Suez Canal. To date, however, this hypothesis has not been tested. This study examines the mode of invasion, using as a model the mussel Brachidontes pharaonis, an acclaimed 'Lessepsian migrant' that thrives along the eastern Mediterranean coast. Our findings reveal two distinct lineages of haplotypes, and five possible explanations are discussed for this observation. We show that the genetic exchange among the Mediterranean, Gulf of Suez and the northern Red Sea is sufficiently large to counteract the build up of sequential genetic structure. Nevertheless, these basins are rich in unique haplotypes of unknown origin. We propose that it is historic secondary contact, an ongoing anthropogenic transport or both processes, that participate in driving the population dynamics of B. pharaonis in the Mediterranean and northern Red Sea.     

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.     

Why is the mediterranean more readily colonized than the Red Sea,by organisms using the Suez Canal as a passageway?

Summary Since the opening of the Suez Canal, more than 120 Red Sea species colonized the eastern Mediterranean, whereas less than 10 Mediterranean species colonized the Red Sea. For most of the species involved in this colonization, the mode of dispersal from the source to the colonized area is through free-drifting propagules. In order to examine whether the current regime of the Suez Canal may be involved in this assymetry in colonization, a mathematical hydraulic model that forecasts the direction and velocity of water currents through the year, along the length of the Canal, was utilized. The movements of free-floating propagules that occur at either entrance of the Canal, was simulated on a computer, and it was found that the completion of a Mediterranean-bound passage of Red Sea propagules is far faster and much more likely than a completion of a Red Sea-bound passage of Mediterranean propagules.Paper No. 7 in the series Colonization of the Eastern Mediterranean by Red Sea species immigrating through the Suez Canal     

Mitochondrial DNA variation in a species with two mitochondrial genomes: the case of Mytilus galloprovincialis from the Atlantic, the Mediterranean and the Black Sea

We have examined mitochondrial DNA (mtDNA) variation in samples of the mussel Mytilus galloprovincialis from the Black Sea, the Mediterranean and the Spanish Atlantic coast by scoring for presence or absence of cleavage at 20 restriction sites of a fragment of the COIII gene and at four restriction sites of the 16S RNA gene. This species contains two types of mtDNA genomes, one that is transmitted maternally (the F type) and one that is transmitted paternally (the M type). The M genome evolves at a higher rate than the F genome. Normally, females are homoplasmic for an F type and males are heteroplasmic for an F and an M type. Occasionally molecules from the F lineage invade the paternal transmission route, resulting in males that carry two F-type mtDNA genomes. These features of the mussel mtDNA system give rise to a new set of questions when using mtDNA variation in population studies and phylogeny. We show here that the two mtDNA types provide different information with regard to amounts of variation and genetic distances among populations. The F genome exhibits higher degrees of diversity within populations, while the M genome produces higher degrees of differentiation among populations. There is a strong differentiation between the Atlantic and the Black Sea. The Mediterranean samples have intermediate haplotype frequencies, yet are much closer to the Black Sea than to the Atlantic. We conclude that in this species gene flow among the three Seas is restricted and not enough to erase the combined effect of mutation and random drift. In one sample, that from the Black Sea, the majority of males did not contain an M mtDNA type. This suggests that a molecule of the maternal lineage has recently invaded the paternal route and has increased its frequency in the population to the point that the present pool of paternally transmitted mtDNA molecules is highly heterogeneous and cannot be used to read the population's history. This liability of the paternal route means that in species with doubly uniparental inheritance, the maternal lineage provides more reliable information for population and phylogenetic studies.     

Parhyale plumicornis (Crustacea: Amphipoda: Hyalidae): is this an anti-lessepsian Mediterranean species? Morphological remarks,molecular markers and ecological notes as tools for future records

Hyalid amphipods living in coastal marine habitats are frequently included in ecological studies. The systematics of this taxon has been subject to profound changes, with an emphasis on the North Pacific fauna. Since a proper species delimitation is a prerequisite in taxonomic and ecological studies, Parhyale plumicornis (Heller, 1866) has been herein re-described, showing the criticisms and mismatches of various characters, which were previously used in dichotomous keys. This species was collected for the first time off the western coast of Sicily Island (Italy: central Mediterranean Sea). The male is peculiar, due to the second antennae heavily setose posteriorly and bearing long tufts of plumose ventral setae. In this paper, the species will be illustrated, and morphological polymorphism, molecular tags and ecological features will be reported. The species does not appear to be frequent in the Mediterranean Sea but it is important that marine biologists identify it accurately. A recent record, possibly ascribable to Parhyale plumicornis in the Red Sea, could indicate that this species is the first anti-lessepsian amphipod, which has migrated from the Mediterranean Sea towards the Red Sea.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:770BEDA1-3E06-464F-9D34-8AFE43592FCA     

Population genetic structure and connectivity of the harmful dinoflagellate Alexandrium minutum in the Mediterranean Sea

The toxin-producing microbial species Alexandrium minutum has a wide distribution in the Mediterranean Sea and causes high biomass blooms with consequences on the environment, human health and coastal-related economic activities. Comprehension of algal genetic differences and associated connectivity is fundamental to understand the geographical scale of adaptation and dispersal pathways of harmful microalgal species. In the present study, we combine A. minutum population genetic analyses based on microsatellites with indirect connectivity (C(i)) estimations derived from a general circulation model of the Mediterranean sea. Our results show that four major clusters of genetically homogeneous groups can be identified, loosely corresponding to four regional seas: Adriatic, Ionian, Tyrrhenian and Catalan. Each of the four clusters included a small fraction of mixed and allochthonous genotypes from other Mediterranean areas, but the assignment to one of the four clusters was sufficiently robust as proved by the high ancestry coefficient values displayed by most of the individuals (>84%). The population structure of A. minutum on this scale can be explained by microalgal dispersion following the main regional circulation patterns over successive generations. We hypothesize that limited connectivity among the A. minutum populations results in low gene flow but not in the erosion of variability within the population, as indicated by the high gene diversity values. This study represents a first and new integrated approach, combining both genetic and numerical methods, to characterize and interpret the population structure of a toxic microalgal species. This approach of characterizing genetic population structure and connectivity at a regional scale holds promise for the control and management of the harmful algal bloom events in the Mediterranean Sea.