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.     

Seagrasses in the Zeit Bay area and at Ras Ghârib (Egyptian Red Sea coast)

The distribution and ecology of seagrasses along parts of the Egyptian Red Sea coasts have been investigated. Five of the 10 species ever reported from the Red Sea have been found in this area, i.e. Halodule uninervis (Forssk.) Aschers., Thalassodendron ciliatum (Forssk.) den Hartog, Halophila stipulacea (Forssk.) Aschers., Halophila ovalis (R. Br.) Hook. f. and Halophila ovata Gaud. H. ovata is recorded here for the first time for the west coast of the Red Sea. The finding of Halophila decipiens Ostenfeld represents a new record to the Red Sea; this species has been found at a depth of 30 m, near Ras Ghârib in the Gulf of Suez.     

The distribution and expansion of the invasive seagrass Halophila stipulacea in Dominica, West Indies, with a preliminary report from St. Lucia

The seagrass Halophila stipulacea Forsskål, native to the Red Sea, is an invasive species in the Mediterranean that was recently observed offshore Grenada, in the Caribbean. Here, we document the presence of this seagrass in Dominica and St. Lucia, demonstrating it has spread across part of the eastern Caribbean. H. stipulacea in Dominica was present in seven locations along the west coast covering more than 22.9 ha of the benthos, at depths from 2 to 18 m. Populations were concentrated in or adjacent to bays frequented by recreational or commercial boats, likely vectors for the introduction. Morphological features varied from bed to bed, with depth being the predominant driving factor. H. stipulacea had a rapid mean lateral bed expansion rate of 0.5 cm d⁻¹, with a maximum rate of >6 cm d⁻¹. H. stipulacea patches often occurred exclusive of the otherwise dominant seagrasses of the Caribbean. The potential for the expansion of H. stipulacea, combined with its tolerance for a wide spectrum of environmental conditions, positions it as a potential threat to local and regional biodiversity.     

Productivity and respiration of three seagrass species from the gulf of Aqaba (Jordan) and some related factors

Net productivity and rates of respiration of the seagrasses Halodule uninervis (Forsk.) Aschers., Halophila ovalis (R. Brown) Hooker fil. and Halophila stipulacea (Forsk.) Aschers., from Aqaba (Jordan) are presented in this paper. The effect of some physical and chemical parameters on the productivity of H. stipulacea is discussed in some detail. Of the three species, H. stipulacea is considered a shade plant since it is photoinhibited at light intensities near 100 Wm^?2 and has a productivity/respiration ratio of 1.8 at 30 m depth.     

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.     

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     

New insights into DNA barcoding of seagrasses

Taxonomists find some plant genera challenging because of the few morphological differences or unclear characters among closely related species, which leads to the misidentification of taxa. DNA barcoding is an approach to identify species by using short orthologous DNA sequences, known as ‘DNA barcodes’. Concatenated rbcL and matK sequences are considered DNA barcodes for seagrasses. However, these markers are not applicable to all members of seagrasses at the species level, especially within the genus Halophila. Our previous studies indicated that the internal transcribed spacer (ITS) showed higher species resolution than the concatenated rbcL and matK sequences in the case of Halophila ovalis and closely related species. In this study, 26 ITS, two rbcL and two matK consensus sequences from 18 seagrass taxa belonging to four families collected in India, Vietnam, Germany, Croatia and Egypt were processed. Molecular ITS analysis resolved five clades. The results also indicate that the Cymodoceaceae family might be a non-monophyletic group. In conclusion, ITS could be applied as a DNA barcode for seagrasses instead of the rbcL/matK system previously proposed.     

Untersuchungen zur Zoogeographie der Euphausiaceen (Crustacea) des Roten Meeres

A joint programme on “Biota of the Red Sea and the Eastern Mediterranean” has been carried out by the Hebrew University of Jerusalem and the Smithsonian Institution of Washington between 1967 and 1972 in order to study the role played by the Suez Canal as a route for faunal exchanges between the Red Sea and Eastern Mediterranean. The euphausiids of this material have been examined and additional samples of zooplankton have been taken in the Gulf of Elat (Aqaba) in 1973 in order to study the vertical distribution of the euphausiids. Four species have been found to occur in the Northern Red Sea and the Gulf of Elat:Euphausia diomedeae, E. sanzoi, Stylocheiron affine andS. abbreviatum. Seven species were found in the Eastern Mediterranean of whichEuphausia brevis is the dominant species. The present studies revealed that an exchange of Euphausiacea through the Suez Canal does not take place. The species of the Red Sea belong to the Indopacific fauna. Despite the particular historic-geographical and ecological situation of the Red Sea, no endemisms or subspecies have evolved there within the Euphausiacea.     

The Japanese threadfin bream Nemipterus japonicus, a new Indo‐Pacific fish in the Mediterranean Sea

A specimen of the Japanese threadfin bream Nemipterus japonicus was recorded for the first time from the Mediterranean Sea. The presence of this Indo‐Pacific fish in the Mediterranean is evidently due to migration from the Red Sea via the Suez Canal.     

The Arabian scad Trachurus indicus, a new Indo-Pacific species in the Mediterranean Sea

The Arabian scad Trachurus indicus is recorded for the first time from the Mediterranean Sea (Iskenderun Bay, Turkey). The presence of this Indo-Pacific fish in the Mediterranean Sea is probably because of migration from the Red Sea via the Suez Canal.     

Chromosome numbers of the European seagrasses

The chromosome numbers of the five European seagrasses have been determined in material from several sites along the coasts of the Atlantic Ocean, the North Sea and the Mediterranean:Zostera marina L., 2n = 12;Z. noltii Hornem., 2n = 12;Posidonia oceanica (L.)Delile, 2n = 20;Cymodocea nodosa (Ucria)Aschers., 2n = 14, 2n = 28;Halophila stipulacea (Forsk.)Aschers., 2n = 18. The difference in chromosome morphology betweenZ. marina andZ. noltii supports the division of the genus into two subgenera.     

Lack of a genetic bottleneck in a recent Lessepsian bioinvader, the blue-barred parrotfish, Scarus ghobban

The present study investigates the genetic diversity of Scarus ghobban, a recently introduced parrotfish in the Mediterranean Sea via the Suez Canal. Two mitochondrial and one nuclear DNA regions were sequenced and phylogenetic relationships investigated, from samples collected from Lebanon and across its natural range. Scarus ghobban clustered in two major clades, Pacific Ocean and Indian Ocean, indicating strong population structure, or cryptic speciation. Expectedly, Mediterranean samples clustered with Indian Ocean-Red Sea individuals. However, unlike other recent Lessepsian invaders, S. ghobban displayed high genetic diversity. These results underscore that genetic diversity is a poor predictor of success of an invasive species.     

First confirmed record of Lagocephalus sceleratus(Gmelin, 1789) in the Mediterranean Sea

One specimen of the Indo‐Pacific silverstripe blaasop Lagocephalus sceleratus (Gmelin, 1789)(Tetraodontidae) is recorded from the Aegean coast of Turkey and is confirmed for the Mediterranean. Dispersal of the species to the Mediterranean is due to migration from the Red Sea via the Suez Canal.     

In situ nitrogen fixation associated with seagrasses in the Gulf of Elat (Red Sea)

In situ nitrogen fixation associated with the seagrass Halophila stipulacea, at the northern Gulf of Elat (Red Sea), is eight to ten times higher than that of nearby plant-free areas. A daily cycle of nitrogen fixation is evident, with rates during the day being seven times greater than during the night. Removal of seagrass leaves only from a patch within a seagrass bed gradually decreases nitrogen fixation activity, reaching the rates of plant-free areas after ten hours. A method devised for the in situ measurement of nitrogen fixation rates using belljars is described in detail. Nitrogen fixation rates in situ are higher than in the laboratory and lack the lag period typical to laboratory measurements. In laboratory experiments using intact plant samples, glucose enhances nitrogen fixation rates both in light and dark. Photosystem II inhibitor (3-3,4-dichloro-phenyl-1,1-dimethylurea) doubles nitrogen fixation rates in light. Both field and laboratory results indicate that light is essential for nitrogen fixation activity in the H. stipulacea bed possibly through its effect on cyanobacterial population that occupy the aerobic niches of the phyllosphere and on photosynthetic Rhodospirillacean bacteria that inhabit the anaerobic ones. Nitrogen fixation rates evident in H. stipulacea beds in situ account for a considerable portion of the biomass production by the seagrasses. The dependence of high nitrogenase activity by the diazotrophs on the presence of the seagrasses indicates the great importance of the seagrass community to the nitrogen cycle in its highly oligotrophic surroundings of the Gulf of Elat.     

Carbon Metabolism in Seagrasses: III. ACTIVITIES OF CARBON-FIXING ENZYMES IN RELATION TO INTERNAL SALT CONCENTRATIONS

The internal salt content and distribution in photosynthetictissues as well as the effect of NaCl on photosynthetic carbonfixation enzymes was investigated in two seagrass species fromthe Red Sea. Concentrations of both Na⁺ and Cl^– were lower in the chloroplast-richepidermis than in underlying cell layers in Halophila stipulacea.In Halodule uninervis, the concentration of Na⁺ was lower inthe epidermis than in the underlying cells, while K⁺ was evenlydistributed between cell layers. The epidermal concentrationsof Na+ were estimated to be 0.17 and 0.10 M for Halophila stipulaceaand Halodule uninervis, respectively, which were about to the average leaf concentrations. Epidermal Cl^– concentrationof Halophila stipulacea was estimated to be 0.08 M, a valueonly about of the overall leaf concentration. Phosphoenolpyruvate carboxylase (PEPcase) extracted from leavesof these seagrasses showed increased activity at 0.05–0.3M NaCl in vitro. Ribulose-l, 5-bisphosphate carboxylase (RuBPcase)activity, on the other hand, was inhibited by NaCl at all testedconcentrations. At epidermal NaCl concentrations, PEPcase activitywas thus stimulated while RuBPcase was inhibited. The reducedRuBPcase activity at such concentrations compared to salt-freeconditions was still sufficient to account for observed photosyntheticrates. We conclude that these seagrasses have adapted to a saline environmentboth by maintaining relatively low ion concentrations in theepidermis where photosynthesis occurs and by having carbon-fixingenzymes capable of functioning in the presence of salt.     

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.     

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.     

On the occurrence of Hippocampus fuscus in the eastern Mediterranean

The western Indian Ocean seahorse Hippocampus fuscus is recorded for the first time in the Mediterranean. Its presence there is probably due to migration from the Red Sea via the Suez Canal.     

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.