Aim The Red Sea is characterised by a unique fauna and historical periods of desiccation, hypersalinity and intermittent isolation. The origin and contemporary composition of reef-associated taxa in this region can illuminate biogeographical principles about vicariance and the establishment (or local extirpation) of existing species. Here we aim to: (1) outline the distribution of shallow water fauna between the Red Sea and adjacent regions, (2) explore mechanisms for maintaining these distributions and (3) propose hypotheses to test these mechanisms.Location Red Sea, Gulf of Aden, Arabian Sea, Arabian Gulf and Indian Ocean.Methods Updated checklists for scleractinian corals, fishes and non-coral invertebrates were used to determine species richness in the Red Sea and the rest of the Arabian Peninsula and assess levels of endemism. Fine-scale diversity and abundance of reef fishes within the Red Sea were explored using ecological survey data.Results Within the Red Sea, we recorded 346 zooxanthellate and azooxanthellate scleractinian coral species of which 19 are endemic (5.5%). Currently 635 species of polychaetes, 211 echinoderms and 79 ascidians have been documented, with endemism rates of 12.6%, 8.1% and 16.5% respectively. A preliminary compilation of 231 species of crustaceans and 137 species of molluscs include 10.0% and 6.6% endemism respectively. We documented 1071 shallow fish species, with 12.9% endemic in the entire Red Sea and 14.1% endemic in the Red Sea and Gulf of Aden. Based on ecological survey data of endemic fishes, there were no major changes in species richness or abundance across 1100 km of Saudi Arabian coastline.Main conclusions The Red Sea biota appears resilient to major environmental fluctuations and is characterized by high rates of endemism with variable degrees of incursion into the Gulf of Aden. The nearby Omani and Arabian Gulfs also have variable environments and high levels of endemism, but these are not consistently distinct across taxa. The presence of physical barriers does not appear to explain species distributions, which are more likely determined by ecological plasticity and genetic diversity.
Trochoidea are a large superfamily of morphologically and ecologically diverse marine gastropods. We present here an appraisal of the composition and relationships among trochoidean families based on molecular data, with an especial focus on the family Trochidae. Bayesian analyses of sequences from three genes (18S rRNA, 28S rRNA and COI) including data from 162 vetigastropod species show that the gastropod family Trochidae (sensu Hickman & McLean (1990), Natural History Museum Los Angeles County Science Series, 35, 1–169) is not monophyletic. Recognition of Chilodontidae, Solariellidae and Calliostomatidae at the family level is supported. Our new, more limited, definition of Trochidae includes the subfamilies Stomatellinae, Lirulariinae and Umboniinae and redefined Trochinae, Cantharidinae and Monodontinae. Halistylinae are provisionally retained in the Trochidae based on previous morphological studies. As redefined, Trochidae are a predominantly shallow‐water radiation in the tropics and subtropics. Some subfamilies and genera previously included in Trochidae have been moved to an enlarged family Turbinidae. The family Turbinidae has been redefined to include Turbininae, Skeneinae, Margaritinae, Tegulinae, Prisogasterinae and most surprisingly the commercially important genus Tectus Montfort, 1810. The new definition of Turbinidae means that the family includes both predominantly shallow and deep‐water clades as well as genera that are distributed across the globe from the poles to the tropics. A greater range of habitat is now seen in Turbinidae than in Trochidae. The redefined Trochidae and Turbinidae, together with Solariellidae, Calliostomatidae and Liotiidae, make up the superfamily Trochoidea. Phasianellidae and Colloniidae are recognized as belonging in a new superfamily, Phasianelloidea, and Angaria Röding, 1798 is recognized as belonging in a new superfamily, Angarioidea. Placement of Areneidae into a superfamily awaits further work.
The phylum Mollusca is highly speciose, and is the largest phylum in the marine realm. The great majority of molluscs are shelled, including nearly all bivalves, most gastropods and some cephalopods. The fabulous and diverse colours and patterns of molluscan shells are widely recognised and have been appreciated for hundreds of years by collectors and scientists alike. They serve taxonomists as characters that can be used to recognise and distinguish species, however their function for the animal is sometimes less clear and has been the focus of many ecological and evolutionary studies. Despite these studies, almost nothing is known about the evolution of colour in molluscan shells. This review summarises for the first time major findings of disparate studies relevant to the evolution of shell colour in Mollusca and discusses the importance of colour, including the effects of visual and non-visual selection, diet and abiotic factors. I also summarise the evidence for the heritability of shell colour in some taxa and recent efforts to understand the molecular mechanisms underpinning synthesis of shell colours. I describe some of the main shell pigments found in Mollusca (carotenoids, melanin and tetrapyrroles, including porphyrins and bile pigments), and their durability in the fossil record. Finally I suggest that pigments appear to be distributed in a phylogenetically relevant manner and that the synthesis of colour is likely to be energetically costly.
A new molecular phylogeny is presented for the highly diverse, bivalve molluscan subclass Heterodonta. The study, the most comprehensive for heterodonts to date, used new sequences of 18S and 28S rRNA genes for 103 species from 49 family groups with species of Palaeoheterodonta (Trigoniidae, Margaritiferidae and Unionidae) as outgroups. Results confirm previous analyses that the Carditidae/Astartidae/Crassatellidae clade is basal to all other heterodonts including Anomalodesmata (often classified as a separate subclass or order). Thyasiroidea occupy a near basal position between the Crassatelloidea and Anomalodesmata. Lucinidae form a well‐supported monophyletic group distinct from Thyasiridae and Ungulinidae. The Solenoidea and Hiatelloidea link as sister groups distant from the Tellinoidea and Myoidea, respectively, where they had been previously associated. The position of the Gastrochaenidae is unstable but does not group with myoidean taxa. Species of four families of Galeommatoidea form a clade that also includes Sportellidae of the Cyamioidea. The Cardioidea and Tellinoidea form highly supported, long branched, individual clades but group as sister taxa. A major clade including Veneroidea, Mactroidea, Myoidea and other families is given the unranked name Neoheterodontei. There is no support for a separate order Myoida (Myoidea and Pholadoidea). Dreissenidae group within the clade including Myidae, Corbulidae, Pholadidae and Teredinidae. The Corbiculoidea is confirmed as polyphyletic with the Sphaeriidae and Corbiculidae forming separate clades within the Neoheterodontei; Corbiculidae grouping with the Glauconomidae. Hemidonacidae are unrelated to the Cardiidae, as previously proposed, but nest within the Neoheterodontei. The Gaimardiidae group near to the Ungulinidae and not with Cyamioidea where most recently classified. The family Ungulinidae, previously classified in the Lucinoidea, forms a well‐supported clade within the Neoheterodontei and is elevated to superfamily rank — Ungulinoidea. The monophyletic status of Glossoidea, Arcticoidea and Veneroidea is unconfirmed. A brief review of the fossil record of the heterodonts indicates that the basal clades of Crassatelloidea, Anomalodesmata and Lucinoidea diverged very early in the Lower Palaeozoic. Other groups such as the Hiatelloidea, Solenoidea, Gastrochaenidae probably were of late Palaeozoic origins. The Cardioidea and Tellinoidea originated in the Triassic while major groups of Neoheterodontei radiated in the Late Mesozoic. The phylogenetic position of the Thyasiroidea and Galeommatoidea suggests a longer fossil history than has so far been recognized.
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