Molecular phylogeny of Thoracotremata crabs (Decapoda, Brachyura): Toward adopting monophyletic superfamilies, invasion history into terrestrial habitats and multiple origins of symbiosis.

The Thoracotremata is a large and successful group of "true" crabs (Decapoda, Brachyura, Eubrachyura) with a great diversity of lifestyles and well-known intertidal representatives. The group represents the largest brachyuran radiation into terrestrial and semi-terrestrial environments and comprises multiple lineages of obligate symbiotic species. In consequence, they exhibit very diverse physiological and morphological adaptations. Our understanding of their evolution is, however, largely obscured by their confused classification. Here, we resolve interfamilial relationships of Thoracotremata, using 10 molecular markers and exemplars from all nominal families in order to reconstruct the pathways of lifestyle transition and to prepare a new taxonomy corresponding to phylogenetic relationships. The results confirm the polyphyly of three superfamilies as currently defined (Grapsoidea, Ocypodoidea and Pinnotheroidea). At the family level, Dotillidae, Macrophthalmidae, and Varunidae are not monophyletic. Ancestral state reconstruction analyses and divergent time estimations indicate that the common ancestor of thoracotremes already thrived in intertidal environments in the Late Cretaceous and terrestrialization became a major driver of thoracotreme diversification. Multiple semi-terrestrial and terrestrial lineages originated and radiated in the Early Eocene, coinciding with the global warming event at the Paleocene-Eocene Thermal Maximum (PETM). Secondary invasions into subtidal regions and colonizations of freshwater habitats occurred independently through multiple semi-terrestrial and terrestrial lineages. Obligate symbiosis between thoracotremes and other marine macro-invertebrates evolved at least twice. On the basis of the current molecular phylogenetic hypothesis, it will be necessary in the future to revise and recognize seven monophyletic superfamilies and revisit the morphological character states which define them.

The Identity of Homoioplax haswelli (Miers, 1884) (Crustacea: Decapoda: Brachyura).

The brachyuran crab Pseudorhombila haswelli Miers, 1884, described on the basis of two juveniles from the Arafura Sea, is a poorly known species of uncertain systematic position. It was made the type and only species of Homoioplax Rathbun, 1914, and assigned to the Prionoplacinae (Goneplacidae). Subsequent revisions of the goneplacids showed Prionoplacinae to be a junior synonym of Eucratopsinae Stimpson, 1871, as a member of the Panopeidae Ortmann, 1893, but no one has re-evaluated the position of Homoioplax. As a result, Homoioplax has remained in the Panopeidae as the only Indo-West Pacific representative of the family. This study assesses the systematic status of Homoioplax haswelli based on a re-examination of the type and other material of the species. Homoioplax haswelli is here attributed to Carcinoplax H. Milne Edwards, 1852 (Goneplacidae), justifying the recognition of the species described by Miers as C. haswelli. Moreover, Carcinoplax haswelli is shown to be a senior synonym of C. sinica Chen, 1984, described from the South China Sea. Therefore, C. haswelli is now known to range from southern Taiwan and the Philippines, through the South China Sea, northern Vietnam, the eastern waters of Singapore to the Madura Straits in eastern Java and Arafura Sea, northern Australia, at 25-187 m, but usually less than 100 m depth.

Carcinoplax fasciata, a new species of deep-water goneplacid crab from southwestern India (Crustacea: Decapoda: Brachyura: Goneplacoidea).

A new species of Carcinoplax H. Milne Edwards, 1852 (family Goneplacidae) is described from southern India. Carcinoplax fasciata n. sp. is closest to C. specularis Rathbun, 1914, but can be distinguished by its different coloration in life as well as structures of the carapace surface, anterolateral armature, supraorbital margin, chela and male first gonopod.

On the identity of Cancer urania Herbst, 1801 (Crustacea: Decapoda: Brachyura: Leucosiidae).

Cancer urania Herbst, 1801, is the type species of the leucosiid genus Coleusia Galil, 2006. Its identity has been a subject of confusion due to various taxonomical and nomenclatural issues. We redescribe the species, discuss its complex history and taxonomy, and a female syntype is designated as the lectotype of the species to clarify any lingering ambiguities concerning the type material of Cancer urania Herbst, 1801.

Discovery of a new genus and new species of Indo-West Pacific pilumnoidid crab from a semisubmersible oil platform (Crustacea: Brachyura: Pseudozioidea).

A new genus and new species of pseudozioid crab of the family Pilumnoididae is described from a fouling community on a semisubmersible oil platform in Singapore that had been operating in the Timor Sea and South China Sea. Setozius incertus gen. et sp. nov. superficially resembles species of Pilumnus (Pilumnidae, Pilumnoidea) but has male first and second gonopod structures characteristic of the Pseudozioidea. The form of the carapace, male anterior thoracic sternum and male abdomen indicates that it should be classified in the Pilumnoididae. Setozius is the first member of the family to be recorded from the Indo-West Pacific; all other known pilumnoidids occur in the Atlantic and eastern Pacific.

A new record of the giant freshwater prawn, Macrobrachium spinipes (Schenkel, 1902) (Crustacea: Decapoda: Palaemonidae) from Taiwan, with notes on its taxonomy.

The giant freshwater prawn Macrobrachium spinipes (Schenkel, 1902) is recorded from Taiwan for the first time and extends the distribution of the species to north of the Tropic of Cancer. The Taiwanese specimens differ slightly from material from Indonesian Papua in the density of the spination of the adult second pereipods, the relative length of the ridge of the posterior submedian plate of thoracite sternite 4, and the color of the carapace, abdomen and pleural condyles.

Semisubmersible oil platforms: understudied and potentially major vectors of biofouling-mediated invasions.

Biofouling has long been recognised as a major pathway for the introduction of non-indigenous species. This study records the decapods and stomatopod crustaceans fouling a semisubmersible oil platform dry docked for hull cleaning in Jurong Port, Singapore. Of the 25 species of decapods identified, 13 were non-indigenous and represent new records to Singapore waters. Of these, the crabs Glabropilumnus seminudus and Carupa tenuipes are known to be invasive in other parts of the world. The stomatopod, Gonodactylaceus randalli, is the first mantis shrimp recorded in a biofouling community. The richness and diversity of this fouling community, consisting of many vagile species, highlights the difference between platforms and ships. With the expansion of maritime oil and gas exploration, the threat posed by an expanded fleet of semisubmersible oil platforms translocating non-indigenous fouling communities across biogeographical boundaries is very serious. Scientists, policy-makers, and stakeholders should turn their attention to this growing problem.

Phylogenetics of the brachyuran crabs (Crustacea: Decapoda): the status of Podotremata based on small subunit nuclear ribosomal RNA.

The true crabs, the Brachyura, are generally divided into two major groups: Eubrachyura or 'advanced' crabs, and Podotremata or 'primitive' crabs. The status of Podotremata is one of the most controversial issues in brachyuran systematics. The podotreme crabs, best recognised by the possession of gonopores on the coxae of the pereopods, have variously been regarded as mono-, para- or polyphyletic, or even as non-brachyuran. For the first time, the phylogenetic positions of the podotreme crabs were studied by cladistic analysis of small subunit nuclear ribosomal RNA sequences. Eight of 10 podotreme families were represented along with representatives of 17 eubrachyuran families. Under both maximum parsimony and Bayesian Inference, Podotremata was found to be significantly paraphyletic, comprising three major clades: Dromiacea, Raninoida, and Cyclodorippoida. The most 'basal' is Dromiacea, followed by Raninoida and Cylodorippoida. Notably, Cyclodorippoida was identified as the sister group of the Eubrachyura. Previous hypotheses that the dromiid crab, Hypoconcha, is an anomuran were unsupported, though Dromiidae as presently composed could be paraphyletic. Topologies constrained for podotreme monophyly were found to be significantly worse (P < 0.04) than unconstrained topologies under Templeton and S-H tests. The clear pattern of podotreme paraphyly and robustness of topologies recovered indicates that Podotremata as a formal concept is untenable. Relationships among the eubrachyurans were generally equivocal, though results indicate the majoids or dorippoids were the least derived of the Eubrachyura. A new high level classification of the Brachyura is proposed.