Convergent adaptation of true crabs (Decapoda: Brachyura) to a gradient of terrestrial environments.

For much of terrestrial biodiversity, the evolutionary pathways of adaptation from marine ancestors are poorly understood, and have usually been viewed as a binary trait. True crabs, the decapod crustacean infraorder Brachyura, comprise over 7,600 species representing a striking diversity of morphology and ecology, including repeated adaptation to non-marine habitats. Here, we reconstruct the evolutionary history of Brachyura using new and published sequences of 10 genes for 344 tips spanning 88 of 109 brachyuran families. Using 36 newly vetted fossil calibrations, we infer that brachyurans most likely diverged in the Triassic, with family-level splits in the late Cretaceous and early Paleogene. By contrast, the root age is underestimated with automated sampling of 328 fossil occurrences explicitly incorporated into the tree prior, suggesting such models are a poor fit under heterogeneous fossil preservation. We apply recently defined trait-by-environment associations to classify a gradient of transitions from marine to terrestrial lifestyles. We estimate that crabs left the marine environment at least seven and up to 17 times convergently, and returned to the sea from non-marine environments at least twice. Although the most highly terrestrial- and many freshwater-adapted crabs are concentrated in Thoracotremata, Bayesian threshold models of ancestral state reconstruction fail to identify shifts to higher terrestrial grades due to the degree of underlying change required. Lineages throughout our tree inhabit intertidal and marginal marine environments, corroborating the inference that the early stages of terrestrial adaptation have a lower threshold to evolve. Our framework and extensive new fossil and natural history datasets will enable future comparisons of non-marine adaptation at the morphological and molecular level. Crabs provide an important window into the early processes of adaptation to novel environments, and different degrees of evolutionary constraint that might help predict these pathways.

A new shallow-water mud crab of the genus Hexapanopeus Rathbun, 1898 from Belize (Crustacea: Decapoda: Panopeidae).

A new species of panopeid crab from shallow coastal waters of Belize is formally described and named based upon comparative study of three small specimens obtained in the course of a regional biodiversity survey of decapod crustaceans. The herein designated holotype of Hexapanopeus klausruetzleri n. sp. is a specimen previously included in a comparative molecular phylogenetic analysis that showed it to be closely related to, but genetically distinct from, the western Atlantic species Hexapanopeus angustifrons (Benedict Rathbun, 1891). Hexapanopeus klausruetzleri n. sp. differs from this widely reported warm-temperate species in both its smoother carapace and pereopod surfaces and its generally more lobiform anterolateral teeth. The unique male first gonopod of the new species also strongly supports its morphological separation from all known congeners and other superficially similar western Atlantic panopeids with which it might be confused. To date, mature females have not been found, and the limited available collections are restricted to the vicinity of Carrie Bow Cay and nearby waters. Fourteen additional species of small panopeid and pseudorhombilid crabs, similar to Hexapanopeus klausruetzleri n. sp. in size and general habitus, were taken in the course of our studies in Belize. Most have not been previously reported from these waters, and several warrant further study to confirm their identifications.

A comprehensive molecular phylogeny of the brachyuran crab superfamily Xanthoidea provides novel insights into its systematics and evolutionary history.

Maximum likelihood and Bayesian phylogenies for the brachyuran crab superfamily Xanthoidea were estimated based on three mitochondrial and four nuclear genes to infer phylogenetic relationships and inform taxonomy. Habitat data was then used in conjunction with several diversification rates analyses (BAMM, BiSSE, HiSSE, and FiSSE) to test evolutionary hypotheses regarding the diversification of xanthoid crabs. The phylogenies presented are the most comprehensive to date in terms of global diversity as they include all four constituent families (Xanthidae, Panopeidae, Pseudorhombilidae, and Linnaeoxanthidae) spanning all oceans in which xanthoid crabs occur. Six Xanthoidea families are recognised. Panopeidae and Xanthidae sensu stricto are the two largest family-level clades, which are reciprocally monophyletic. Pseudorhombilidae is nested within and is here treated as a subfamily of Panopeidae. Former subfamilies or tribes of Xanthidae sensu lato are basally positioned clades in Xanthoidea and are here assigned family-level ranks: Garthiellidae, Linnaeoxanthidae, Antrocarcinidae, and Nanocassiopidae. The genera Linnaeoxantho and Melybia were recovered in separate clades with Linnaeoxantho being sister to the family Antrocarcinidae, while Melybia was recovered within the family Panopeidae. The existing subfamily classification of Xanthidae and Panopeidae is drastically restructured with 20 xanthid and four panopeid subfamilies provisionally recognised. Diversification-time analyses inferred the origin of Xanthoidea and Garthiellidae in the Eocene, while the other families originated during the Oligocene. The majority of genus- and species-level diversification took place during the Miocene. Ancestral state reconstruction based on depth of occurrence (shallow vs. deep water) shows some ambiguity for the most recent common ancestor of Xanthoidea and Nanocassiopidae. The most recent common ancestors of Antrocarcinidae and Panopeidae were likely deep-water species, while those of Garthiellidae and Xanthidae were probably shallow-water species. Several shifts in net diversification rates were detected but they were not associated with depth-related habitat transitions.

A new genus and species of mud crab (Crustacea, Brachyura, Panopeidae) from shoreline waters of the western Gulf of Mexico.

Several specimens of a small panopeid crab from coastal waters of the western Gulf of Mexico were long suspected to represent an undescribed species and are herein designated as representatives of a new genus. While the originally collected specimens from over four decades ago were not of gene-sequence quality, later collections from the same locality produced materials that yielded sequence data for inclusion in molecular phylogenetic studies. Building on results of those analyses, the present taxonomic description draws upon morphology to support the description of a unique species in which especially the male first gonopods differ from those of all other described panopeid genera. To date, the species remains known from only two western Gulf of Mexico sites, both of which are wave-washed intertidal rocky habitats where substrates are heavily burrowed by boring bivalves and sipunculans. While we cannot exclude the possibility that the species was introduced, recurrent collections show its populations to be at very least persistent, the species most likely being a long-overlooked among a confusing hard-substrate assemblage of small panopeid crabs.

A new genus and species of xanthoid crab (Decapoda: Brachyura) from offshore hard bank habitats in the Gulf of Mexico.

Speciose populations of small xanthoid crabs on offshore banks and reefs of the northern Gulf of Mexico include a new species that is not assignable to presently named genera. Morphological diagnoses of the new genus and species are underpinned by previously published gene sequence analyses, originally misattributed to another species but now known to apply to this taxon. Herein named Guinope tiara n. gen., n. sp., the species shows molecular phylogenetic affinities with the family Linnaeoxanthidae Stevcic, 2005, an ally of panopeid and pseudorhombilid crabs. Specimens from Occulina banks off the Florida Atlantic coast, previously regarded to represent Garthiope barbadensis (Rathbun, 1921), are not that species but instead morphologically assignable to Guinope n. gen. Whether they represent variants of Guinope tiara n. gen, n. sp. or a second species of the genus awaits the collection of fresh materials for DNA analyses.