Phylogenetic relationships within the coral crab genus Carpilius (Brachyura,Xanthoidea, Carpiliidae) and of the Carpiliidae to other xanthoid crab families based on molecular sequence data

The coral crab genus Carpilius currently includes three widely distributed species that inhabit tropical coral reefs and adjacent waters. The relationship of Carpilius to other xanthoid crabs is unknown. Previously, carcinologists considered Carpilius to be allied with crabs of the family Xanthidae (e.g., Euryozius, Liagore, and Liomera), however, recent workers have considered it to be a monotypic genus within its own family, Carpiliidae. Mitochondrial 12S- and 16S-rDNA gene fragments confirm the monophyly and distinct status of the family Carpiliidae. Within the genus Carpilius, the Caribbean species C. corallinus is basal to the two Pacific species C. maculatus and C. convexus. The Pacific species are sister taxa, despite the greater morphological resemblance of C. corallinus to the Pacific C. convexus. The relationship of the Carpiliidae (Carpilius) to other xanthoid crabs is investigated, and results of a preliminary analysis of higher xanthoid relationships did not resolve the relationships of Carpiliidae, "Xanthidae," Menippidae, Trapeziidae, and Ocypodidae to one another. A Menippidae and Carpilius relationship could not be rejected, although a Liomera, Liagore, and Carpilius relationship was.     

The evolutionary significance of heterochrony in the abbreviated zoeal development of pilumnine crabs (Crustacea: Brachyura: Xanthoidea)

The zoeal development of Pilumnus hirtellus (Linnaeus, 1761) is redescribed and the four stages are compared with the abbreviated development of Actumnus setifer (de Haan, 1835) with three stages, and Pilumnus sluiteri De Man, 1892 with two stages. A number of characters are not affected by abbreviated zoeal development and do not change during successive stage moults. Of these, some traits remain conservative at higher taxonomic level, whereas others varied between closely related pilumnid taxa, but neither provided phylogenetic information within the three pilumnines studied. However, abbreviated zoeal development affected 23 pilumnine characters that change with successive stage moults. Their timing of appearance and rate of development occur at different stages relative to the homologous process in an ancestral sequence with more zoeas, and can be attributed to three heterochronic mechanisms; postdisplacement, predisplacement and acceleration. These processes collectively appear to provide the predominant mechanism underlying the evolution of oligomerization within pilumnine zoeas.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 417–446.     

Patterns of morphological and genetic variability in UK populations of the shore crab, Carcinus maenas Linnaeus, 1758 (Crustacea: Decapoda: Brachyura)

Previous research has identified extensive inter-population variability in the morphology of the shore crab (Carcinus maenas L.). To determine the source of this variation (genetic or environmental), morphological and genetic data were analysed from crabs collected from eight sites around the coast of the UK. Ten morphometric traits were measured from over 800 crabs and the degree of morphological similarity among sites was calculated using multivariate techniques. Allozyme electrophoresis was used to investigate patterns of genetic similarity. Extensive morphological variability was detected: eight out of the ten morphometric traits analysed were useful when discriminating between crabs from each site. Discriminant function analysis revealed that over 35% of individuals could be classified to their site of origin on the basis of their morphology. In contrast, the allozyme analysis revealed low levels of genetic variability, both within the meta-population and among the crab population at each site. Pairwise comparisons revealed a moderate correlation between the degree of morphological and genetic similarity of crabs at each site, which suggests that the observed phenotypic variability has a genetic component. However, only around 20% of the phenotypic variability detected was associated with the patterns of genetic similarity. This means that patterns of morphological variability in this species are largely determined by the local environmental conditions: local factors could have a within-generation selective influence on mean trait values or C. maenas may exhibit phenotypic plasticity.     

The complete mitogenome of the hydrothermal vent crab Gandalfus yunohana (Crustacea: Decapoda: Brachyura): a link between the Bythograeoidea and Xanthoidea

Yang, J.‐S., Nagasawa, H., Fujiwara, Y., Tsuchida, S. & Yang, W.‐J. The complete mitogenome of the hydrothermal vent crab Gandalfus yunohana (Crustacea: Decapoda: Brachyura): a link between the Bythograeoidea and Xanthoidea. —Zoologica Scripta, 39, 621–630. Metazoan mitochondrial genomes (mitogenomes) are often used for all‐level phylogenetic analyses and evolution modelling. Although mitochondrial fragments facilitate studying the occurrence and dispersal of hydrothermal‐vent species, few complete mitogenomes have been determined for comprehensive analyses. We determined the complete nucleotide sequence of the bythograeid crab Gandalfus yunohana. The G. yunohana mitogenome is 15 567 bp in length and with an AT content of 69.9%. A putative control region of 625 bp was identified due to its position (between rrnS and trnI) and AT richness (72.8%), which exhibits high similarity with that of the Australian giant crab Pseudocarcinus gigas. The mitochondrial gene order is identical to the typical brachyuran mode. Codon usage, nucleotide composition and bias are well conserved as the Brachyura. Phylogenetic analyses from protein‐coding genes indicated its closest relationship with P. gigas. All the results support the close evolution distance between the Bythograeoidea and Xanthoidea, which might imply the possible origin that the only superfamily of vent crabs underwent. The G. yunohana mitogenome exhibits highly conserved characteristics with those of other decapods, especially its close relative brachyurans. A recent origin rather than the relic fauna was suggested. The present study will supply considerable data of use for both genomics and evolutionary research on hydrothermal vent ecosystems.     

Evolutionary relationships among American mud crabs (Crustacea: Decapoda: Brachyura: Xanthoidea) inferred from nuclear and mitochondrial markers,with comments on adult morphology

Members of the brachyuran crab superfamily Xanthoidea sensu Ng, Guinot & Davie (2008) are a morphologically and ecologically diverse assemblage encompassing more than 780 nominal species. On the basis of morphology, Xanthoidea is presently regarded to represent three families: Xanthidae, Pseudorhombilidae, and Panopeidae. However, few studies have examined this superfamily using modern phylogenetic methods, despite the ecological and economic importance of this large, poorly understood group. In this study we examine phylogenetic relationships within the superfamily Xanthoidea using three mitochondrial markers, 12S rRNA, 16S rRNA, and cytochrome oxidase I (COI), and three nuclear markers, 18S rRNA, enolase (ENO) and histone H3 (H3). Bayesian and maximum‐likelihood analyses indicate that the superfamily Xanthoidea is monophyletic; however, the families Xanthidae, Panopeidae, and Pseudorhombilidae, as defined by Ng et al., are not, and their representative memberships must be redefined. To this end, some relevant morphological characters are discussed. © 2013 The Linnean Society of London     

Global diversity of crabs (Crustacea: Decapoda: Brachyura) in freshwater

An assessment of the global freshwater crab diversity is presented. A total of 1,476 species in 14 families are currently known from all zoogeographical regions (except Antarctica), including 1,306 species in eight exclusively freshwater families (Pseudothelphusidae, Trichodactylidae, Potamonautidae, Deckeniidae, Platythelphusidae, Potamidae, Gecarcinucidae and Parathelphusidae). Estimates of true freshwater crab diversity including likely numbers of undescribed taxa suggest that the field remains largely in a “discovery” phase. Main ideas on the origins, diversification, and phylogeny of true freshwater crabs are briefly discussed. The economic importance of freshwater crabs is also highlighted. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Taxonomy and distribution of the hydrozoan and protozoan epibionts on Pagurus bernhardus (Linnaeus, 1758) (Crustacea, Decapoda) from Scotland

Protozoan and hydrozoan epibionts on the hermit crab Pagurus bernhardus and its shell, collected near Cumbrae Island (Scotland) were studied. The epibionts found were the following: (1) protozoans: the suctorian ciliates Ephelota plana, Acineta compressa, Conchacineta constricta, Corynophrya anisostyla; the peritrich ciliates Cothurnia mobiusi and Zoothamnium plumula; the chonotrich ciliate Chilodochona quennerstedti; and (2) hydrozoans: the species Leuckartiara sp. and Clytia sp. The morphological characteristics of the epibionts were analysed, as well as their taxonomic position. The distribution of epibionts on the crab surface and its shell was studied, and the density and biomass of epibionts were calculated on each anatomic unit. There was a differential distribution according to the type of epibiont: hydrozoans dominated in biovolume and were present mainly on the shell, meanwhile protozoans represented the highest fraction of density and they were found exclusively on the crab (principally on eyes, antennulae, antennae, maxillipeds, pereopods and uropods). The anterior area of the cephalothorax was the most colonized. On this area, the maxillipeds and second pereopods showed the highest densities. The location of each epibiont species was described. There was a correlation between the length of the crab and the total number of hydrozoans. There was a significant correlation between the right and the left units of the crab, taking into consideration the mean densities of epibionts on each anatomical unit. The shell was colonized by the same species of hydrozoa that appeared on the crab, although in a much higher density (mean 3024.38 per shell; 6.9 per crab). There was a significant difference between both species of hydrozoan epibionts with respect to the mean densities on the different areas of the shell. The zone of the shell more occupied by Clytia sp. was the apical zone of the shell, while the highest densities of Leuckartiara sp. were registered near the aperture of the shell. The hydrozoan and protozoan epibiont species found on P. bernhardus in this study represent the first mention of their presence on this hermit crab.     

Phylogenetics of Cancer crabs (Crustacea: Decapoda: Brachyura).

We used morphological, mitochondrial DNA sequence, paleontological, and biogeographical information to examine the evolutionary history of crabs of the genus Cancer. Phylogenies inferred from adult morphology and DNA sequence of the cytochrome oxidase I (COI) gene were each well resolved and well supported, but differed substantially in topology. Four lines of evidence suggested that the COI data set accurately reflected Cancer phylogeny: (1) in the phylogeny inferred from morphological data, each Atlantic species was sister taxon to an ecologically similar Pacific species, suggesting convergence in morphology; (2) a single trans-Arctic dispersal event, as indicated by the phylogeny inferred from COI, is more parsimonious than two such dispersal events, as inferred from morphology; (3) test and application of a maximum likelihood molecular clock to the COI data yielded estimates of origin and speciation times that fit well with the fossil record; and (4) the tree inferred from the combined COI and morphology data was closely similar to the trees inferred from COI, although notably less well supported by the bootstrap. The phylogeny inferred from maximum likelihood analysis of COI suggested that Cancer originated in the North Pacific in the early Miocene, that the Atlantic species arose from a North Pacific ancestor, and that Cancer crabs invaded the Atlantic from the North Pacific 6-12 mya. This inferred invasion time is notably prior to most estimates of the date of submergence of the Bering Strait and the trans-Arctic interchange, but it agrees with fossil evidence placing at least one Cancer species in the Atlantic about 8 mya.     

The ovarian maturation cycle of the Norway lobster Nephrops norvegicus (Linnaeus, 1758) (Crustacea,Decapoda) from the western Mediterranean Sea

Summary

The Norway lobster Nephrops norvegicus lives in the continental margins of the western Mediterranean Sea at depths between 100 and 600 m. It constitutes an important fisheries resource and presents a seasonal reproductive pattern. Female Norway lobsters were obtained each month from a vessel fishing off Barcelona. One hundred females caught in June 2002 were kept in the laboratory. After spawning, ovarian samples were taken every 30 days with the objective of monitoring the first steps in ovarian maturation. The gonado-somatic index (GSI) remained low over the 6 months during which the females carried their eggs, plus for a further 2–3 months. However, this study suggests that ovarian maturation is a continuous process with two different phases, taking at least 6–8 months during which the female carries its eggs. There is an increase in oocyte numbers; the germinal zone produces oogonia; and the oocytes that develop migrate to the periphery, pushing the post-ovulatory follicles to the wall of the ovary and reinforcing it for subsequent spawning. Besides this increase in the number of oocytes, vitellogenesis begins 2–3 months after the eggs hatch. Oocytes then grow and the ovaries gain weight and change from a cream color to a blackish-green. When the GSI reached 10, spawning occurred and, from then on, the ovary is mainly composed of post-ovulatory follicles.     

Xanthidae MacLeay, 1838 (Decapoda: Brachyura: Xanthoidea) systematics: A multi-gene approach with support from adult and zoeal morphology

Currently, 13 subfamilies are recognised in the brachyuran family Xanthidae: Actaeinae, Antrocarcininae, Chlorodiellinae, Cymoinae, Etisinae, Euxanthinae, Kraussiinae, Liomerinae, Polydectinae, Speocarcininae, Xanthinae, Zalasiinae and Zosiminae. This classification has been based on shared adult features like a transversely ovate carapace, well defined dorsal carapace regions, usually with lateral dentition, stout chelipeds and relatively short ambulatory legs. Such characters are now considered to be convergent. Consequently a number of higher xanthid taxa may be artifical and not monophyletic. A broad sample of 147 xanthid species representing 75 out of 124 genera from all 13 xanthid subfamilies were sampled in a multi-gene analysis. Four markers (three mitochondrial and one nuclear) were used and yielded a tree with ca. 30 xanthid clades. Monophyletic support was demonstrated for the Antrocarcininae (although substantially redefined), Cymoinae, and Polydectinae. Almost every other subfamily was para- or polyphyletic. Furthermore, the two other families of the Xanthoidea, Pseudorhombilidae and Panopeidae, were found nested within the Xanthidae. The molecular results were consistent with phylogenetic relationships implied by a suite of novel and/or neglected “ventral” adult characters including sternal characters, position of genital openings and morphology of the first zoea, instead of “dorsal” characters traditionally used to infer xanthid relationships.     

Larval development of Hexapanopeus caribbaeus (Stimpson, 1871) (Crustacea, Decapoda, Xanthoidea, Panopeidae) reared under laboratory conditions

The genus Hexapanopeus Rathbun, 1898 contains eight species,of which three are found in the Brazilian coast. The larvaldevelopment of two of these species has already been studiedin the laboratory. This article describes in detail the morphologyof the complete larval stages of Hexapanopeus caribbaeus rearedunder laboratory conditions, and the morphology is comparedwith that of the larvae of the other species of the same genus.     

The ecology and biology of Southeast Asian false spider crabs (Crustacea: Decapoda: Brachyura: Hymenosomatidae)

The false spider crabs of the family Hymenosomatidae are one of the most poorly known group of brachyuran crabs in Southeast Asia. This is largely attributed to their small size and cryptic behaviour. Despite the many studies of decapods in Southeast Asia, only eleven species of hymenosomatids are known. Of these, two genera and six species were only described in the last two years. From Singapore the largest number of species (five) has been reported probably because it has been the best explored, while from Thailand three species and Peninsular Malaysia two species are known. There are no published records of hymenosomatids from Borneo or Java. The hymenosomatid fauna is perhaps best known in Australia and New Zealand. It is anticipated that when proper collections are made and studies implemented, the Southeast Asian hymenosomatid fauna will prove to be more diverse than what has been reported. The ecology of the Southeast Asian species is reviewed, especially in the view that two species are completely freshwater, one of which is a troglobite. Most species are littoral or sublittoral in habit and very sensitive to human activities (e.g. pollution).     

Descriptions of some planktonic larvae of the Calappidae (Crustacea: Decapoda; Brachyura)

Some unidentified planktonic larval stages have been assignedto the Calappidae using Rice's key of Brachyura families. Zoeastage I from the eastern region of the Algerian coast (westernMediterranean Sea) has been attributed, according to Manningand Holthuis (1980), to Calappa granulata (L.), the only recordedrepresentative of the family from the Mediterranean Sea. Zoealstages 1-lV from the Jordanian coast of the Gulf of Aqaba, NorthernRed Sea, have been reported to the subfamily Matutinae. Comparisonsof larvae of Calappa and of Hepatus and then, of larvae of Hepatusand of Matuta, show that Hepatus larvae are more closely relatedto those of Calappa, while those of Matuta are clearly distinct.It s suggested to place the genus Hepatus with the Calappinae.Guinot (1966. 1978) placed some genera, including Hepatus. ina 'unit parthenoxystomienne, lea Aethrinae'. Larval developmentis unknown for Aethra, Actaeomorpha. Hepatella andOsachila.However, comparison of Hepatus larvae with those of Parthenopidaeshows that larval characters, particularly the setation of theendopod of the first maxilliped, are not conclusive for supportingsuch a regrouping. Moreover, the Calappidae have a combinationof primitive and advanced features which suggest that theirgrouping in the Oxystomata seems more likely to be an adaptationto a similar way of life. The morphotypal taxon Oxystornatahas been definitively rejected by Guinot (1978).     

Isolation of polymorphic microsatellite loci from the endangered mudflat crab Chasmagnathus convexus (Crustacea: Decapoda: Varunidae)

Chasmagnathus convexus is a mudflat crab that is distributed throughout East Asia. However, most populations of this species in Korea are endangered due to continuing seashore developments and habitat fragmentations. In this study, we isolated and characterized nine microsatellite loci from C. convexus. The number of alleles per locus ranged from 3 to 23. The expected heterozygosities ranged from 0.145 to 0.974 and the observed heterozygosities ranged from 0.150 to 0.947. Cross-species amplifications for 10 other Varunid crab species were performed. These microsatellite markers could be useful for planning conservation strategies for C. convexus and related species.     

On the marine sister groups of the freshwater crabs (Crustacea: Decapoda: Brachyura)

Freshwater crab sister group relationships with marine eubrachyuran families were investigated. A morphology-based cladistic analysis was conducted on representatives of the freshwater crab families Deckeniidae, Gecarcinucidae, Parathelphusidae, Potamidae, Potamonautidae, Pseudothelphusidae, and Trichodactylidae using a disparate assemblage of marine heterotreme and thoracotreme brachyurans as possible sister groups. The monophyly of the freshwater crabs sensu lato is falsified. The family Trichodactylidae and the marine portunid subfamily Carcininae form basal groups within the superfamily Portunoidea. The monophyly of the Pseudothelphusidae and the Paleotropical freshwater crab families is supported, and this clade is the sister group of the Thoracotremata (Gecarcinidae, Grapsidae s.l., and Ocypodoidea). The origin, groundplan, and diversification of freshwater crabs are discussed in the context of previously published scenarios of their evolution.     

蟹类精子超微结构的比较研究

应用光镜和电镜,比较研究了三疣梭子蟹,中华绒螯蟹和长江华溪蟹的成熟精子。揭示３种蟹精子都是不能游动的无鞭毛精子,呈球形,前后略扁,精子前端出现一光滑的小圆面,圆面四周有内陷的沟环。沟环之后,精子表面凹凸不平,并伸出多数辐射臂。３种蟹精子均为高度特化的细胞,外被质膜,内含细胞核,顶体及退化的细胞质。     

The Grapsidae and Ocypodidae (Decapoda: Brachyura) of Tanzania

Twenty-nine species of the Grapsidae and 15 species of the Ocypodidae were collected from the littoral zone in the area around Dar es Salaam. Nine species of Grapsidae and three of Ocypodidae were new records for the East African area, bringing the total of recorded species for the region to 35 and 26 respectively. The distribution of these crabs indicates a very limited endemism in the western Indian Ocean, with the majority of species forming an attenuated extension of the abundant Indo/West Pacific brachyuran fauna.
The habitat preference of each species is described. Within each subfamily there is a relative uniformity in general habitat preference, mode of life and manner of feeding. The species of each subfamily are then separated by substrate choice and preferred level on the shore. The greatest numbers of species occur in the most diverse habitats—the creek mangrove, the coastal mangrove and the reef flat.     

Morphology and ultrastructure of the adult ovarian cycle in Mithracidae (Crustacea,Decapoda, Brachyura,Majoidea)

The ultrastructure of the ovary during development and yolk production is poorly known in Brachyura and Majoidea in particular. Here, we describe the histology, histochemistry and ultrastructure of the adult ovarian cycle in four Mithracidae species from three different genera: Mithrax hispidus, Mithrax tortugae, Mithraculus forceps and Omalacantha bicornuta. All species showed a similar pattern of ovarian development and vitellogenesis. Macroscopically, we detected three stages of ovarian development: rudimentary (RUD), developing (DE) and mature (MAT); however, in histological and ultrastructural analyses, we identified four stages of development. The oocytes of the RUD stage, during endogenous vitellogenesis, have basophilic cytoplasm filled with dilated rough endoplasmic reticulum. The reticulum lumen showed many granular to electron-dense materials among the different stages of development. The Golgi complexes were only observed in the RUD stage and are responsible for releasing vesicles that merge to the endogenous or immature yolk vesicles. At the early DE stage, the oolemma showed many coated and endocytic vesicles at the cortex. The endocytic vesicles merge with the endogenous yolk to form the exogenous or mature yolk vesicles, always surrounded by a membrane, characterizing exogenous vitellogenesis. The exogenous yolk vesicles comprise glycoproteins, showing only neutral polysaccharides. At the late DE stage, endocytosis still occurs, but the amount of endogenous yolk decreases while the exogenous yolk increases. The late DE stage is characterized by the beginning of chorion production among the microvilli. The MAT stage is similar to the late DE, but the endogenous yolk is restricted to a few cytoplasmic areas, the ooplasma is filled with exogenous yolk, and the oolemma has very few coated vesicles. In the MAT stage, the chorion is fully formed and shows two electron-dense layers. The ovarian development of the species studied has many similarities with the very little known Majoidea in terms of the composition, arrangement and increment of the yolk vesicles during oocyte maturation. The main differences are in the vitellogenesis process, where immature yolk formation occurs without the direct participation of the mitochondria but with the participation of the rough endoplasmic reticulum in the endogenous phase.     

Variants in the histochemical patterns of lipids in the hepatopancreas of Scylla serrata (Forskal) (Brachyura, Decapoda).

The highest concentrations of phospholipid, neutral lipid and fatty acids were observed in the R cells and connective tissue of the hepatopancreas of Scylla serrata. The basal parts of the B cells and apical parts of the cells lining the main duct also showed moderate presence of these substances. The E cells however, except at their cell membranes were found to be devoid of lipids. F cells on the other hand exhibited lipoid complexes. Considerable reduction in the staining intensity of fatty acids were noticed 4 h after the bilateral ablation of eyestalks, neutral lipid undergo depletion 24 h after the operation whereas phospholipid reserves increase 48 h after the eyestalk removal. A fall in the quantity of neutral lipid and phospholipid was conspicuous when eyestalk extract was injected into normal or destalked crabs. From the present data it appears that R cells and connective tissue form major sites of lipid storage and in an intermolt animal eyestalk factor(s) may have an important role in the control of lipid metabolism.