The invasive green crab and Japanese shore crab: behavioral interactions with a native crab species,the blue crab

Blue crabs, Callinectes sapidus (Rathbun), are an ecologically and commercially important species along the East coast of North America. Over the past century and a half, blue crabs have been exposed to an expanding set of exotic species, a few of which are potential competitors. To test for interactions with invasive crabs, juvenile C. sapidus males were placed in competition experiments for a food item with two common non-indigenous crabs, the green crab Carcinus maenas (L.) and the Japanese shore crab, Hemigrapsus sanguineus (De Haan). Agonistic interactions were evaluated when they occurred. In addition, each species’ potential to resist predators was examined by testing carapace strength. Results showed that C. maenas was a superior competitor to both C. sapidus and H. sanguineus for obtaining food, while the latter two species were evenly matched against each other. Regarding agonism, C. sapidus, was the “loser” a disproportionate number of times. C. sapidus carapaces also had a significantly lower breaking strength. These experiments suggest that both as a competitor, and as potential prey, juvenile blue crabs have some disadvantages compared with these common sympatric exotic crab species, and in areas where these exotics are common, juvenile native blue crabs may be forced to expend more energy in conflict that could be spent foraging, and may be forced away from prime food items toward less optimum prey.     

The first-stage zoeas of Carpilius convexus (Forskal, 1775) and Carpilius maculatus (Linnaeus, 1758) (Crustacea: Decapoda: Brachyura: Xanthoidea: Carpiliidae): an example of heterochrony

The first-stage zoeas of Carpilius convexus (Forskål,1775) and Carpilius maculatus (Linnaeus, 1758) are describedand fully illustrated. Both these Indo-West Pacific speciesexhibited a unique xanthoidean character for the basis of thesecond maxilliped, which possesses five (arranged 1,1,1,2) setaeinstead of the expected four (arranged 1,1,1,1). A comparisonwith the zoeal stages of Carpilius corallinus (Herbst, 1783)as reported by Laughlin et al. (Laughlin et al., 1983) revealedmarked differences including the possession of carapace lateralspines (v. absent in C. convexus only), the subterminal setationof the distal maxillule endopod segment with two subterminalsetae (versus one subterminal seta in Indo-West Pacific species),the terminal setation of the distal maxillule endopod segmentwith four setae (versus three setae in C. convexus only), twolateral spines on the telson (versus three in Indo-West Pacificspecies) and the number of zoeal stages. The first-stage zoeasof the two Indo-West Pacific species appear to have hatchedin a more advanced state of development than those of C. corallinus,and the expression of a number of characters has been accelerated(early onset). In fact, the zoeal stages of both Indo-West Pacificspecies and carpiliid species appear to be abbreviated becausethe first zoeas are considered to be equivalent to the third-stagezoeas of C. corallinus.     

Molecular ecology, speciation, and evolution of the reef fish genus Anisotremus

Grunts (family Haemulidae) in the genus Anisotremus comprise 10 described species which occur predominantly on coral reefs and subtropical rocky reefs in the Neotropics of the Tropical Eastern Pacific the Caribbean and adjacent waters. In this study, the phylogenetic relationships for all described species were examined based on one mitochondrial gene (cytochrome b) and one nuclear marker (the first intron of the ribosomal protein S7). Closely related species of the genus Haemulon and the single representative of the genus Genyatremus, G. luteus were also included in the analysis, following the suggestion, based on morphological data, that they may be closely related to, or possibly placed within, Anisotremus. Two sweetlips, genus Plectorhinchus, which belong to the recognized sister subfamily Plectorhynchinae, were used as outgroups. Two species pairs in the ingroup genus Anisotremus are geminate species, which presumably diverged after the rise of the Isthmus of Panama, thus providing an internal molecular clock calibration. This allowed for a comprehensive evolutionary scenario for the genus. Our data were consistent with two independent allopatric speciation events (geminate species), and one likely sympatric speciation event. Our data were also consistent with the placement of Genyatremus and Haemulon within the genus Anisotremus, thus breaking its monophyletic status. As proposed for other coral reef species, Anisotremus seems to have colonized coral reefs from non-coral reef habitats. In addition, ancestral species seem to have been widespread, since species that are currently found in Brazilian and Pacific waters are basal on the phylogenetic tree. Finally, this study adds to the hypothesis of a secondary evolutionary center in the New World.     

Tri-locus sequence data reject a “Gondwanan origin hypothesis” for the African/South Pacific crab genus Hymenosoma

Crabs of the family Hymenosomatidae are common in coastal and shelf regions throughout much of the southern hemisphere. One of the genera in the family, Hymenosoma, is represented in Africa and the South Pacific (Australia and New Zealand). This distribution can be explained either by vicariance (presence of the genus on the Gondwanan supercontinent and divergence following its break-up) or more recent transoceanic dispersal from one region to the other. We tested these hypotheses by reconstructing phylogenetic relationships among the seven presently-accepted species in the genus, as well as examining their placement among other hymenosomatid crabs, using sequence data from two nuclear markers (Adenine Nucleotide Transporter [ANT] exon 2 and 18S rDNA) and three mitochondrial markers (COI, 12S and 16S rDNA). The five southern African representatives of the genus were recovered as a monophyletic lineage, and another southern African species, Neorhynchoplax bovis, was identified as their sister taxon. The two species of Hymenosoma from the South Pacific neither clustered with their African congeners, nor with each other, and should therefore both be placed into different genera. Molecular dating supports a post-Gondwanan origin of the Hymenosomatidae. While long-distance dispersal cannot be ruled out to explain the presence of the family Hymenosomatidae on the former Gondwanan land-masses and beyond, the evolutionary history of the African species of Hymenosoma indicates that a third means of speciation may be important in this group: gradual along-coast dispersal from tropical towards temperate regions, with range expansions into formerly inhospitable habitat during warm climatic phases, followed by adaptation and speciation during subsequent cooler phases.     

Phylogenetic relationships and natural hybridization in rabbitfishes (Teleostei: Siganidae) inferred from mitochondrial and nuclear DNA analyses

Phylogenetic relationships of rabbitfishes (the family Siganidae), ecologically important components as primary consumers in coral reef communities, were studied using mitochondrial cytochrome b gene and nuclear ITS1 (internal transcribed spacer 1) sequence analyses. The analyses of 19 out of 22 species known in the Western Pacific region revealed that siganids are genetically clustered into three major clades, which are characterized by some morphological and ecological traits. Between closely related species, such as Siganus guttatus-S. lineatus and S. virgatus-S. doliatus, and also between two morphs recognized in S. corallinus, small but discernible genetic differentiation was found, implying that the components of each pair are incipient species. On the other hand, between some species, such as S. fuscescens-S. canaliculatus and S. unimaculatus-S.vulpinus, individuals of the components of each pair were found to construct a genetic mosaic, suggesting that the components are genetic color morphs within a single biological species, respectively. Moreover, evidence from morphological characters, mtDNA, and nuclear DNA gave an inconsistent picture of identity and relationships for several individuals. They were regarded as hybrids or individuals with hybrid origin. Such instances were observed not only between closely related species, such as S. guttatus-S. lineatus, S. virgatus-S. doliatus, and two morphs (incipient species) in S. corallinus, respectively, but also between distantly related ones, such as S. corallinus-S. puellus. In fact, more than half of the species examined (11/20, when treating the two morphs in S. corallinus as independent species) were involved in hybridization. These suggest that hybridization is much more prevalent in marine fishes than previously assumed, and may have some relevance to their diversification.     

Molecular phylogeny of Eastern Pacific porcelain crabs, genera Petrolisthes and Pachycheles, based on the mtDNA 16S rDNA sequence: phylogeographic and systematic implications

Porcelain crabs, genera Petrolisthes and Pachycheles, are diverse and abundant members of the eastern Pacific near-shore decapod crustacean community. Morphology-based taxonomic analyses of these crabs have determined groupings of affiliated species, but phylogenetic relationships remain unknown. We used sequence data from the mitochondrial 16S rRNA gene of 46 species of eastern Pacific porcelain crabs to perform phylogenetic analyses by distance and parsimony methods. Our results are used to compare the taxonomic significance of morphological and molecular characters, to examine sequence divergence rates of crab 16S rRNA genes, and to analyze the phylogeographic history of these crabs. Our phylogenetic trees indicate that the genus Petrolisthes is divided into two main clades, reflecting morphological features. One clade contains primarily tropical species, and the other contains species from throughout the eastern Pacific, as well as species in the genera Allopetrolisthes and Liopetrolisthes. Phylogenetic trees of Pachycheles suggest an antitropical distribution; north and south temperate species form one clade and tropical species form a second clade. Sequence divergence rates of the 16S rRNA gene from three pairs of geminate species can be used to date divergence times, and we discuss porcelain crab phylogeographic patterns in relation to paleogeographic events.     

Phylogenetic ecology of gall crabs (Cryptochiridae) as associates of mushroom corals (Fungiidae)

Coral‐associated fauna is a relatively understudied topic. Hence, the nature of the relationship between an associated organism and its host is usually unknown. In the present study, the obligate associations between gall crabs (Decapoda: Cryptochiridae) and mushroom corals (Scleractinia: Fungiidae) are reviewed from a phylogenetic perspective. Based on field surveys, examination of museum material and a literature review, a total of 35 fungiid species have been found that act as hosts for four gall crab species. Fungiid‐associated gall crabs appear to be more geographically widespread than previously known, with new records showing their occurrences from the Red Sea and western Indian Ocean all the way to the central Pacific Ocean. The obligate nature of the association between cryptochirids and their hosts makes them an ideal model taxon to test for possible cospeciation events. The congruence between their phylogenies was tested by using the program Jane 4.0, resulting in cospeciation and duplication events between the crabs and their host corals. The sharing of several closely related host coral species by the same gall crab species or genus may provide support to models indicating phylogenetic relationships within the Scleractinia.     

Re-evaluation of the systematics of the endemic corals of Brazil by molecular data

Recent genetic work on various coral genera has shown that morphological convergence between Atlantic and Pacific corals obscures evolutionary relationships and inferred levels of endemicity between the regions. Based on DNA sequences from nuclear and mitochondrial loci that provide higher resolution than those previously presented, this study shows that relationships within parts of the Atlantic coral fauna are also in need of substantial revision. The data presented here indicate that (1) the endemic Brazilian genus Mussismilia is a monophyletic clade, (2) Mussismilia is more closely related to the Caribbean Faviidae than Mussidae, the family in which it is currently placed, (3) the Brazilian endemic coral Favia leptophylla is much more closely related to Mussismilia than other species of Favia and has most likely been incorrectly placed in the genus Favia and (4) the other endemic Favia species found in Brazil, Favia gravida, is genetically distinct from Favia fragum, a Caribbean congener with which it is frequently synonymized. The nuclear data also suggest the possible presence of a cryptic species within Mussismilia, but additional sampling and morphological information is required to confirm this finding.     

Hemicyclops columnaris sp.n. (Copepoda, Poecilostomatoida, Clausidiidae) Associated with a Coral in Panama (Pacific Side)

Hemicyclops columnaris sp.n. is described from the scleractinian coral Porites lobata Dana on the Pacific coast of Panama. The new species may be distinguished from its 23 congeners by a combination of the body size and the shape of the genital segment in the female. Although certain members of the genus Hemicyclops are associated with crustaceans, polychaetes and other marine invertebrates, this is the first record of a species of this genus from a hard coral.     

Peltidiphonte gen. n., a New Taxon of Laophontidae (Copepoda: Harpacticoida) from Coral Substrates of the Indo-West Pacific Ocean

A new genus of the harpacticoid family Laophontidae is described and named Peltidiphonte gen. n. Eight new species are assigned to this genus; they were collected from different locations in the Indo-West Pacific Ocean, including the Comoros, the Kenyan coast, the Red Sea, the Andaman Islands, the northern coast of Papua New Guinea, the Solomon Islands and the northeastern coast of Australia. Most of the specimens were collected from dead coral substrates, suggesting a close affinity between the members of the new genus and this substrate. Peltidiphonte gen. n. can easily be discriminated from other genera of the family by the extremely depressed body and by the shape of the antennule, bearing two (or three) processes on the first segment and a hook-like process along the outer margin of the second segment. An identification key for the new genus is provided.     

Does the handedness of the pebble crab Eriphia smithii influence its attack success on two dextral snail species?

The predaceous crab Eriphia smithii (Xanthidae) has one larger claw with molar teeth on either the right or the left cheliped, which it uses to crush the shell of prey. Whether the handedness of crabs affected successful predation on two snail species, Nerita albicilla (Neritidae) and Planaxis sulcatus (Planaxidae) was experimentally investigated. The fate of snails of each species was analysed by multiple logistic regression with three explanatory variables: handedness, shell-size index and individuality of crabs. No effect of handedness was detected in attacks on N. albicilla , probably as a result of the spherical and more symmetrical shell morphology of this species. In contrast, right-handedness contributed to greater attack success on P. sulcatus , which has more conical shells. Further investigation of how snail shells were broken revealed that left-handed crabs had more difficulty breaking the aperture of larger P. sulcatus , which was thought to cause the difference in attack success between right- and left-handed crabs. The advantages conferred by handedness are discussed.     

Hermit crab biocoenoses: a worldwide review of the diversity and natural history of hermit crab associates

The symbiotic associates of hermit crabs (excluding parasites and flora) are reviewed worldwide. The review includes species found on the shells occupied by hermit crabs (epibiotic species), species boring into these shells (endolithic species), species living within the lumen of the shell (either free-living or attached to the shell), species attached to the hermit crabs themselves, and hypersymbionts. In total over 550 invertebrates, from 16 phyla are found associated with over 180 species of hermit crabs. Among these associates, 114 appear to be obligate commensals of hermit crabs, 215 are facultative commensals, and 232 are incidental associates. The taxa exhibiting the highest number of associates are arthropods (126), polychaetes (105), and cnidarians (100). The communities of species associated with Dardanus arrosor, Paguristes eremita, Pagurus bernhardus, Pagurus cuanensis, and Pagurus longicarpus are the best studied and harbor the most diverse assemblages of species. While trends in biodiversity of hermit crab assemblages do not follow predicted patterns (e.g., hermit crabs within the Indo-West Pacific do not harbor more species than those from temperate regions), this is suggested to reflect a lack of sampling rather than a true representation of the number of associates. Hermit crabs date to at least the Cretaceous and provided a niche for a number of groups (e.g., hydractinians, bryozoans, polydorids), which were already associates of living gastropods. Apparently hermit crab shells initially supplied a substrate for settlement and then these symbiotic relationships were reinforced by enhanced feeding of symbionts through the activity of the hosts. Through their use and recycling of gastropods shells, hermit crabs are important allogenic ecosystem engineers in marine habitats from the intertidal to the deep sea. Hermit crabs benefit from some symbionts, particularly cnidarians and bryozoans, through extension of shell apertures (alleviating need to switch into new shells) and by providing protection from predators. However, hermit crabs are also negatively impacted (e.g., decreased reproductive success, increased predation) by some symbionts and a review of egg predators is provided. Thus, the symbiotic relationships between hermit crabs and many associates are difficult to characterize and often exhibit temporal changes depending on environmental and biological factors. Research on the biology of these symbionts and the costs/benefits of their associations with hermit crabs are analyzed. While some associates (e.g., Hydractinia spp.) have been studied in considerable detail, for most associations little is known in terms of the impacts of symbionts on hosts, and future experimental studies on the multitude of relationships are suggested.     

Folliculinid ciliates: a new threat to Caribbean corals?

This is the first report of a putative pathogenic ciliate protozoan that has been associated with Caribbean corals. Previously, only 2 species of the phylum Ciliophora had been linked to coral diseases, and they were exclusive to the Indo-Pacific region. In this study, a ciliate of the genus Halofolliculina was found on 10 hard coral species at the National Parks of Los Roques and Morrocoy, Venezuela. The general morphology of this ciliate is very similar to that of Halofolliculina corallasia from the Indo-Pacific, which is known to cause skeletal eroding band. None of the other 31 genera in the family Folliculinidae are known to cause diseases in corals or in any other animal species. The presence of this ciliate, which shows a prevalence comparable to that of other epizootics in the Caribbean, suggests it could be a new threat to the coral reefs of this region.     

A preliminary assessment of the invasiveness of the Indo-Pacific sponge Chalinula nematifera on coral communities from the tropical Eastern Pacific

This is the first report of a sponge that overgrows live corals in the tropical Eastern Pacific ocean. Chalinula nematifera, native from the Indo-Pacific region, is an invasive sponge recorded for the first time in 2003 on coral communities from the Isla Isabel National Park (Mexican Pacific Ocean). Later, in 2006, it was found also on coral reefs from the Cabo Pulmo National Park; 217 nautical miles far away. It has been suggested that C. nematifera was introduced as fouling on ship hulls that have arrived at Isla Isabel from the Indo-Pacific. In this paper we examined the habitat specificity and the distribution and abundance through time of C. nematifera. While there were no significant variations in abundance through time, this species showed a very high specificity for living on live corals of the genus Pocillopora (94% vs. 6% on rocks). One of the environmental parameters that may explain this specificity for ramified corals is the low light intensity inside the coral colony, which was 96% lower than outside it. Coral reefs are currently struggling with a multitude of impacts that have weakened their resilience and pushed them away from equilibrium. As a result, more attention on ecology of corals is necessary. Although the abundance of C. nematifera seems to be stable, long-term monitoring programs (including studies of growth rates and recruitment) are needed to determine if this species could represent a threat to the Mexican coral ecosystem in the future.     

Global diversity of crabs (Aeglidae: Anomura: Decapoda) in freshwater

The freshwater anomuran crabs of the family Aeglidae are all restricted to southern South America occurring in Chile, Brazil, Bolivia, Uruguay, Paraguay, and Argentina. The family consists of a single genus, Aegla, containing 63 currently described species. There are another 5–10 known yet undescribed species to complement this diversity. The aeglids occur in freshwater lakes, streams, rivers, and in caves with freshwater. The origin of the family appears to be from marine ancestors from the Pacific invading streams in Chile about 75 mya radiating both in Chile and again on the eastern side of the Andes, particularly in Brazil. Of the 63 species, 23 or 36.5% are considered under threat and are in need of conservation action. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

A comparison of the predatory impacts of an invasive and native crab species using a functional response approach

The European green crab (Carcinus maenas) is invasive on the West coast of North America, but the ecological consequences of this invasion remain poorly understood. Comparative functional response analysis has arisen as a method of elucidating ecological consequences of invasive species by comparing the impact of these species to native analogues. Through comparative functional response experiments of green crabs and native red rock crabs (Cancer productus) we found that green crab predation increased asymptotically (Type II functional response) when fed increasing densities of Pacific oysters (Magallana gigas), while red rock crab predation displayed a sigmoidal (Type III) response. At high oyster densities red rock crabs consume more Pacific oysters than green crabs do, due to their reduced handling time, though green crabs consume more Pacific oysters relative to their size than red rock crabs. However, compared to red rock crabs, green crabs consume more oysters at low prey densities, which implies that they have a larger, potentially destabilizing impact on low densities of Pacific oysters. As green crabs continue to spread across the West coast of North America, Pacific oysters will face increased predation pressure. Our results show the advantage of using functional response analysis to compare density dependent predation between an invasive species and a native species to predict the ecological consequences of invasions.

     

Cryptic invasions of the crab Carcinus detected by molecular phylogeography

Coastal marine ecosystems world-wide are threatened by invasions of nonindigenous species. The ubiquity of marine sibling species identifiable only by genetic analysis suggests that many invasions are cryptic and therefore undetected, causing an underestimation of the actual number and impacts of invading species. We test this hypothesis with European crabs in the genus Carcinus that have invaded five regions globally. Partial 16S ribosomal RNA gene sequences confirm sibling species status of morphologically similar Atlantic C. maenas and Mediterranean C. aestuarii . Based on 16S rRNA haplotypes, crabs from California, New England and Tasmania were all C. maenas . However, we report the cryptic multiple invasion of both species in Japan and South Africa, where only C. aestuarii and C. maenas , respectively, were previously recognized.     

Delimitation of morphologically similar sponge crab species of the genus Pseudodromia (Crustacea, Decapoda, Dromiidae) from South Africa

Stewart, B. A., Gouws, G., Daniels, S. R. & Matthee, C. A. (2004). Delimitation of morphologically similar sponge crab species of the genus Pseudodromia (Crustacea, Decapoda, Dromiidae) from South Africa. — Zoologica Scripta , 33 , 45–55.
Presently, three Pseudodromia sponge crab species are recognized, all of which are endemic to the continental shelf off the coast of South Africa. Two of these differ only in the morphology of their rostral teeth, making them difficult to distinguish, and can thus be considered as cryptic species. In addition they have very similar distribution ranges, thus raising doubts as to their specific status. Discriminant function analysis of morphometric data, differentiation at 10 allozyme loci, and sequence data derived from the 12S rRNA mitochondrial gene were used to test whether specimens identified as P. latens and P. rotunda are morphological forms of a single, widespread species, or represent two, distinct, reproductively isolated species, and to establish whether these two taxa are sister species, and thus form a monophyletic entity. The presence of fixed allele differences at three, and strong genetic heterogeneity at five other allozyme loci, indicating no gene flow occurring between sympatric populations, as well as the relatively high degree of 12S rRNA and allozyme genetic differentiation observed, supported the recognition of P. latens and P. rotunda as separate species. The 12S rRNA topology suggested that the genus Pseudodromia , as presently constituted, is paraphyletic, thus inferring that the morphological characters used to define this taxon might not be useful for phylogenetic inferences. It was concluded that in view of the uncertainties raised regarding the designation and composition of certain genera within the family Dromiidae, further rigorous analyses of morphological and genetic data are needed to further our understanding of the taxonomy of the sponge crabs.     

A revision of the octocoral genus Ovabunda Alderslade, 2001 (Anthozoa,Octocorallia, Xeniidae)

The family Xeniidae (Octocorallia) constitutes an abundant benthic component on many Indo-West Pacific coral reefs and is ecologically important in the Red Sea. The genus Ovabunda Alderslade, 2001 was recently established to accommodate previous Xenia species with sclerites comprised of a mass of minute corpuscle-shaped microscleres. The aim of the present study was to examine type material of Xenia species in order to verify their generic affiliation. We present here a comprehensive account of the genus Ovabunda, using scanning electron microscopy to depict sclerite microstructure. We assign three Xenia species to the genus: O. ainex comb. n., O. gohari comb. n., and O. crenata comb. n.; and synonymize several other species of Ovabunda. We provide a key to Ovabunda species and conclude that they are mainly confined to the Red Sea, with some occurrence in the West Indian Ocean.     

Host specificity of <Emphasis Type="Italic">Sacculina carcini</Emphasis>, a potential biological control agent of the introduced European green crab Carcinus maenas in California

The European green crab, Carcinus maenas, is an introduced marine predator established on the west coast of North America. We conducted laboratory experiments on the host specificity of a natural enemy of the green crab, the parasitic barnacle Sacculina carcini, to provide information on the safety of its use as a possible biological control agent. Four species of non-target, native California crabs (Hemigrapsus oregonensis, H. nudus, Pachygrapsus crassipes and Cancer magister) were exposed to infective larvae of S. carcini. Settlement by S. carcini on the four native species ranged from 33 to 53%, compared to 79% for green crabs. Overall, cyprid larvae tended to settle in higher numbers on individual green crabs than on either C. magister or H. oregonensis. However, for C. magister this difference was significant for soft-shelled, but not hard-shelled individuals. Up to 29% of the native crabs arrested early infections by melanizing the rootlets of the parasite. Most native and green crabs settled on by S. carcini became infected, especially when settled on by >3 cyprids. Infected green crabs died at more than twice the rate of uninfected green crabs. In contrast to green crabs, all infected native crabs died without producing an externa (reproductive sac). At high settlement intensities, infected native crabs frequently exhibited neurological symptoms (twitching, loss of movement) before death. These results indicate that use of S. carcini as a biological control agent could result in the death of native crabs. The magnitude of this effect would be proportional to the density of infected green crabs in the environment and the probability that cyprids would contact native crabs in the wild. Potential benefits of biological control should be assessed in relation to these potential non-target effects.