A key for the identification of the known larval stages of the Mediterranean Brachyura

Summary

Based on the accounts by various authors, an identification key has been constructed for the planktonic zoeae of the 70 species of Mediterranean Brachyura whose larvae are known. The key is based on primary (spines of the carapace, exopodite of the antenna, spines of telson forks) as well as secondary (setation of mouthparts, processes and spines on abdominal somites) morphological characteristics. It allows the identification of the zoeae of 64 species plus those of Liocarcinus genus (six species).     

Hydromedusae of the Mediterranean Sea

Mediterranean hydromedusae can be classified in several categories: common species, widely distributed throughout the Mediterranean and rare species reported from the Pacific, Indian or Atlantic Oceans; seasonal species comprising cold-season species with a boreal affinity and warm-season species with a hot to temperate affinity; neritic and mid-ocean species; surface species and meso-, infra-, and bathypelagic species. Holoplanktonic species are known to be associated with stable hydrological conditions (offshore waters, deep waters, summer and winter homothermy), whereas meroplanktonic species are more likely to be found in more disturbed hydrological conditions (coastal waters, surface waters, spring and autumn periods of temperature fluctuation). A general pattern of medusa distribution is established based upon comparative study throughout the Mediterranean.     

A catastrophic mass-mortality episode of gorgonians and other organisms in the Ligurian Sea (North-western Mediterranean), summer 1999

In the late summer of 1999, an extensive mortality of gorgonians and other epi-benthic organisms was observed in the Ligurian Sea (Mediterranean Sea) from the Tuscan Archipelago to Marseille. Quantitative data from Tino Island and Portofino Promontory indicated that the proportion of affected gorgonians ranged from 60% to 100% in populations having a density of 9–27.8 colonies m⁻², suggesting that millions of sea fans died along the coast of Liguria. This mass mortality episode coincided with a sudden increase of sea water temperature down to more than 50 m depth. Laboratory analyses showed that the colonies stressed by high temperature also underwent extensive attack by microrganisms (protozoans and fungi), which are interpreted as opportunistic pathogens.     

Genetic differentiation of white sea bream within the Lion's Gulf and the Ligurian Sea (Mediterranean Sea)

An allozymic study was conducted on 190 individuals of white sea bream, Diplodus sargus sampled on five locations throughout the Lion's Gulf and the Ligurian Sea (Mediterranean Sea), in order to identify genetic structure. Electrophoretic surveys carried out on muscle and liver tissues identified 25 loci of which 12 were polymorphic ( P 0.95). G -test analysis shows significant differences on allelic frequencies between the five stations at six loci. Examination of the spatial structure was performed using Nei's distances and F statistics, and indicated significant genetic differences between three groups. A group which clustered Blanes (Spain), Marseille (France) and Livorno (Italy) where the absence of genetic difference can be explained by migration of larvae and adults along a coastal 'continuum'. The Elba sample (Italy) shows genetic divergence from other samples and this difference may result from isolation due to limited migration of larvae and adults. Banyuls (France), differs from all other stations. Several hypotheses are proposed to explain genetic patterns including local current systems, larval dispersal, geographic isolation and historical effects, and variation in the size classes of sampled individuals between sites.     

Changes of zooplankton communities in the Gulf of Tigullio (Ligurian Sea, Western Mediterranean) from 1985 to 1995. Influence of hydroclimatic factors

Year-to-year variations in abundance and composition of zooplanktonwere studied in the Ligurian Sea at a station sampled two timesa month between 1985 and 1995. As a break of 2 years (April1989–December 1990) occurred in the time series, the STATISmethod was chosen instead of time series analysis. Each of thenine sampled years was a single table of monthly or seasonalaverage densities of 26 plankton taxa. STATIS allowed (i) estimationof similarity between each yearly table, (ii) visualizationof the trajectories of both species and observations (seasons)from one year to another, and (iii) associations of particularspecies, which showed similar temporal variations, to be determined.A strong seasonal variation was evident for most species, andyears 1987, 1992 and 1994 were different from the others. Trajectoriesindicated which species were stable and which were characterizedby small or large fluctuations during the nine years. Five differenttaxa associations were detected. For each association, the mostrepresentative period was identified, where each period wasa group of months obtained by clustering on species abundances.Taking into account hydro-climatic factors in the representativeperiods, a contingency discriminant analysis allowed us to identifyand characterize the most discriminant environmental parametersassociated with each group of species. Environmental factorsthat best discriminated the different representative periodswere atmospheric pressure, current speed and direction, andwater temperature.     

Molecular phylogeny of grapsoid crabs (Decapoda, Brachyura) and allies based on two mitochondrial genes and a proposal for refraining from current superfamily classification

Grapsoid and ocypodoid crabs receive a lot of attention in the literature due to their predominance and important role as primary and secondary consumers in intertidal as well as supratidal marine habitats. They are especially species‐rich in the tropics, where they have been found to repeatedly invade terrestrial and freshwater habitats. However, the systematics of the crabs belonging to these two superfamilies is still not settled, despite recent steps clarifying phylogenetic relationships and introducing new taxa. In this study, a molecular phylogeny of grapsoid crabs primarily based on East African representatives is constructed based on DNA sequences of the mitochondrial small and large ribosomal subunits (12S and 16S rRNA), thus complementing previous molecular taxonomic studies that had been carried out with the American and East Asian fauna. In addition, selected representatives of all ocypodoid families and subfamilies were included. The monophyly of Grapsidae, Ocypodidae (sensu stuctu), Sesarmidae and Varunidae is well confirmed, if the genera Cyclograpsus, Helice are considered Varunidae and Euchirograpsus a Plagusiidae, as previously suggested. The monophyly of the family Gecarcinidae cannot be supported with our data. The family Plagusiidae in its present composition is polyphyletic. Special attention was given to the large family Sesarmidae, which has many endemic genera in the Indo‐West Pacific. According to this study, two of the most speciose genera, Chiromantes and Parasesarma, are not monophyletic and need to be redefined. On the higher taxonomic level, it becomes evident that both superfamilies, Grapsoidea and Ocypodoidea, are not monophyletic in their current composition, as exemplified by a proposed sister group relationship of Varunidae and Macrophthalmidae. These results confirm those from previous molecular studies and we therefore propose to refrain from the traditional use of the Grapsoidea and Ocypodoidea as monophyletic superfamilies and treat the constituent families separately.     

Foregut morphology and ontogeny of the mud crab Dyspanopeus sayi (Smith, 1869) (Decapoda,Brachyura, Panopeidae)

The morphology of the foregut of the Say's mud crab Dyspanopeus sayi was described in adults and larvae. The ossicle system was illustrated based on a staining method with Alizarin-Red. The gastric teeth and cardio-pyloric valve were dissected and examined using optical and scanning electron microscopy. In the adults, the morphology of ossicles and gastric teeth of D. sayi is very similar to the related species Rhithropanopeus harrisii. The foregut of first zoea (ZI) presented a functional cardio-pyloric valve while the filter press was lacking. The filter press was observed in the pyloric chamber from ZII. The most significant changes in morphology take place after metamorphosis from ZIV to megalopa, including the occurrence of the gastric mill. The organization and morphology of many megalopal foregut ossicles are recognizable in the adult phase, although the morphology of the gastric teeth differs from the morphology of adults. A correlation of gastric mill structures with food preferences and their contribution to the phylogeny are briefly discussed.     

Biogeography of subterranean decapods in North and Central America and the Caribbean region (Caridea,Astacidea, Brachyura)

A significant number of decapod crustaceans (81 troglobites and 58 other cavernicoles) has been described from various subterranean waters in North and Central America (United States south to Costa Rica) and from the islands in the western north Atlantic and the Caribbean Sea, posing puzzling questions concerning their evolution and biogeography. Of these troglobitic species, 36 are shrimps (1 procarid, 11 atyids, 2 agostocarids, 15 palaemonids, 2 alpheids, 5 hippolytids), 35 are cambarid crayfishes, and 10 are crabs (1 grapsid, 7 pseudothelphusids, 2 trichodactylids). They are known to occur in caves, springs, cenotes, blue holes, anchialine environments, and various crevicular habitats in localized areas throughout the region. Many, if not the majority, of the troglobites appear to have arisen independently from epigean progenitors rather than sharing common subterranean precursors.     

Correlations between chaetodontid fishes and coral communities of the Gulf of Aqaba (Red Sea)

Synopsis The relationships existing between the chaetodontid fishes and the surrounding coral communities were investigated in the Gulf of Aqaba. Quantitative data were analysed by a correspondence and a cluster analysis. The results demonstrated a similarity in the spatial distribution of both communities. Significant correlations were found between the density of chaetodontid fishes and the diversity of the coral community as well as the substratum coverage by the coral colonies. The density of exclusive coral browsers was also correlated to the abundance of branching colonies. Among the different genera of branching corals, correlations were significant only for the genusAcropora. These results suggested the existence of strong links between coral and chaetodontid fish assemblages.     

New species and new records of pinnotherid crabs (Crustacea: Decapoda: Brachyura) from the Yellow Sea

Five species belonging to two genera of pinnotherid crabs are reported from the Yellow Sea. Two are new to science, namely: Sakaina glabra sp. nov. and Sakaina granulata sp. nov.; while three species are recorded for the first time from China: Pinnaxodes majorOrtmann, 1894, Pinnaxodes mutuensisSakai, 1939, and Sakaina japonicaSerène, 1964. Sakaina glabra can be easily distinguished from congeners by lacking a thick rim of pubescence along the anterolateral margin of the carapace. Sakaina granulata closely resembles S. asiatica (Sakai, 1933), but differs mainly in having a row of granule-like denticles along the anterolateral margin of its carapace.     

Factors Controlling the Planktonic Community in the Shallow-Lake of Créteil,France

A two-year study on the seasonal pattern of plankton was performed in a shallow, polymictic, eutrophic lake. Autogenic successions typical of stratified lakes did not occur. Only small edible algae developed throughout the two years and Cyanophyceae rarely appeared, and did not bloom. Algal sequences depended on wind-induced periodic mixing of water. The seasonal pattern of Rotifers varied greatly from year to year and some species depended on sporadic increases of algae. In contrast, the seasonal pattern of microcrustacea was more regular. Resting stages played an important role in zooplankton development in this variable environment. Successful fish reproduction depended a great deal on climatic conditions. The changes from year to year in predation pressure from young fish appeared to be an important factor controlling zoo-plankton structure.     

Biological Aspects of Medorippe lanata (Linnaeus, 1767) (Brachyura: Dorippidae) from the Eastern Ligurian Sea (Western Mediterranean)

The aim of the present study is to investigate the demographic structure and to identify some aspects of the biology of an exploited population of Medorippe lanata (Brachyura: Dorippidae) in the eastern Ligurian Sea, western Mediterranean. 1364 specimens (639 males and 725 females) of M. lanata were collected on a monthly basis from January to December 2001, in a wide area of the eastern Ligurian Sea usually exploited by the Viareggio ‘rapido’ trawl fleet. M. lanata represented an important fraction of the discard, both in weight and in number of individuals. Maximum abundance of this species occurred in late summer-early autumn (up to 3369 ind. km⁻² and 50.6 kg km⁻² in August). The overall females:males sex-ratio was 1.13:1, while the monthly sex-ratio did not differ statistically from 1:1 in all months, except in September and October, when females significantly outnumbered males. The sampled population was composed of two cohorts from November to April. Sizes ranged from 10 to 29 mm carapace length (CL) for females and from 9 to 29 mm CL for males. The von Bertalanffy growth curve, computed for both sexes, gave a higher growth rate in males than in females. Recently moulted males and females were observed throughout the year, except in summer, when the highest number of ovigerous females was present. Females with external eggs were collected from March to November, with peaks in August and September. The monthly evolution of the ovarian maturity stages showed no clear temporal trend. At 21 mm CL, 50% of females were ovigerous or showed macroscopically mature ovaries. According to the dimorphism in chelae size, the presence of adult males (post-puberty stage) was observed all year round, from 18 to 29 mm CL, without evident temporal trends.     

Notes on the distribution and biology of the deep-sea crab Bathynectes maravigna (Brachyura: Portunidae) in the Mediterranean Sea

Data on the distribution and biology of the deep-sea portunid crab Bathynectes maravigna are reported for the Mediterranean Sea, based on several fisheries research surveys. Densities are low and, therefore, biological data are scarce. In the western Mediterranean, the species is much commoner in Alborán Sea than in the Catalano-Balearic Sea. Occurrences are also scarce in the southern Adriatic and northwestern Thyrrenian Sea, as well as in the Ionian Sea. The Alborán Sea and the seas surrounding the southern Italian peninsula are the areas where densities are the highest. The occurrence depth range was found to be 245–786 m, but most of the occurrences took place deeper than 500 m. Sizes ranged between 9 and 51 mm carapace length in males and between 12 and 51 mm in females. Ovigerous females have been only reported in October–December and March–May. Eighty three percent of both males and females are right-handed. Sexual dimorphism was present in cheliped length with males having longer chelae than females. The species appears to be much commoner in those areas where Atlantic influence is the highest.     

Phylogeny, biogeography and a new taxonomy for the Gecarcinucoidea Rathbun, 1904 (Decapoda: Brachyura)

Phylogenetic relationships of gecarcinucoid freshwater crabs were investigated, based on morphology of the male second gonopod. In addition, a comparison of sequences from the mitochondrial large subunit rRNA gene helped to resolve the phylogeny of this group and relationships to other Old World freshwater crabs. As a result, we recognise two sister groups within the Gecarcinucoidea, the African Deckeniidae and the Asian Gecarcinucidae. Deckeniidae includes three monophyletic clades, the Deckeniinae in East Africa and on the Seychelles, the West African Globonautinae and the Malagasy Hydrothelphusinae. Gecarcinucidae comprises two sister groups, the Gecarcinucinae with representatives in Sri Lanka, India and southeast Asia, and the Parathelphusinae in India, southeast Asia, the Sundaic Islands and Australia. Interpretation of our phylogenetic results leads us to propose a new biogeographic hypothesis for the Gecarcinucoidea. Most likely, the gecarcinucoid freshwater crabs have an African origin; their distribution can be explained by successive events of dispersal. This model can be correlated with palaeogeographical and palaeoclimatological data for the Cenozoic, suggesting a gecarcinucoid dispersal to Asia via the “Lemurian Stepping-Stones”, a chain of islands in the West-Indian Ocean that were emergent in times of low sea levels during the Oligocene.     

Heterochely and handedness in the shore crab Carcinus maenas (L.) (Crustacea: Brachyura)

A large sample ofadult male Carcinus maenas was 79% right-handed and 21% left-handed. A separate sample of 207 intact adult males was divided into left-handed and right-handed crabs and four measurements were taken from all major and minor chelae. Correlation and regression analyses against carapace width on log-transformed data showed that major chelae of right-handed crabs grow proportionately higher with increasing size and the ideal mechanical advantage increases; concurrently, the fingers of the minor chelae grow proportionately longer. The data for left-handed crabs showed greater variability, especially for minor chelae, providing evidence for the concept that left-handedness arises by reversal of handedness following loss of the major chela from the right-hand side. Records of handedness in large samples of non-ocypodid heterochelous brachyuran crabs show a preponderance of right-handedness.     

On the morphology of the central nervous system in larval stages ofCarcinus maenas L. (Decapoda,Brachyura)

We investigated the morphology of the central nervous system throughout the larval development ofCarcinus maenas. For that purpose single larvae were reared in the laboratory from hatching through metamorphosis. Complete series of whole mout semithin sections were obtained from individuals of all successive larval stages and analysed with a light microscope. Morphological feature and spatial arrangement of discernable neural cell clusters, fibre tracts and neuropile are described and compared with the adult pattern. We found that most of the morphological features characterizing the adult nervous system are already present in the zoea-1. Nevertheless, there are marked differences with respect to the arrangement of nerve cell bodies, organization of cerebral neuropile, and disposition of ganglia in the ventral nerve cord. It appears that complexity of the central nervous neuropile is selectively altered during postmetamorphotic development, probably reflecting adaptive changes of sensory-motor integration in response to behavioural maturation. In contrast, during larval development there was little change in the overall structural organization of the central nervous system despite some considerable growth. However, the transition from zoea-4 to megalopa brings about multiple fundamental changes in larval morphology and behavioural pattern. Since central nervous integration should properly adapt to the altered behavioural repertoire of the megalopa, it seems necessary to ask in which respect synaptic rearrangement might characterize development of the central nervous system.     

Population Dynamics,Growth and Productivity of Abra ovata (Mollusca,Bivalvia) in the Evros Delta (North Aegean Sea)

Monthly samples of Abra ovata were collected during February 1983-January 1984 in the Evros Delta (N. Aegean Sea). Population density (mean annual value = 2407.5 ind · m⁻²) was characterized by seasonal variation. An analysis of the length frequency distributions shows that one annual recruitment of juveniles (> 2 mm) occurred in October-January; and also that, throughout the year, two age groups existed in the population. One growth ring was formed on the shells of the oldest age group during July-August. Mean growth in shell length can be described by Bertalanffy function. A positive correlation existed between shell length and decalcified dry weight. Secondary production in A. ovata, calculated by the instantaneous growth method, showed a mean biomass of 29.221 g dry weight m⁻²yr⁻¹, a productivity of 17.086 g dry weight m⁻²yr⁻¹ and an annual turnover ratio of 0.59.     

A comparative study of zoeal morphology in the genus Liocarcinus (Crustacea: Brachyura: Portunidae)

The zoeal morphology of Liocarcinus arcuatus, L. corrugatus and L. depurator is described from laboratory reared material and the larvae of L. holsatus, L. puber, L. pusillus and L. marmoreus are reexamined from specimens deposited in the BM (NH). No single meristic character was found that would adequately separate the seven described species of Liocarcinus. Characters that may facilitate separation of later zoeal stages only are listed.     

Maternal control of calcium concentration in the larval nursery of the bromeliad crab,Metopaulias depressus (Grapsidae)

It has been proposed that parental care in invertebrates controls for physicochemical conditions. This suggests that the parent detects deviations from optimal conditions and responds by correcting the deviation for the benefit of the offspring. In the bromeliad crab, Metopaulias depressus, mothers accumulate snail shells (CaCO₃) in the leaf axil containing the larvae, which results in an increase in Ca²⁺ and pH, both beneficial to the young. I tested the hypotheses that mothers: (i) control for calcium levels and only collect shells when calcium is too low; (ii) always collect shells, independent of the calcium content of the nursery; (iii) collect shells if encountered, but increase their activity if the calcium content becomes critical. In a field experiment, I reduced the initial calcium concentration in the nurseries of one group and increased it in another group. In both groups mothers collected shells. However, they collected significantly more shell mass per day in the group with reduced levels than in the group with high levels. The results support hypothesis (iii) and unequivocally demonstrate maternal control of physicochemical conditions in the nursery. This is the first proof that crustacean mothers actively regulate abiotic conditions for their offspring.     

Agonistic Behaviour and Development of Territoriality During Ontogeny of the Sea Anemone Dwelling Crab Allopetrolisthes Spinifrons (H. Milne Edwards, 1837) (Decapoda: Anomura: Porcellanidae)

Allopetrolisthes spinifrons is an ectosymbiotic crab of the sea anemone Phymactis clematis. As a consequence of low host abundance, these represent a scarce and limited resource for the crab. Additionally, the relatively small size of the sea anemone host suggests that few symbiotic crabs can cohabit on one host individual, forcing crabs to adopt a territorial behaviour. In order to examine the potential presence and ontogenetic development of territoriality, the agonistic behaviour between crabs of various ontogenetic stages (adults, juveniles, and recruits) was studied in the laboratory. Laboratory experiments demonstrated that adult or juvenile crabs aggressively defended their sea anemone hosts against adult or juvenile intruders, respectively, but both adult and juvenile crabs tolerated recruits. Adult crabs behaved indifferently towards juvenile crabs, sometimes tolerating them, sometimes expelling them. Recruits never showed agonistic behaviour among themselves. The agonistic interactions observed in the laboratory and the uniform population distribution pattern on sea anemones recently described for A. spinifrons indicate that this species exhibits territorial behaviour, which develops during ontogeny. Territoriality in this species and other symbiotic decapods may function as a density-dependent mechanism of population regulation, being mediated by the availability of hosts. Resource monopolisation behaviours may be common among other symbiotic and free-living marine invertebrates inhabiting discrete habitats that represent a limiting resource.