Studies on portunid crabs from the Eastern Pacific. II. Significance of the unusual distribution of Euphylax dovii

Euphylax dovii Stimpson (Brachyura: Portunidae: Podophthalminae), a tropical Eastern Pacific swimming crab, has distinctive morphological adaptations for pelagic existence. Crabs in collections from the open ocean had a sex ratio approximating 1:1, with no crabs bearing eggs. Samples from the continental shelf of Colombia contained thousands of females, mostly ovigerous, but no males. Egg attachment has posed a major problem in the evolution of decapod crustaceans, and the two genera of portunid crabs thus far observed cannot attach eggs unless females can bury partly in soft sediments. This suggests that mated E. dovii females must migrate into shallow shelf waters to encounter sediments necessary for spawning. The high energetic cost of swimming while carrying eggs and the presence of abundant food for larvae are factors favoring residence of females in shelf waters until hatching is complete.     

Diurnal and lunar patterns of larval recruitment of Brachyura into a mangrove estuary system in Ranong Province, Thailand

The present study followed the temporal recruitment pattern of brachyuran larvae in a mangrove tidal creek on the Andaman Sea coast of Ranong Province, Thailand, based on the assumption that the processes governing recruitment are important for the overall population dynamics of mangrove brachyuran crabs. Plankton net samples were taken on five occasions: on two new moon spring tides, one waxing moon neap tide, one full moon spring tide and one waning moon neap tide during October and November 1997. In addition collectors for larval crab megalopae were employed every 3 days through one dry season and one wet season (March–October 1998). Both the plankton net samples and collector samples revealed four major brachyuran groups in three families: Ocypodidae, Grapsidae and Portunidae. The grapsid group was further separated into two morphotypes which were identified as Metaplax and sesarmid species. Identified group mean numbers per cubic metre were ocypodids 3.0, sesarmids 0.8 and Metaplax 0.5, while portunid megalopae were very scarce (≪0.1 m⁻³). Further analysis of plankton net samples showed that when considering the parameters date, depth, current direction and the diel cycle, Metaplax and ocypodids distribute differently in the tidal and lunar cycle. Metaplax recruitment dominates on flood tides and on bottom layers, followed by middle and surface layers. Conversely, ocypodid abundance varied significantly with date only. Notably recruitment was not dependent on the diel cycle for either group. The collector samples of megalopae showed that recruitment of ocypodids, Metaplax and sesarmids occurred on full and new moon spring tides, while portunid megalopae preferred to settle on full moon spring tides. Since tidal currents were related to the lunar cycle megalopa groups are also cross-correlated with tidal amplitude, except for the portunid group. It is concluded that megalopae recruit in a similar manner to what has been found in other regions of the world, except that the abundance of ocypodids and Metaplax is not influenced by the diel cycle. Received: 14 February 2000 / Accepted: 24 November 2000     

Population structure of four species of porcellanid crabs (Decapoda: Anomura) occurring on the coast of Karachi

The population structure of 4 species of porcellanid crabs, including 2 species of Petrolisthes (P. boscii and P. rufescens) and 2 of Pachycheles (P. tomentosus and P. natalensis) was investigated. Sex ratios deviated from the expected 1:1 distribution in some size classes of the 4 species, but conformed in general to this pattern on the entire population basis. Probability curves developed from sex ratios differ markedly in shape and slope and resemble Wenner's (1972) anomalous and intermediate types. Ovigerous females of Petrolisthes boscii occurred for 9 months, of P. rufescens for 11 months; those of Pachycheles tomentosus and P. natalensis occurred throughout the year. In every size class, a considerably higher number of female Pachycheles tomentosus and P. natalensis became ovigerous than in Petrolisthes boscii and P. rufescens. Size-frequency distributions of the 4 species were unimodal, indicating that recruitment to crab populations was gradual. Juveniles of Petrolisthes boscii and P. rufescens were most abundant towards the commencement of the southwest monsoon season, in June–July; juveniles of Pachycheles tomentosus and P. natalensis were most abundant during the premonsoon month of March. Thirteen percent of Pachycheles natalensis were infected by a bopyrid parasite; the bopyrid also infected 5.2% of Petrolisthes rufescens and 0.6% of P. boscii. The latter two species were also infected by a rhizocephalan parasite, the infection rates being inverse to those by the bopyrid, i.e., 0.3 and 6.4%, respectively.     

The mechanism of crustacean salinity tolerance: Cell volume regulation by K+ and glycine effluxes

The regulation of muscle fiber K⁺ and free amino acid (FAA) concentrations during hypoosmotic stress was investigated in the moderately euryhaline crab Cancer irroratus. After 6 h of exposure to 60% ASW, muscle fiber K⁺ concentration declined from 185 mM to 140 mM. Following this, the blood glycine levels began to increase, indicating an FAA efflux from the cells. These data indicate that both muscle fiber K⁺ and FAA contribute to cell volume regulation in C. irroratus. The early release of K⁺ limits the initial rate of cell hydration. The subsequent efflux of glycine accounts for the volume regulation response of the muscle fibers. The cell volume regulatory system of C. irroratus is a coordinated use of both inorganic ions and FAA.     

Survival and chloride ion regulation of the porcelain crab Petrolisthes armatus exposed to mercury

Acute toxicity bioassays conducted at various salinities demonstrated that mercury (as mercuric chloride) at low concentrations was lethal to Petrolisthes armatus. Ninety-six hour LC₅₀ values varied from 50 to 64 parts per billion (ppb) of mercury, depending on test salinities. Lower salinities. decreased the time to death of mercuryexposed crabs. Differences in survival after 96 h due to salinity were not statistically significant. Blood chloride concentrations were regulated hyperchloride to the medium at low salinities and hypochloride at high salinities by acclimated crabs. The salinity isochloride to blood was 20 S. Transfer of crabs from 15 S to salinities ranging from 7 to 35 S resulted in new steadystate chloride levels within 12 h. Exposure to 50 ppb mercury did not alter chloride ion regulation of either acclimated crabs or crabs adjusting to new salinities.     

Morphology of grooming limbs in species of Petrolisthes and Pachycheles (Crustacea: Decapoda: Anomura: Porcellanidae): a scanning electron microscopy study

The morphology of the grooming limbs, the 5th pair of pereiopods, was studied by scanning electron microscopy in six species of porcellanid crabs, Petrolisthes cabrilloi, P. cinctipes, P. armatus, P. galathinus, Pachycheles monilifer, and Pachycheles rudis, and their function was inferred by comparison with findings from previous studies. Grooming limb morphology was almost identical among the four Petrolisthes species and differed little compared to that of the two Pachycheles species. The 5th pereiopods bore a basal tuft of mechanoreceptive setae, three different types of grooming setae armed with setules or denticles, two types of smooth sensilla for location and identification of fouling objects, and a terminal, toothed chela for picking firmly attached objects off the gills and body. The grooming limb was extremely flexible and could reach most parts of the body, including the gill chamber on the opposite side. The grooming limb morphology in Petrolisthes cabrilloi is consistent with its wellknown effectiveness in preventing parasitism by the rhizocephalan Lernaeodiscus porcellanae. Grooming setae remove recently attached cyprids, while the chela can grip and remove the much smaller, firmly attached kentrogons. Porcellanid crabs not known to host rhizocephalans, however, had grooming limbs almost identical to those of Petrolisthes cabrilloi despite their previously demonstrated failure to prevent settlement and infestation by L. porcellanae larvae. The effectiveness of P. cabrilloi in removing kentrogons, therefore, seems also to depend on behavioral adaptations whereby this species recognizes the parasite larvae as high-threat objects.     

Occurrence of a shadow response among brachyuran larvae

A closed-circuit television system was used to study phototaxis and behavioral responses upon sudden decreases in light intensity in light-adapted Stage I zoeae from 7 species of Brachyura (Panopeus herbstii, Menippe mercenaria, Uca pugilator, Callinectes sapidus, Sesarma cinereum, Pinnotheres maculatum, and Libinia emarginata). All species except one show positive phototaxis to high intensities of 500 nm light and negative phototaxis to low intensities. Upon a sudden light intensity decrease, larvae show a shadow response, which depends upon the initial and final intensities. If the initial intensity is sufficient to induce positive phototaxis, and the light is extinguished, the larvae of 6 species stop swimming and passively sink (sinking response). Reducing the intensity to an absolute level that causes negative phototaxis, induces a sinking response followed by negative phototaxis. For intensity reductions that occur in the intensity range above that for negative phototaxis, a sinking responseis observed. The intensity decrease that initiates the sinking response is independent of initial stimulus intensity and duration, is consistent between species, and is equivalent to a decrease by 0.5 OD (optical density) units. The sinking response is greatest at intensity decreases of at least 1.0 to 1.1 OD units. It is concluded that the shadow response is common in brachyuran larvae inhabiting coastal-estuarine areas.     

The metabolic demands of swimming behavior influence the evolution of skeletal muscle fiber design in the brachyuran crab family Portunidae

We investigated the influence of intracellular diffusion on muscle fiber design in several swimming and non-swimming brachyuran crabs. Species with sustained swimming behavior had aerobic dark fibers subdivided into small metabolic functional units, creating short diffusion distances necessary to support the high rates of aerobic ATP turnover associated with endurance activity. This dark fiber design was observed in all swimming species including Ovalipes ocellatus, which has apparently evolved swimming behavior independently of other Portunidae. In addition, we observed fiber and subdivision size-dependent differences in organelle distribution. Mitochondria, which rely on oxygen to function, were uniformly distributed in small fibers/subdivisions, but were clustered at the fiber periphery in larger fibers. The inverse pattern was observed for nuclei, which are not oxygen dependent, but rely on the transport of slow diffusing macromolecules. Phylogenetically independent contrast analysis revealed that these relationships were largely independent of phylogeny. Our results demonstrate cellular responses to diffusion that were necessary for the evolution of swimming and that are likely to be broadly applicable.     

Interception and dispersal of artificial food falls by scavenging fishes in the abyssal Northeast Atlantic: early-season observations prior to annual deposition of phytodetritus

The hypothesis that the behaviour of deep-sea scavenging fishes is influenced by seasonal input of organic matter from the ocean surface was investigated by observing responses to baits placed on the sea floor at 4800 m depth in the NE Atlantic (48°50′N; 16°30′W) during spring (April 1994). Data from the present study are compared with those from previous studies of the same location made in summer 1989. The first fishes to arrive at baits were the grenadier Coryphaenoides (Nematonurus) armatus and the eel Histiobranchus bathybius, after delays of 28 and 29 min, respectively; these results are not significantly different from those of summer 1989. Similarly, other indices of activity (staying time and swimming speed) showed no evidence of differences between years/seasons. However, the rate of radial dispersal of bait (0.009 m s⁻¹) by C. (N.) armatus was much slower than in all previous studies. A change in the size distribution of C. (N.) armatus to smaller individuals in spring 1994 was also evident. It is suggested that the fish on the abyssal plain may not comprise a steady-state population and that major episodic or seasonal migrations may occur. Received: 18 October 1996 / Accepted: 20 December 1996     

Electrophoretic and biometric variability in the abyssal grenadier Coryphaenoides armatus of the western North Atlantic,eastern South Pacific and eastern North Pacific Oceans

Specimens of the abyssal grenadier Coryphaenoides armatus (Hector, 1875), from the western North Atlantic and eastern North Pacific Oceans were compared electrophoretically at 27 presumptive gene loci. At 6 of the 7 polymorphic loci there were only minor differences in allelic frequencies but a nearly fixed difference was found at one locus, phosphogluconate dehydrogenase. Eastern North Pacific grenadiers typically have a narrower interorbital space, a shorter dorsal interspace, more soft rays in the 1st dorsal fin (9–10 versus 8–9) and more pelvic fin rays (21–23 versus 18–21) than grenadiers from the western North Atlantic (as well as grenadiers from the eastern South Pacific, which were included in the biometric analysis). There is an apparent disjunction in the distribution of C. armatus in the eastern Pacific at the Gulf of Panamá which coincides with the change of morphology. It is suggested that North Pacific grenadiers comprise a subspecies, C. armatus variabilis Günther, 1878, which is morphologically distinct from the subspecies C. armatus armatus (Hector, 1875) of the other areas.     

Seasonal changes of benthic megafauna in the Ría de Muros e Noia (Galicia,North-West Spain)

A community of decapod crustaceans (Brachyura) was sampled seasonally (October 1978–July 1979) from three habitats (raft, middle and beach) in the Ría de Muros e Noia (North-West Spain), with the purpose of studying spatial and temporal changes in the community and comparing with communities in the neighbouring Ría de Arousa, which supports an intense mussel (Mytilus edulis)-raft culture. The Portunidae family dominated the decapod community. Polybius henslowi, a species with pelagic stages, which enters the rías periodically in large numbers, was the dominant species at all the stations throughout the sampling period, attaining densities of 1.6 individuals m^-2 (18.7 g wet wt m^-2) in summer. Macropipus depurator and M. puber were the next most important species at the raft station, M. depurator at middle stations, and M. vernalis and Carcinus maenas at beach stations. The highest population densities were recorded in summer, due to the great abundance of Polybius henslowi at this time. When the data were reconsidered omitting p. henslowi, highest densities were in autumn and winter in the inner ría. In general, the nature of the substratum, the presence of mussel rafts, depth and salinity were the main factors determining the structure of the community. Density and biomass in the Ría de Muros e Noia (P. henslowi omitted) were lower than in the Ría de Arousa (up to six times lower in some areas). The Ría de Muros lacks the extensive number of mussel rafts present in the Ría de Arousa which constitute an important food resource for decapods.     

Origin and nature of spermatophoric mass of the spiny lobster Panulirus homarus

In the spiny lobster Panulirus homarus (Linnaeus) the spermatophoric mass consists of a highly convoluted tube, containing the sperm mass and a gelatinous matrix. The spermatophoric components have their origin in the proximal vas deferens and the completed spermatophoric mass is stored within the distal dilated part of the vas deferens. In the proximal vas deferens the inner glandular epithelial cells give rise to leaf-like typhlosoles which gradually diminish in size and finally disappear in the distal vas deferens. The secretions originating from the typhlosole form the outer gelatinous matrix of the spermatophoric mass. Histochemical observations reveal that the wall of the spermatophore tube consists of neutral mucopolysaccharide, whereas the sperm mass and the gelatinous matrix are rich in acidic mucopolysaccharides. Both polyacrylamide gel electrophoresis and agarose gel electrophoresis were used to separate the acidic mucopolysaccharide (AMPS) complexed with the proteins and the simple proteins. In addition, AMPS have been isolated following the method of Rahemtulla and Lovtrup (1974) and used for electrophoresis. In both the polyacrylamide gel electrophoresis and agarose gel electrophoresis, the extracted acid mucopolysaccharides give only a single fraction which, in comparison with the standard acid mucopolysaccharides, corresponds to chondroitin sulphate. The functional significance of the chondroitin sulphate and neutral mucopolysaccharides in the spermatophoric components of P. homarus is discussed in relation to their functional role in spermatophore hardening and protection of the delicate spermatozoa during their prolonged storage on the sternum of the female.     

Experimental examination of movement of the giant scallop,Placopecten magellanicus

We quantified movements of tagged giant scallops, Placopecten magellanicus (Gmelin, 1791), (40 to 115 mm in shell height) released in 1991 and 1992 at nine stations in Port Daniel Bay, Baie des Chaleurs, Gulf of St. Lawrence, Canada, for periods of 10 to 52 d. Both the mean distances moved per day and dispersion distances from release points were usually greater at the two stations with sand substratum, low scallop densities and high rock crab, Cancer irroratus, densities. Movement rate and dispersion of scallops from the release points at the other seven stations were similar, even though there were marked differences in substratum type (gravel, cobble or bedrock), predator density, and scallop density. Most mean dispersal directions were random, and scallops did not appear to migrate from unsuitable to suitable habitats. Although movement did reduce predation rate, scallop movement was weakly correlated with the abundance of only one predator, C. irroratus.     

Vertical distribution and feeding of the copepod<Emphasis Type="Italic"> Chiridius armatus</Emphasis>

Adult females of the copepod Chiridius armatus inhabited the lower half of a 200-m water column during winter and spring. Their distribution became gradually shallower during summer and autumn. Female C. armatus carried out diel vertical migrations, with shallower population distribution at night compared to day, although normally with few individuals captured in the upper 50 m. Enumeration of fecal pellets produced by freshly collected individuals showed that C. armatus was foraging throughout the year. Production of fecal pellets was at a minimum in February and at a maximum in April. Individuals from all depths produced fecal pellets, but the numbers seemed to decrease by depth. Numbers of fecal pellets were greatest for individuals captured at night. The omnivorous C. armatus consistently produced many more fecal pellets than the concurrently studied, strictly carnivorous copepod Paraeuchaeta norvegica. We ascribe the relatively high number of fecal pellets to the higher content of indigestible components in plant material than in animal food. The assumption of phytoplankton as a prevailing food source got qualitative support from scanning electron microscope pictures, showing that fecal pellets mainly contained phytoplankton.Communicated by L. Hagerman, Helsingør     

Metals in Cancer irroratus (Crustacea: Decapoda): Concentrations,concentration factors,discrimination factors,correlations

Nine metals were analysed in whole specimens of the decapod crustancean Cancer irroratus, and three in the exoskeleton of Carcinus maenas. In terms of concentrations (expressed as parts per millions) and concentration factors, 2 groups of metals could be distinguished in Cancer irroratus: Zn, Cu, Fe and Mn in one group, Ca, Mg, Sr, Na and K in the other group. Na excepted, all the metals had concentration factors significantly higher (P<0.01) than 1. The concentration factor of Na was significantly lower (P<0.01) than 1, demonstrating the existence in C. irroratus of a barrier to the movement of Na into the crustacean from its environment. Studies of coefficients of discrimination (D), in C. irroratus, for metals considered in pairs, showed, in each case that D was significantly different (Plt;0.01) from 1. Only the discrimination factor between, Zn and Mn was less significantly different (0.05lt;Plt;0.1) from 1. Correlations were made between the concentrations of metals and other parameters: water content, size, and ash content. Significant correlations were found, especially in the cases of the pairs Ca?Sr, Na?K, Cu?Zn, and Mn-animal size.     

Acquisition of particle processing capability in juvenile oyster <Emphasis Type="Italic">Crassostrea gigas</Emphasis>: ontogeny of gill mucocytes

Acquisition of particle processing capability in postlarval oysters depends upon the structural development of the pallial organs, as well as the development of cilia and mucocytes used (either directly or indirectly) in particle capture and transport. Mucocyte mapping was therefore used to identify mucocyte types and distributions throughout gill development in juvenile oyster Crassostrea gigas (Thunberg 1793) specimens from 2.9 mm to 2.4 cm in shell length. Three categories of gill filaments were identified: apical, lateral and principal filaments, corresponding to filament location or future location in gill plicae. Mucocyte densities were recorded per linear μm (l μm) of frontal surface, and converted to potential total volumes, using the mean volumes of each of the two major mucocyte types: acid mucopolysaccharide (AMPS)-mucocytes and mixed mucopolysaccharide (MMPS)-mucocytes. While AMPS secretions were dominant up to 1.0 cm (flat homorhabdic gill, to semi-heterorhabdic differentiation and plication), MMPS secretions increased progressively, dominating in 2.4 cm and adult specimens (fully heterorhabdic and plicated). Mucus composition, and hence mucus viscosity, thus appears to evolve in relation to the degree of enclosure of the gill frontal surfaces. Total (AMPS + MMPS) potential mucus secretion increased allometrically with juvenile growth, characterized by a sharp increase between 10 and 24 mm shell length, suggesting a marked improvement in particle processing capability. Mucocyte distributions on the gill were heterogeneous from the onset of heterorhabdic differentiation (7.5 mm): the apical filaments of the plicae contained much greater mucocyte total volumes, compared to the lateral and principal filaments. In addition to mucus composition, total potential mucus volume thus also evolved in relation to the degree of enclosure of the gill frontal surfaces. These results show that functional specialization in mucocyte distribution precedes the complete anatomical heterorhabdic differentiation. The completely functional adult gill system is thus attained in 2.4 cm juveniles. This information should be of use in understanding the dynamics of juvenile feeding, growth, and mortality, both in natural systems and in rearing operations.     

Specific effects of thiosulphate and L-lactate on hemocyanin-O2 affinity in a brachyuran hydrothermal vent crab

The hydrothermal vent crab Bythograea thermydron Williams (Brachyura: Bythograeidae) is exposed to high environmental concentrations of hydrogen sulphide. Hydrogen sulphide has previously been shown to be oxidized to a non-toxic form, thiosulphate (S₂O₃ ^2-), that accumulates in the hemolymph (to concentrations>1 mmoll^-1). Hemocyanin-oxygen (Hc-O₂) affinity was determined in dialysed, fresh or frozen hemolymph samples from B. thermydron. Although freezing is known to alter the affinity or cooperativity of some decapod crustacean hemocyanins, neither Hc-O₂ affinity nor cooperativity was significantly altered in B. thermydron hemolymph samples that had been frozen, consistent with previous findings. Oxygen affinity of B. thermydron hemocyanin was significantly increased by the presence of L-lactate. Likewise, Hc-O₂ affinity was significantly increased by the presence of 1.5 mmol S₂O₃ ^2- I^-1. The magnitude of this effect was the same as for similar concentrations of L-lactate. This specific effect of S₂O₃ ^2- does not appear to be a general property of crustacean hemocyanins, as there was no significant effect of S₂O₃ ^2- on Hc-O₂ affinity or cooperativity in dialysed hemolymph samples from the brachyuran crabs Cancer anthonyi Rathbun and C. antennarius Stimpson, or the thalassinid ghost shrimp Callianassa californiensis Dana. In the context of high environmental sulphide concentrations coupled with low P_{O 2}, and the subsequent accumulation of S₂O₃ ^2- in crab hemolymph, the increase in Hc-O₂ affinity due to thiosulphate appears to be an adaptive response in B. thermydron.     

Deep-sea bottom-living fishes at two repeat stations at 2 200 and 2 900 m in the Rockall Trough,northeastern Atlantic Ocean

Repeated sampling of deep-sea bottom-living fishes was conducted at two stations in the Rockall Trough at depths of 2 200 and 2 900 m. The 2 200 m station (M) was sampled 16 times by an Agassiz trawl between April 1978 and April 1985 and yielded 473 fish belonging to 17 species. The 2 900 m permanent station (PS) yielded 781 fish belonging to 11 species from 9 Agassiz trawls, 2 small box otter trawls, 4 semi-balloon otter trawls and 7 epibenthic sledges between March 1975 and April 1985. Macrourid fishes were numerically dominant at both stations with Coryphaenoides guentheri and Coryphaenoides (Nematonurus) armatus being the most abundant species at M and PS, respectively. In terms of biomass the morid, Antimora rostrata, was dominant at M but C. (N.) armatus was dominant at PS. The only species that showed clear evidence of a seasonal reproductive cycle was C. guentheri. Many species showed no indication of reproducing in the Rockall Trough. The diets of almost all the species are described and considered in relation to the food resources exploited. Investigations on the pelagic and benthic invertebrate faunas at these stations should reveal the extent to which the bottom-living fishes exploit the available resources.     

A genetic analysis of the pea crabs (Decapoda: Pinnotheridae) of New Zealand

Seven populations of the pea crabPinnotheres atrinicola Page were sampled from around the North Island of New Zealand from February to October 1987, and individuals were scored at 23 presumptive enzymatic loci. For a brachyuran crab,P. atrinicola revealed high levels of polymorphism and heterozygosity. Of the loci scored, phosphoglucose isomerase (Gpi) and phosphoglucomutase (Pgm) were distinguished by high variability ( =0.602 and 0.526, respectively). A clinal variation in electromorph frequency was evident at several loci, and atGpi in particular. Statistical analyses revealed that, despite relatively small genetic distance separation, a high degree of structuring was present between the geographic populations. The degree of population subdivision observed in this study is atypical of brachyuran crabs. It is suggested that the genetic differentiation observed between pea crab populations is maintained by life-history attributes and current movements which restrict gene flow between populations and, to some extent, by random genetic drift.Publication No. 41 from the Evolutionary Genetics Laboratory, University of Auckland