Interactive effects of predation pressure and insecticide (temefos) toxicity on populations of the marsh fiddler crab Uca pugnax

The interaction of predation pressure with lethal and sublethal effects of temefos (Abate^®) an organophosphorous insecticide, were studied in field populations of Uca pugnax (S.I. Smith). Changes in fiddler crab population densities were followed in open-marsh temefos-treated and untreated test plots and in treated and untreated plots which were caged over to reduce predation by marsh birds. Temefos significantly reduced the population density of U. pugnax in the open test plots but not in the caged plots. These results indicate that temefos has a primarily sublethal effect on the crabs, the effect becoming lethal only after interaction with avian predation. Evidence from the presence of a time-lag effect in the population decrease, from a calculated predation index, and from laboratory studies reported elsewhere of behavioral alteration by temefos also supports the conclusion that temefos primarily impairs the escape response of U. pugnax: this leads to increased predation and subsequently to a decreased fiddler crab population. Such studies of sublethal effects of toxicants and field studies of interactions of lethal and sublethal effects of such compounds with natural population dynamics of affected species are necessary to evaluate possible effects of toxicants on populations.     

Kinetics of uranium uptake by the crab Pachygrapsus laevimanus and the zebra winkle Austrocochlea constricta

The crab Pachygrapsus laevimanus and the zebra winkle Austrocochlea constricta were exposed for 40 d to uranium (1.5 to 10 mg l^-1) in continuous-flow sea water in separate starved and fed treatments, and the kinetics of uranium bioaccumulation were estimated from an exponential model. Starved and fed crabs took up U at a similar rate, which suggests that sea water was the major source of U to the crab; the fed crabs excreted U more rapidly than the starved crabs and this led to a lower net uptake of U by fed crabs. Fed and starved winkles took up U at similar rates and excreted it at similar rates, so the sea water was also the major source of U to winkles. Crabs took up more U than winkles; the concentration factors were 7 to 18 and 4, respectively. Uranium turnover was quite slow for both species (11 to 36 d) as it was also for winkle shells (6 d); this suggests that the rate-limiting processes which control turnover are biological (e.g. growth or tissue replacement) or physical (e.g. diffusion into the shell) rather than chemical (e.g. precipitation, adsorption or exchange). There was no effect of increasing U concentration in water on the U kinetic parameters.     

Influence of extracts of adult crabs on molting of fiddler crab megalopae (<Emphasis Type="Italic">Uca pugnax</Emphasis>)

Larvae of many benthic invertebrates metamorphose in response to habitat cues, which include the presence of adult conspecifics. Prior research showed that fiddler crab [Uca pugnax (Smith)] megalopae advance molting to the first crab stage in seawater in which conspecific adult crabs were maintained. In the present study, extracts of adult crabs were prepared and the specificity and protein content were characterized. U. pugnax megalopae were reared in the laboratory to minimize their prior exposure to potential molting cues. Then they were presented with extract solutions (in 1995, 1996, 1997, and 2000) to determine the specificity of the molting response to extracts of several crab species and the effects of the protein concentration of the extract, age of the megalopae at exposure to extract, and the duration of exposure on the molting response. Megalopae of U. pugnax molted sooner in seawater containing extract from adult conspecifics than in filtered seawater without extract or in seawater containing extract from the congener U. minax. Extract from the mud crab Dyspanopeus sayi retarded molting of U. pugnax megalopae. The stimulatory effect of U. pugnax extract on molting of megalopae was dependent on extract concentration. U. pugnax megalopae did not respond to extract until 6 days after molting, and only 1 day of exposure was sufficient to stimulate molting. The boiled extract was effective after being frozen (–15°C) for 2 years. The preparation of stable extract provides a uniform stimulus for multiple experiments examining the specificity of molting cues for brachyuran crustacean larvae and the onset of receptivity to cues.Communicated by J.P. Grassle, New Brunswick     

Toxicological and ecological implications of biotransformation enzymes in the tropical teleost Chaetodon capistratus

Hepatic cytochromes P450 (phase I monooxygenases) and glutathione transferases (phase II conjugating enzymes) were investigated in Chaetodon capistratus (Linnaeus) collected in Florida and Belize in June and December 1991, respectively. These biotransformation enzymes play major roles in the detoxification of xenobiotics by converting lipophilic chemicals to more hydrophilic, readily excretable metabolites. Content of total microsomal P450 (0.501 to 0.821 nmol mg^-1 microsomal protein) and rates of NADPH-cytochrome c (P450) reductase (270.7 to 330.2 nmol min^-1 mg^-1 microsomal protein) and glutathione transferase (2.81 to 3.12 g min^-1 mg^-1 cytosolic protein) in these fish were greater than in most untreated fish species, i.e., fish that have not been exposed to PAHs (polycyclic aromatic hydrocarbons) or PCBs (polycyclic biphenyls). Ethoxyresorufin O-deethylase (EROD) rates (0.029 to 0.171 nmol min^-1 mg^-1) were also comparable to those in most untreated marine fish. Immunoblot analysis with monoclonal antibody (MAb) 1-12-3 to scup P450E (the EROD catalyst and a teleost representative of the PAH-inducible CYP1A gene subfamily) showed slight amounts of cross-reacting protein in C. capistratus liver microsomes. Hepatic CYP1A content and EROD activity did not differ significantly between fish collected in Florida and Belize, suggesting that the two sites differed little in contamination by CYP1A inducers. Immunochemical analyses with polyclonal antibodies to scup P450B (a teleost representative of the CYP2B subfamily) and human CYP3A4 cross-reacted strongly with C. capistratus hepatic proteins. The CYP2B and CYP3A subfamilies in mammals are believed to have partially evolved in response to toxic dietary allelochemicals. C. capistratus regularly feeds on terpenoid-rich gorgonian corals, suggesting that biotransformation enzymes may be involved in the metabolism of dietary allelochemicals as well as anthropogenic xenobiotics in this species.     

Oxygen uptake and heat production in a metabolic conformer (Littorina irrorata) and a metabolic regulator (Uca pugnax)

The heat production of Littorina irrorata and Uca pugnax in air was measured with a twin calorimeter while oxygen tension was measured with a pO₂ electrode. Both L. irrorata, an oxyconformer, and U. pugnax, a metabolic regulator, showed a rapid decrease in oxygen uptake (below 1.3 mm Hg in L. irrorata and 13.4 mm Hg in U. pugnax) while heat production decreased more slowly. Consequently, during the period of minimum oxygen uptake, the oxycalorific coefficient increased from about 4.8 for both species to an average value of 8.3 in L. irrorata and 19.9 cal ml^-1 O₂ in U. pugnax, indicating the onset of anaerobic metabolism and accumulation of metabolic and products. Above their respective critical pO₂, the oxycalorific coefficient was the same as the commonly used conversion factor of 4.8 cal ml^-1 O₂. From one time interval to the next, however, the coefficient varied from 3.8 to 5.4 in L. irrorata and from 2.9 to 6.0 in U. pugnax, indicating, that the processes of oxygen consumption and heat production are more or less independent of each other and usually not in phase.     

Activity,track and speed of movement of the crab Scylla serrata in an estuary

Ultrasonic transmitters were used to track the movements of the crab Scylla serrata (Forskal) over 24 h periods in the Kowie estuary, South Africa. Laboratory experiments using infra-red time-lapse photography to record activity indicated that the transmitters did not affect duration of emergence, amount of movement or feeding. In the estuary, S. serrata was active on average for 13 h. out of 24 h, most activity was at night. The distance moved per night by continuously tracked crabs averaged 461 m, but ranged between 219 and 910 m. Most movement was slow, modal speed was 10 to 19 m h^-1. Slow movements were independent of direction of current and are assumed to be related to use of contact chemoreception for location of prey. About one-seventh of movements were faster than 70 m h^-1; these were most frequently against the current and may be related to olfactory location of food. The crabs did not occupy a distinct territory, but tended to remain in the same general area although they were capable of moving at least 800 m along the length of the estuary at night.     

Influence of simulated tidal cycles on aromatic hydrocarbon uptake and elimination by the shore crab Hemigrapsus nudus

Specimens of the intertidal crab Hemigrapsus nudus (Dana, 1851) were collected near Sitka, Alaska, in 1983, and were used in laboratory experiments to determine the influence of periodic exposure to air on the toxicity, uptake, and elimination of two aromatic hydrocarbons, toluene and naphthalene. Three tidal cycles were simulated; i.e., crabs spent 0, 33, or 66% of the time in air and the remainder of the time in toxicant or seawater solutions. Naphthalene was 10 to 20 times more toxic than toluene: naphthalene caused locomotory impairment or killed the crabs more rapidly and at concentrations lower than those of toluene. In tissues, naphthalene was accumulated faster and to higher concentrations than was toluene. Exposure to air 66% of the time reduced mortality as well as the rate at which toluene and naphthalene were accumulated and lost. Crabs exposed to air had significantly higher respiration rates than those submerged in seawater; thus, reduced rates of accumulation and loss of hydrocarbons were not caused by general metabolic quiescence. Exposure to air reduced the time the crabs were in contact with water-borne hydrocarbons and, therefore, reduced toxicity of the hydrocarbons. Once crabs had accumulated the toxicants, however, elimination was hindered by exposure to air, even for the somewhat volatile compound toluene.     

Aspects of population ecology in two populations of fiddler crabs, <Emphasis Type="Italic">Uca pugnax</Emphasis>

Fiddler crabs, Uca pugnax, were collected from a highly contaminated site and a relatively clean site, both in New Jersey to determine if and how environments with varying levels of pollutants may impact aspects of population biology including individual size, morphology (major cheliped size), population density, fecundity, recruitment and survivorship of early benthic phases. Crabs from the highly contaminated site were significantly larger in size, but had lower population density, lower recruitment, reduced reproductive season and lower survivorship of early benthic phases. Our study suggests that contamination may play a role in population ecology of U. pugnax. This study also determined that the reproductive season for U. pugnax in New Jersey is much longer than reported in the literature and could potentially be impacted by global climate change.     

Respiration and excretion rates of Calanus glacialis in arctic waters of the Barents Sea

The respiration and excretion rates of Calanus glacialis (Jaschnov) Copepodite Stages III, IV, V, and adult females from the drift-ice area east of Svalbard (Barents Sea) were measured in shipboard experiments during the period from 27 May to 13 June, 1983. The phytoplankton biomass and abundance varied considerably between localities, but these variations were not generally reflected in the respiration and excretion rates of the copepod. The respiration and excretion rates of C. glacialis at the ambient temperature of-1.8°C (average respiration rates of 0.95, 0.73, 0.57, and 0.60 l O₂ mg^-1 dry wt h^-1 for Copepodite Stage III, IV, V, and adult females, respectively) were similar to those previously reported for other large-sized copepods from cold or temperate areas. Average respiration and excretion rates tended to decrease with incubation time or time after capture. Measurements on ten occasions within a period of 27 h after capture revealed excretion rates of ammonium ranging between 2.9 and 16.8 for C III, 3.7 and 21.1 for C IV, 1.3 and 28.4 for C V, and 1.6 and 18.7 for adult females, all expressed as nmol mg^-1 dry wt h^-1. In all experiments, excretion rates of inorganic phosphate varied between 0.7 and 1.5 (C III), 0.5 and 1.1 (C IV), 0.2 and 0.8 (C V), and 0.3 and 1.0 (adult females) nmol mg^-1 dry wt h^-1. Ratios of O:N, O:P, and N:P indicated that much of the metabolic energy was derived from catabolism of proteins. Comparison of the turnover rate of carbon and nitrogen showed, however, that nitrogen turnover was between 2.6 and 8.9 times higher than that of carbon. This may indicate that the copepods deaminate ingested protein, with the carbon skeleton of the amino acids subsequently being used in the synthesis of lipid compounds, possibly wax esters.     

Natural food,foregut clearance-rate and activity of the crab Scylla serrata

The natural diet, rate of foregut clearance and diurnal activity of the crab Scylla serrata were determined. The gut volume is related to size of crab as gut volume (ml)=0.07e^0.033x , where x=carapace width in millimetres. Fifty per cent of crabs collected in Australia and South Africa contained molluscan remains and about 21% contained crustacean remains — chiefly grapsid crabs. Fish remains were rarely found, and it was concluded that S. serrata does not normally catch mobile forms such as fish and penaeid prawns. Gut clearance of organic tissue was rapid and almost complete after 12 h. Fish bone was retained for a mean time of 2 to 3 days, and shell for 5 to 6 days. Time-lapse photography, using infra-red light, was used to record activity. Visible light flashes reduced activity. S. serrata remained buried during the day, emerging at sunset to spend the night feeding, which occurred intermittently even when unlimited food was available. If no food was present the amount of time spent on the substrate surface was halved.     

Leaf consumption by <Emphasis Type="Italic">Sesarma plicata</Emphasis> in a mangrove forest at Jiulongjiang Estuary,China

The feeding ecology of Sesarma plicata (Grapsidae: Sesarminae), the most abundant crab species in a mangrove forest dominated by Kandelia candel at Jiulongjiang Estuary, China, was investigated through field and laboratory experiments. Feeding preference and consumption rates were determined on mature, senescent and decomposed leaves of Kandelia candel, Bruguiera gymnorrhiza and Aegiceras corniculatum. In the laboratory, S. plicata preferred leaves of K. candel over those of B. gymnorrhiza and A. corniculatum, and consumed significantly more decomposed leaves than mature and senescent ones, irrespective of crab size. Field experiments with limited power failed to reveal detectable species preferences despite more consumption of K. candel, but decomposed leaves of each species were again preferred. Leaf characteristics associated with preference changed with plant species and leaf state. Low tannins and high water content characterized the preference for decomposed state of leaves. Species preference was significantly and negatively related to crude fibers and C:N ratios for mature leaves, and crude fiber for senescent leaves, but significantly and positively related to water content for decomposed leaves. Leaf consumption rates averaged for all leaf categories from laboratory no-choice feeding experiments were 0.101, 0.055 and 0.017 gDW ind⁻¹ d⁻¹ for large, medium and small crabs, respectively. In this forest, mean density of S. plicata was 20.5 ind m⁻² as assessed by a manual catching method. Leaf litter removal rate during neap tides by sesarmid crabs was about 1.33 gDW m⁻² d⁻¹ in April 2006. The leaves removed by crabs were grazed on the sediment surface or taken into crab burrows, shredded and stored before being eaten.     

Biodeposition by salt-marsh invertebrates

The importance of salt-marsh invertebrate biodeposits in biogeochemical cycles is discussed. Yearly biodeposition rates of nitrogen, phosphorus, potassium, magnesium, maganese, copper, zinc, and molybdenum are reported for 5 invertebrate species. The 5 species (Littorina irrorata, Arcatula demissa, Polymesoda caroliniana, Uca pugnax and U. pugilator) deposit 1709 g dry weight m² year^-1, of which 455 g m² year^-1 is organic material. Calculations suggest that the invertebrates are capable of processing 53% of the marsh's yearly production, but the actual amount utilized is probably substantially less than this.Contribution No. 716 from the Virginia Institute of Marine Science, and Contribution No. 295 from the University of Georgia Marine Institute.     

Nitrogenous excretion by the sediment-living bivalve Nucula tenuis from the Oslofjord,Norway

The resting rate of ammonia excretion for the sediment living bivalve Nucula tenuis (Montagu) was found to be 38.8 gN mg^-1 dw h^-1×10^-4 in August and November 1985 in the Oslofjord. The excretion rate of experimental individuals was 37% higher when placed in artificial glass bead sediment. The regression between dry weight and excretion was logN excretion=1.338+1.192 log x, where excretion is gN individual^-1 h^-1×10^-4 and log x=mg dry weight.     

Fate of petroleum hydrocarbons taken up from food and water by the blue crab Callinectes sapidus

Radiolabeled paraffinic and aromatic hydrocarbons, including benzopyrene, fluorene, naphthalene, methylnaphthalene, methylcholanthrene, hexadecane, heptadecane and dotriacontane, were taken up from food and water by the blue crab Callinectes sapidus. In 2 days, approximately 10% of the benzopyrene and fluorene were taken up from the water when their concentrations were 2.5 and 30 g/l, respectively. When given food with radiolabeled hydrocarbons, 2 to 10% of the hydrocarbons were assimilated by the carbs, with the remainder excreted. After uptake of hydrocarbon from water or food, a major pathway for the elimination of hydrocarbon and metabolites was through fecal material. All hydrocarbons used in the study were metabolized, with similar rates for both paraffinic and aromatic hydrocarbons. More than 50% of the radioactivity assimilated by the crabs was in the hepatopancreas, suggesting that the hepatopancreas was the site of hydrocarbon metabolism. Twenty-five days aftex exposure to radiolabeled hydrocarbons, radioactivity was found only in the hepatopancreas. The hepatopancreas contained highly polar hydrocarbon metabolites, including dihydroxy-compounds and their conjugates, while blood contained both monohydroxy-and dihydroxy-compounds. No evidence was found of storage of hydrocarbons by any of the crab tissues.     

Responses of estuarine crab megalopae to pressure,salinity and light: Implications for flood-tide transport

The megalopal larval stage of many estuarine brachyuran crabs appears to return to adult habitats by undergoing rhythmic vertical migrations which result in saltatory up-estuary transport on flood tides. Larval ascent into the water column during rising tides may be cued by changing hydrologic variables. To test this hypothesis, we investigated the responses of field-caught megalopae of the blue crab Callinectes sapidus and the fiddler crab Uca spp. to constant rates of pressure and salinity change under laboratory conditions. For both genera, pressure changes resulted in increased movement (barokinesis) and upward migration in the test chamber, with C. sapidus megalopae having a lower response threshold (2.8×10^-2 mbar s^-1) than Uca spp. larvae (5×10^-2 mbar s^-1). Similarly, larvae ascended in response to increasing salinity, with C. sapidus larvae being more sensitive. Larvae were negatively phototactic and failed to respond to pressure increases at light levels above 1.0×10¹⁵ and 1.0×10¹³ photons m^-2 s^-1 for C. sapidus and Uca spp. megalopae, respectively. Such responses are thought to explain the low abundances of larvae in the water column during daytime flood tides. Nevertheless, threshold sensitivities to increasing pressure for both genera were above levels experienced during floodtide conditions in the field. Similarly, it is unlikely that increasing salinity is sufficient to induce ascent in Uca spp. postlarvae. However, rates of salinity increase during midflood tide typically reach levels necessary to induce an ascent in C. sapidus megalopae. These results are consistent with the hypothesis that fiddler crab megalopae utilize an endogenous activity rhythm for flood-tide transport, while blue crab megalopae rely upon external cues, especially salinity changes, to time their sojourns in the water column.     

Shell-size selection by intertidal sympatric hermit crabs

This study evaluated selection for shell size by three species of tropical intertidal hermit crabs, Clibanarius antillensis, C. sclopetarius, and C. vittatus, from species of shells which are frequently used in nature. Crab size and weight were strongly and significantly related to all measured parameters of the selected shells. The strength of these relationships (r² values) depended neither on the crab nor on the shell variables taken into account. The relationships between crab size and the dimensions of the selected shells showed higher r² values than the corresponding relationships with the shells that the crabs had occupied when they were collected (0.482–0.903 in comparison to 0.091–0.652, respectively), indicating that the crabs were occupying sub-optimal shells in nature. Negative allometry was frequently found in the relationships between crab and shell variables, indicating that large crabs select and use proportionally lighter shells than do small crabs. This negative allometry was stronger for the shells used in nature (except for C. antillensis), i.e. larger crabs tended to select heavier shells in the laboratory than in nature. Different allometric relationships were also recorded among the dimensions of shells used in nature and those selected by the hermit crabs in free-access experiments: as shell length increased, the selected shells were heavier and had larger apertures than the shells used in nature. The relationships between crab size and the length and weight of the selected shells did not depend on the species of crab or species of shell, but only on crab size. Therefore, analyses using these variables can be performed without taking the species of crab or shell into account, i.e. data from different crab or shell species can be pooled. The influence of crab and/or shell species was recorded only in the models fitted for aperture length and width, variables which were more related to shell architecture than did shell length or weight. In contrast, if crab weight is used as an independent variable, different crab or shell species can be analyzed together independently of the particular shell parameter. This indicates that crab weight may be less susceptible than crab shield length to shell morphological constraints. Finally, the results indicate that the preferred size of a given shell type chosen by a given hermit crab will depend more on crab size or weight, than on the crab or shell species under consideration, i.e. crab shell-size relationships are not species specific.Communicated by P.W. Sammarco, Chauvin     

Feeding responses of the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta): Influence of sexual maturity

Feeding responses of the tentaculate depositfeeding polychaeteEupolymnia nebulosa (Montagu) were studied by measuring rates of uptake of three different¹⁴C-labelled diatoms (unialgal cultures ofNavicula incerta Grunow,Nitzschia acicularis Wm Smith, andNitzschia sp.). Worms used during this study were collected in the harbor of Port-Vendre (Western Mediterranean) during August 1986 (immature worms) and December 1987 (mature worms). Uptake rates were affected both by the length of the experiments and by the nature of the food offered. The highest rate of uptake (2.98 10^–4 mg ashfree dry wt of algae mg^–1 dry wt of worms h^–1) was obtained during short-term experiments (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate of uptake (0.21 10^–4 mg ash-free dry wt of algae mg^–1 dry wt of worms h^–1) was also obtained withNitzschia sp., but for a long-term (48 h) experiment. There was no significant difference between rates of uptake of immature and mature worms.     

Heavy metal contamination and physiological variability in the Brazilian mangrove crabs Ucides cordatus and Callinectes danae (Crustacea: Decapoda)

 Physiological studies were made on the crabs Ucides cordatus (L.) and Callinectes danae sampled from populations living in “polluted” mangroves on the southeast littoral of Brazil. Analysis of Cu, Cd, Zn, and Fe of sediments and crab tissues showed interspecific differences in tissue concentrations, and significantly higher levels of Cu, Cd, and Zn in “polluted” populations compared to “unpolluted” crabs living in uncontaminated mangrove in the same geographical area. Individuals of both species from the polluted site showed significantly greater capacities for regulating blood osmotic concentrations at low salinity (9‰). However, U. cordatus showed a reduced hypo-regulatory ability in 34‰S. Differences in ionoregulation were also seen. “Polluted”C. danae showed significantly higher Na/ K-ATPase levels in posterior gills compared to “unpolluted” crabs. Oxygen consumption rates (M˙ _O2) were elevated in U. cordatus, but depressed in C. danae from the “polluted” population. Individuals of both species from this site showed significantly lower O:N ratios, mainly because of an increased net efflux of ammonia. Adenylate energy charge (AEC) values of muscle and hepatopancreas in “unpolluted” and “polluted” populations of both species were not significantly different. These physiological differences are discussed in relation to the known acute physiological and metabolic effects of heavy metals in crustaceans, and interpretated in the light of possible adaptive changes following long-term exposure to contamination. Received: 6 August 1999 / Accepted: 22 June 2000     

Interactions of methylmercury,cadmium, and salinity on regeneration in the fiddler crabs Uca pugilator,U. pugnax and U. minax

After multiple autotomy in the fiddler crabs Uca pugilator, U. pugnax and U. minax, both methylmercury and cadmium retard limb regeneration and ecdysis. When crabs in sea water are exposed to a mixture of both metals, the effect is increased, indicating that the two are interacting in an additive way. In 50% sea water (=15% S), the effects of cadmium are greatly intensified so that growth of limb buds is extremely slow, if it occurs at all. When methylmercury is present in the water at the same time, the severe effects of cadmium are somewhat ameliorated, indicating an antogonistic interaction of the two metals under these conditions. Adding additional calcium to the 50% sea water also decreased the severity of the cadmium effect, thus supporting the idea of a calcium-cadmium competition.     

Competitive role of calcium in sodium transport in the crab carcinus mediterraneus Acclimated to low salinities

Influence of calcium on sodium fluxes was investigated in the brackish-water crab Carcinus mediterraneus Csrn., after activation of sodium regulatory mechanisms, during longterm acclimation in diluted (15.9 S) sea water. The ²²Na outflux constants measured in whole crabs are noticeably lower (0.188 to 0.374h^-1) in diluted sea water enriched by calcium (5.8 to 10.4 mM Ca²⁺/l), than in ordinary diluted sea water (0.545 h^-1). The sodium-outflux constants in hemolymph, gills and muscle show the same trend of slower exchange of ²²Na in calcium-enriched sea water. In ordinary sea water, the total sodium-outflux rate from the hemolymph amounts to 46.31 M Na/g/h, while in calcium-enriched sea water (8.23 mM Ca²⁺/l) it is inhibited, amounting to 13.86 M Na/g/h. Sodium and potassium concentrations of intracellular muscles in diluted sea water enriched with calcium and control diluted sea water are similar. The outflux of intracellular sodium from the muscle amounts to 2.84 M Na/g/h in crabs acclimated to diluted sea water.