Carapace movements associated with ventilation and irrigation of the branchial chambers in the semaphore crab,Heloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

Summary Carapace movements in crabs are briefly reviewed. While on land and recirculating branchial water, the Australian semaphore crab Heloecius cordiformis (Decapoda: Ocypodidae), a semi-terrestrial air-breathing mangrove crab, sequentially depresses and elevates its carapace relative to its thorax (0.5–1 mm excursion) in a regular pump-like manner. In quiescent crabs each carapace-pumping cycle lasts about 4 s; carapace depression takes 3 s and elevation 1 s. Carapace movements are brought about by pressures generated within the branchial chambers by the scaphognathites, probably in combination with carapace muscles. Carapace movements are associated with bilaterally synchronised scaphognathite activity. Unilateral scaphognathite activity was not observed. During normal forward recirculation of branchial water the scaphognathites beat at about 1.5 Hz (slow-forward pumping) and the lungs (epibranchial chambers) are not ventilated. In Heloecius, the lungs are not physically separated from the gills below by an anatomical barrier. Lung ventilation is accomplished during the following sequence of events: the carapace is lowered and the scaphognathites pump in a fast-forward mode at about 2.8 Hz. This activity preferentially pumps air out of the lungs and generates suction within the branchial chambers (4–10 cm H₂O below ambient) which draws water from external body surfaces into the hypobranchial space below and around the gills. At the end of the carapace's downward travel the scaphognathites switch from fast-forward to fastreverse beating at about 4 Hz. This pumps air into the lungs and the carapace elevates. As a result, during carapace elevation the water which had previously been drawn into the branchial chambers by fast-forward pumping activity is released and flows out between the legs and into the abdominosternal cavity. When the carapace reaches its original resting or up position the scaphognathites switch from fast-reverse to slowforward beating to re-establish water recirculation through the branchial chambers. This cycle is subsequently repeated. In stationary crabs, there are 2 carapace-pumping cycles per minute, increasing to 14 per minute in active crabs (walking). When water is absent, the lungs are preferentially ventilated by slow-reverse scaphognathite pumping activity. Carapace movements do not occur in the absence of branchial water. Carapace pumping is thought to provide a mechanism which permits the scaphognathites to ventilate the lungs in the presence of recirculating branchial water, without this water interfering with lung ventilation or being lost to the environment.Abbreviations FF, FR, SF, SR fast-forward, fast-reverse, slowforward, slow-reverse scaphognathite pumping - MEA Milne Edwards aperture     

Respiratory behaviour,oxygen consumption and relative dependence on aerial respiration in the African lungfish (Protopterus annectens,owen) and an air-breathing teleost (Clarias lazera,C.).

The relative dependence on branchial and pulmonary organs was studied in the African lungfish P. annectens and in the catfish Clarias lazera. The frequency of pulmonary ventilation varied, in the normal state, with the activity and age of the fish and followed a circadian rhythm. Small specimens of both species exhibited a higher branchial ventilatory rate than older specimens and depended largely on aquatic O₂ uptake (over 85% and 90% in Clarias and Protopterus respectively). The dependence on aerial respiration appeared to develop gradually with age in Clarias but occurred over a limited age-range (200–300 g) in Protopterus. In mature fish (over 400 g), pulmonary respiration constituted 50–60% of the total in Clarias and 80–85% in Protopterus. Partitioning of O₂ uptake between air and water depended on the O₂ content of the water and that of O₂ and CO₂ in the pulmonary organs. Protopterus and Clarias surfaced for air when the O₂ content of the respiratory organs was reduced to 90% and 85% (of that immediately following an air-breath) respectively. An increase in the pulmonary O₂ content lengthened the apnoeic period and reduced pulmonary respiration more markedly in Protopterus than in Clarias whereas an increase of that of CO₂ produced the reverse effects.     

Carapace movements aid air breathing in the semaphore crab,Heloecius cordiformis (Decapoda: Brachyura: Ocypodidae)

Summary While on land and recirculating branchial water the Australian semaphore crab Heloecius cordiformis (Decapoda: Ocypodidae), a semi-terrestrial airbreathing mangrove crab, sequentially depresses and elevates its carapace in a regular pump-like manner. The functional role of these carapace movements in aerial oxygen consumption is investigated. Carapace immobilisation (reversible and non-injurious) did not appear to affect branchial water circulation. In dry crabs (branchial water removed) carapace immobilisation had no effect on the rate of oxygen consumption (VO₂), heart rate or whole-body lactate (WBL) levels. In wet crabs (with branchial water) carapace immobilisation caused VO₂ to drop by 38% from 81 to 46 l O₂ · g^-1 · h^-1, heart rate to decline by 32%, from 2.5 to 1.7 Hz, and WBL levels to increase over 2.5-fold, from 0.27 to 0.67 mg · g^-1, after 3 h of carapace immobilisation. The (VO₂) of carapace-immobilised crabs with branchial water was similar to lung-occluded crabs with branchial water. Severe hypoxia induced physiological responses similar to those of carapace-immobilised crabs with branchial water. After 3 h of severe hypoxia, heart rate had declined by 80%, from 2.2 to 0.43 Hz, and the incidence of carapace pumping slowed by 85%, from 2.4 to 0.37 cycles · min^-1. It is concluded that in the absence of carapace movements branchial water in some way inteferes with lung ventilation. Under normal circumstances water circulation and lung ventilation are mutually exclusive processes (due to their singular dependence on the scaphognathites), yet in Heloecius these processes must be carried out simultaneously. Carapace movements may alleviate this conflict.Abbreviations FF, FR, SF, SR fast-forward, fast-reverse, slow-forward, slow-reverse scaphognathite pumping - MEA Milne Edwards aperture - VO₂ rate of oxygen consumption - WBL whole-body lactate     

Prevalence of bacteria in the spermathecae of female snow crab, Chionoecetes opilio (Brachyura: Majidae)

The spermathecal contents of primiparous, multiparous and barren female snow crabs were observed by a scanning electron microscope. Bacteria colonies were observed with a significantly higher frequency in the spermathecae of primiparous and old barren females. Bacteria infect the spermathecae and destroy the spermatophores and spermatozoa inside. These observations suggest that bacteria in the spermathecae do not exclude opportunistic microbes by modifying pH of the medium as suggested in the literature. The prevalence of bacteria in primiparous and old barren females suggests that they infect individuals with a weak anti-microbial protection. The absence of bacteria in the highly acidic seminal fluid derived from males upon copulation suggests that it may provide anti-microbial protection. Bacteria do not seem to be able to survive in an anaerobic environment for a long period.     

Salinity tolerance and osmoregulation in the freshwater crab,Barytelphusa guerini (Milne Edwards)

The freshwater crab, Barytelphusa guerini, tolerates gradual transfer to higher salinities upto seawater, regulating its blood chlorides fairly well. Its metabolic rate shows a minimum in 50% seawater and increases on either side, during dilution as well as during concentration of the medium. However, this increase is more pronounced in hypotonic than in hypertonic media.     

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.     

Bioaccumulation of manganese in selected tissues of the freshwater crab,Potamonautes warreni (Calman), from industrial and mine-polluted freshwater ecosystems

Manganese concentrations in water and sediments of the Bronkhorstspruit River, Nooitgedacht Dam and especially in the Natalspruit River, did not fall within stated limits for the protection of aquatic life. Tissue manganese concentrations in Potamonautes warreni from the Natalspruit River were generally higher than those in the tissues of crabs from the other two aquatic ecosystems. The highest mean manganese tissue concentration in crabs from the three systems was detected in the carapace (587± 445 µg g^-1 wet weight). It appears that the carapace in these animals acts as a sink in which manganese can be deposited, thus also playing an important role in the detoxification of manganese in these crustaceans. No seasonal or sex-related variation was detected. Body size of the crabs, however, seems to be an important influencing factor in manganese bioaccumulation. A significant increase in carapace manganese concentrations was detected with an increase in body size. However, muscle manganese concentrations were higher in the smaller groups.The concentration factors (BF) calculated for the different tissues with respect to the water were highest in the carapace and ranged from 280.9 to 742.8. The BF with respect to the sediment was comparatively low for all the tissues (0.1 to 0.7). As the manganese concentration in the tissues reflects to some extent the degree of manganese contamination of the surrounding aquatic environment, it appears that P. warreni may be useful as a potential biomonitor of manganese pollution.     

Ventilation and respiratory metabolism in the thermophilic desert ant,Cataglyphis bicolor (Hymenoptera,Formicidae)

The discontinuous ventilation cycle of the Saharan desert ant Cataglyphis bicolor was studied over the range 15–40°C, corresponding to a >2-fold increase in the rate of CO₂ output and hence metabolic rate (Q ₁₀=2.1). Over this range, metabolic rate modulated only ventilation frequency; the volume of CO₂ emitted per ventilation remained constant. The closed-spiracle phase accounted for a small, constant proportion (ca. 14%) of total CO₂ output. In the flutter phase, the rate of CO₂ output increased at a greater than exponential rate from 29% of total CO₂ output at 15°C to 52% at 40°C. CO₂ output rate in the ventilation phase increased, and its duration decreased, exponentially with temperature. Relative to total duration of discontinuous ventilation cycle, the length of each phase was constant over the entire range of metabolic rates measured. These data are the first thorough characterization of the effect of changing metabolic rate on all phases of the discontinuous ventilation cycle of an adult insect. Clearly, C. bicolor maximizes ventilation-phase emission volumes and enhances the contribution of the flutter phase to total CO₂ release relative to other ants for which comparable data are available, and does so in ways that may reduce respiratory water loss rates.Abbreviations BM body mass - C-phase closed-spiracle phase - cVCO₂ rate of carbon dioxyde leakage during the C phase - DVC discontinuous ventilation cycle - F-phase fluttering-spiracle phase - MR metabolic rate - Q ₁₀ factorial increase in MR with 10°C increase in temperature - RQ respiratory quotient - SMR standard metabolic rate - T body temperature (°C) - VCO₂ rate of carbon dioxide output - V-phase ventilation phase - vVCO₂ rate of CO₂ emission during the V-phase - fVCO₂ rate of CO₂ emission during the F-phase - VO₂ rate of oxygen consumption     

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

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

Development of dependence on aerial respiration in Polypterus senegalus (Cuvier)

The respiratory behaviour and partitioning of O₂ uptake between air and water were investigated in Polypterus genegalus using continuous-flow and two-phase respirometers and lung gas replacement techniques P. senegalus rarely resorts to aerial respiration under normal conditions. Partitioning of O₂ consumption depends on the activity and age of fish and the availability of aquatic oxygen. Immature fish (12–22 g) cannot utilize aerial O₂ but older fish exhibit age-dependent reliance on aerial respiration in hypoxic and hypercarbic waters. Pulmonary respiration accounts for 50% of the total requirement at aquatic O₂ concentrations of about 3.5 mg · l^–1 (or CO₂ of about 5%) and fish rely exclusively on aerial respiration at O₂ concentrations of less than 2.5 mg · l^–1. Branchial respiration is initially stimulated by hypercarbia (CO₂: 0.5–0.8%) but increased hypercarbia (CO₂ – 1%) greatly depresses (by over 90%) brancial respiration and initiates (CO₂: 0.5%) and sustains pulmonary respiration.     

Molecular phylogeny of the crab genus Brachynotus (Brachyura: Varunidae) based on the 16S rRNA gene

The crab genus Brachynotus de Haan, 1833 is restricted to the intertidal and shallow subtidal of the Mediterranean and northeastern Atlantic. It is presently recognized to consist of four species, of which three (B. foresti, B. gemmellari and B. sexdentatus) are endemic to the Mediterranean. The fourth species, B. atlanticus, is found along the Atlantic coasts of northern Africa and southern Europe, but also extends into the western Mediterranean. This high level of endemism suggests that speciation within Brachynotus is strongly correlated with the geography and geology of the Mediterranean Sea. A molecular phylogeny based on the mitochondrial large subunit (16S) rRNA gene indicates that the four species of Brachynotus form a monophyletic group within Atlantic Varunidae. The DNA sequence data also show that the genus Brachynotus can be subdivided into two species groups, one comprising B. atlanticus and B. foresti, and the other one B. gemmellari and B. sexdentatus. While B. atlanticus and B. foresti are clearly genetically distinct, B. gemmellari and B. sexdentatus are identical in the studied region of the 16S rRNA gene, suggesting a recent separation or continuing gene flow.     

Organization of a phyllobranchiate gill from the green shore crab Carcinus maenas (Crustacea,Decapoda)

Summary The phyllobranchiate gills of the green shore crab Carcinus maenas have been examined histologically and ultrastructurally. Each gill lamella is bounded by a chitinous cuticle. The apical surface of the branchial epithelium contacts this cuticle, and a basal lamina segregates the epithelium from an intralamellar hemocoel. In animals acclimated to normal sea water, five epithelial cell types can be identified in the lamellae of the posterior gills: chief cells, striated cells, pillar cells, nephrocytes, and glycocytes. Chief cells are the predominant cells in the branchial epithelium. They are squamous or low cuboidal and likely play a role in respiration. Striated cells, which are probably involved in ionoregulation, are also squamous or low cuboidal. Basal folds of the striated cells contain mitochondria and interdigitate with the bodies and processes of adjacent cells. Pillar cells span the hemocoel to link the proximal and distal sides of a lamella. Nephrocytes are large, spherical cells with voluminous vacuoles. They are rimmed by foot processes or pedicels and frequently associate with the pillar cells. Glycocytes are pleomorphic cells packed with glycogen granules and multigranular rosettes. The glycocytes often mingle with the nephrocytes. Inclusion of the nephrocytes and glycocytes as members of the branchial epithelium is justified by their participation in intercellular junctions and their position internal to the epithelial basal lamina.     

Effect of long term starvation on the biochemical composition of the muscles and hepatopancreas in the crab Menippe rumphii (Fabricius) (Crustacea: Brachyura)

During the process of long term starvation both muscles and hepatopancreas are affected in their biochemical composition at different rates. During early days of starvation an increase in the muscular and hepatopancreatic glycogen is observed. At the same time a simultaneous decrease in the muscular lipid content is also observed. At a slightly later period a decrease in the hepatopancreatic lipid content is also noticed. This amount of decrease is slow in the early days of starvation and rapid in later days. Decrease in the muscular and hepatopancreatic protein content is observed when there is not an adequate quantity of hepatopancreatic lipid to be consumed.     

The potential for mitten crab Eriocheir sinensis H. Milne Edwards, 1853 (Crustacea: Brachyura) invasion of Pacific Northwest and Alaskan Estuaries

Eriocheir sinensis H. Milne Edwards, 1853 is on the list of top 100 invaders compiled by the International Union for Conservation of Nature and Natural Resources. The recent establishment of a large Chinese mitten crab population in San Francisco Bay and the potential for introductions from California, Asia and Europe pose a significant invasion potential for estuaries and rivers from California to Alaska. This alien species would place at risk the catchment areas of the Pacific Northwest including the economic and social activities that depend upon intact aquatic systems. An analysis of ecological conditions that define the mitten crab’s native and introduced range suggests that large stable estuaries with long flushing times are necessary to sustain significant populations. Most Pacific Northwest estuaries have limited salinity intrusion, estuarine habitat and short flushing times and face a reduced risk of population establishment. Large, stable estuaries, such as the Puget Sound, may support significant populations. River-dominated estuaries, such as the Columbia River, have flushing times less than the duration of larval development and wouldn’t support populations. An application of a temperature based larval development rate to near-shore and estuary sea surface temperatures suggests that estuaries in Oregon and Washington have sufficient thermal regimes to support larval development. Most estuary systems in Alaska have limited periods where water temperatures are above the mortality threshold for the larval stages and are at a low risk for the establishment of populations. A potential sea temperature rise of two degrees Celsius would permit larval development in Alaskan estuaries, where sufficient estuarine and freshwater habitats exist.     

The complete mitogenome of the hydrothermal vent crab Xenograpsus testudinatus (Decapoda, Brachyura) and comparison with brachyuran crabs

In this study, we analyzed the complete mitochondrial (mt) genome of a hydrothermal vent crab Xenograpsus testudinatus (Decapoda: Brachyura) obtained from the hydrothermal vents off Kueishantao Island, Taiwan, which extend from the deep sea Okinawa Trench. The mitogenome of X. testudinatus was 15,796 bp in length and contained the same 37 genes (e.g. 2 rRNAs, 22 tRNAs, and 13 PCGs) found in other metazoan mitogenomes. Analysis of the structural mt gene order in X. testudinatus revealed that the 13 PCGs, excluding a translocation of ND6-Cyt b cluster, were similarly ordered when compared to the pancrustacean ground pattern; however the tRNAs were severely rearranged. Phylogenetic analysis of decapod mitogenomes showed that the molecular taxonomy of the vent crab was in accordance with its morphological systematics. Together, these findings suggest that the vent crab studied here has little mitochondrial genetic variation when compared with morphologically defined conspecifics from other marine habitats.     

Regulation by the exogenous polyamine spermidine of Na,K-ATPase activity from the gills of the euryhaline swimming crab Callinectes danae (Brachyura, Portunidae)

Euryhaline crustaceans rarely hyporegulates and employ the driving force of the Na,K-ATPase, located at the basal surface of the gill epithelium, to maintain their hemolymph osmolality within a range compatible with cell function during hyper-regulation. Since polyamine levels increase during the adaptation of crustaceans to hyperosmotic media, we investigate the effect of exogenous polyamines on Na,K-ATPase activity in the posterior gills of Callinectes danae, a euryhaline swimming crab. Polyamine inhibition was dependent on cation concentration, charge and size in the following order: spermine > spermidine > putrescine. Spermidine affected K_0.5 values for Na⁺ with minor alterations in K_0.5 values for K⁺ and NH₄⁺, causing a decrease in maximal velocities under saturating Na⁺, K⁺ and NH₄⁺ concentrations. Phosphorylation measurements in the presence of 20 µM ATP revealed that the Na,K-ATPase possesses a high affinity site for this substrate. In the presence of 10 mM Na⁺, both spermidine and spermine inhibited formation of the phosphoenzyme; however, in the presence of 100 mM Na⁺, the addition of these polyamines allowed accumulation of the phosphoenzyme. The polyamines inhibited pumping activity, both by competing with Na⁺ at the Na⁺-binding site, and by inhibiting enzyme dephosphorylation. These findings suggest that polyamine-induced inhibition of Na,K-ATPase activity may be physiologically relevant during migration to fully marine environments.     

Internal anatomy and ultrastructure of the male reproductive system of the spider crab Maja brachydactyla (Decapoda: Brachyura)

The morphology and function of the male reproductive system in the spider crab Maja brachydactyla, an important commercial species, is described using light and electron microscopy. The reproductive system follows the pattern found among brachyuran with several peculiarities. The testis, known as tubular testis, consists of a single, highly coiled seminiferous tubule divided all along by an inner epithelium into germinal, transformation, and evacuation zones, each playing a different role during spermatogenesis. The vas deferens (VD) presents diverticula increasing in number and size towards the median VD, where spermatophores are stored. The inner monostratified epithelium exocytoses the materials involved in the spermatophore wall formation (named substance I and II) and spermatophore storage in the anterior and median VD, respectively. A large accessory gland is attached to the posterior VD, and its secretions are released as granules in apocrine secretion, and stored in the lumen of the diverticula as seminal fluids. A striated musculature may contribute to the formation and movement of spermatophores and seminal fluids along the VD. The ejaculatory duct (ED) shows a multilayered musculature and a nonsecretory pseudostratified epithelium, and extrudes the reproductive products towards the gonopores. A tissue attached to the ED is identified as the androgenic gland.