Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

Small-scale spatial and temporal interactions among benthic crustaceans and one fish species in the Bay of Biscay

In the summer of 2004, a video survey was carried out in the northern part of the central mud bank (Grande Vasière) of the Bay of Biscay to study the small scale relationship between the dominant crustacean megafauna Nephrops norvegicus, Munida rugosa and Goneplax rhomboides and juvenile hake (Merluccius merluccius). Using a towed body, high-resolution videos were recorded in six sampling sites. Statistical modelling using generalised additive models (GAM) revealed variations in activity patterns for two species. More N. norvegicus were observed outside their burrows at dawn and somewhat at dusk (no observations during night) while G. rhomboides was less observed in the morning. In addition, reduced spatial overlap between G. rhomboides and N. norvegicus suggested reduced competition for food but also space as both are burrowing species. The observed temporal and spatial activity patterns may contribute to regulating assemblage structure as competing species may be actively foraging at different times and locations thus reducing direct competition.     

Gender- and oxygen-related irrigation behaviour of the decapod Nephrops norvegicus

The Norway lobster Nephrops norvegicus (L.) inhabits burrows in muddy clay sediments (e.g. on the Swedish west coast), where an autumnal oxygen deficiency in the bottom water can occur. Our experiments investigated whether the irrigation of the burrows would reflect a behavioural adaptation to hypoxia, and whether any gender differences of such behaviour exist. Irrigation is performed by the pleopods which may compensate for a decreasing oxygen tension. Pleopod activity (total number of strokes per sampling time), associated with oxygen concentration and gender, was studied in N. norvegicus kept in artificial burrows resembling their natural habitat. Male and female lobsters were separately exposed to either normoxia (70% oxygen saturation) or hypoxia (30% oxygen saturation). A sexual difference in behaviour was found, where females irrigated the burrow less than males during normoxia. Females showed a significant increase of pleopod activity in hypoxia compared with normoxic conditions, which was not displayed by the males probably due to the degree of individual variation found. However, when only males were studied during progressive hypoxia (from 60 to 5% oxygen saturation), following any changes of irrigational behaviour, a significant increase of accumulated pleopod activity occurred. A major increase of pleopod activity appeared between 60 and 50% oxygen saturation, below which the activity remained high until a critical point (<10% saturation, 11 °C, 33 psu) where irrigation dropped to a level close to that of normoxic values. Activity sessions during hypoxia were longer and had a higher stroke rate than during normoxia. Received: 22 October 1997 / Accepted: 26 February 1998     

Locomotory behaviour and swimming performance of the Norway lobster,Nephrops norvegicus,in the presence of an acoustic tag

Observations have been made on the locomotory behaviour and swimming performances of the Norway lobster, Nephrops norvegicus (L.), fitted with an acoustic transmitter ventrally under the cephalothorax. The walking behaviour of adult males (44 mm carapace length) appeared to be unaffected, but the tag caused significant reductions in certain measures of tail-flip swimming performance such as swimming speed and endurance. Flume-tank experiments in low water currents suggested that the transmitter would increase hydrodynamic drag during swimming by 9 to 32%, depending on lobster size. Given the weight and dimensions of the acoustic transmitters currently available, it is considered advisable to confine acoustic tracking studies to relatively large N. norvegicus.Correspondence to: C. J. Chapman     

The water economy of the sea grass Thalassia testudinum

The osmotic values in the leaves of Thalassia testudinum Banks ex König, collected from the natural habitat, were found to be 32 atm; the relative transpiration was 100%. Electronmicroscopical analysis of the cuticle reveals many perforations. The critical sublethal water deficit lies near 65%. In dense meadows the green surface area of the plants amounts to 18.56 m²/m² bottom area.     

Life history of the sandy-beach amphipod Neohaustorius schmitzi (Crustacea: Haustoriidae)

Neohaustorius schmitzi Bousfield, the dominant macroscopic invertebrate on sandy beaches in North Carolina, USA, is strictly intertidal, and distributed from mean low tide to high tide, with an average density of 790/m². Location of greatest densities within the intertidal zone is dependent upon season and sex. An aggregated distribution is shown. N. schmitzi has 2 generations/year: a winter generation which lives about 8 months, and a summer generation which lives about 4 months. The reproductive season extends from February through October, with spring and summer peaks. Each female appears to produce only 1 brood of young. The number of young ranges from 2 to 14, depending upon the length and generation of the female.     

Between-individual variation in haemocyanin concentrations in the Norway lobster<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> following exposure to hypoxia and manganese

Environmental hypoxia and the release of reduced manganese (Mn) are often combined stress factors in marine sediments. Previous investigations have shown that hypoxia induces variable responses in the concentration of the respiratory pigment haemocyanin (Hc) of crustaceans. A recent study demonstrated that the magnitude and direction of the changes was dependent on the initial level of Hc in individuals of the Norway lobster Nephrops norvegicus. The changes also took place within 24 h rather than days as suggested by previous studies. In this present study we investigated changes of Hc in individual N. norvegicus in uncontaminated sea water after exposure to and release from realistic hypoxic stress (PO ₂=6.0 kPa). Furthermore we investigated how concomitant exposure to realistic concentrations of manganese (20 mg l ^-1) modified the responses we observed. We confirmed a between-individual variation in response and a change in Hc towards an optimum level after hypoxia, but also showed that changes that took place post-hypoxic exposure occurred over a timescale similar to that of the hypoxic response itself. Manganese exposure resulted in no significant changes in Hc whereas Mn exposure combined with hypoxia resulted in a significant decrease (15%) in all individuals independent of initial Hc. N. norvegicus was thus unable to compensate for the effects of hypoxia by synthesis of Hc after exposure to naturally occurring Mn concentrations.     

Factors affecting the distribution of rock-lobster larvae (Panulirus longipes cygnus), with reference to variability of plankton-net catches

One of the major problems in the quantitative study of the distribution of larvae of the western rock lobster Panulirus longipes cygnus George, is the high variability in planktonnet catches taken at night, even when replicate hauls are made at a single station. Some of the possible causes of this variability are considered. The density of early and late-stage phyllosoma larvae caught at the surface at night showed no direct relationship to either temperature or salinity. Early-stage phyllosoma larvae were equally well represented at the surface under all light conditions encountered at night. In contrast, the density of late-stage phyllosoma larvae at the surface was, on an average, four times greater on dark nights than on bright moonlight nights. The mean density for all phyllosoma stages decreased rapidly with depth, i.e., from 0.70/1000 m³ at the surface to 0.04/1000 m³ at 25 to 35 m, indicating that the vertical movements at night are not very extensive. The density of early-stage phyllosoma larvae in the surface at night showed no relationship to micronekton biomass (measured as wet weight). Density of late-stage phyllosoma larvae at the surface at night increased slightly, but significantly, with increase of micronekton biomass, possibly indicating similar behavioural responses to light or some other factor(s). The distribution of cast exuviae of final-stage phyllosoma larvae indicated that metamorphosis to the puerulus phase can occur more than 100 miles (161 km) from shore. Moreover, 80% of the puerulus larvae caught were within 80 miles (129 km) of the shore. Mean densities of puerulus larvae taken at and below the surface at night were 0.04/1000 m³ and 0.10/1000 m³, respectively, although total numbers were low. This evidence indicates that, after metamorphosis, the puerulus larvae return to the coastal environment at some subsurface depth.     

Distribution,life history,and production of three species of <Emphasis Type="Italic">Neomysis</Emphasis> in Akkeshi-ko estuary,northern Japan

Spatial and seasonal distribution pattern, life history and production of three species of Neomysis (Mysidacea) which commonly occur in northwestern subarctic Pacific coastal waters, were investigated throughout the year in the Akkeshi-ko estuary, northern Japan. The most abundant species Neomysis awatschensis (annual mean density: 179.8 inds. m⁻², biomass: 108.8 mg DW m⁻²) occurred at the inner part of the estuary including low salinity areas with no clear preference for the seagrass bed. The second most abundant Neomysis mirabilis (mean density: 95.8 inds. m⁻², biomass: 90.1 mg DW m⁻²) occurred at relatively saline seagrass site throughout the year. Occurrence of Neomysis czerniawskii in the estuary was limited to the seagrass bed during summer when their population mainly consisted of juveniles, suggesting that this species is a seasonal migrant between the estuary and the marine environment. Both N. awatschensis and N. mirabilis populations were composed of two generation types, a larger sized overwintering and smaller sized spring/summer generations; however, each species had a different reproductive strategy. N. awatschensis was characterized by fast growth to maturity at a smaller size than N. mirabilis with a relatively high fecundity during warm season, suggesting that this species is an r-strategist which can utilize opportunistically a wide variety of habitats. In contrast, the seagrass bed resident N. mirabilis was a K-strategist which matures at a larger size producing fewer but larger offspring. The annual production of N. awatschensis (0.57–0.70 g DW m⁻², mean of the whole estuary) and N. mirabilis (0.58–0.68 g DW m⁻², mean of the seagrass bed) at their respective habitats was comparable. Consequently, species-specific life history and distribution pattern are concluded to allow Neomysis spp. to coexist in the estuary and the high carrying capacity of seagrass bed is suggested to contribute to maintain their high biomass level.     

Behavioural and physiological responses of the Norway lobster,<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> (Crustacea: Decapoda), to sulphide exposure

The Norway lobster, Nephrops norvegicus, is moderately tolerant of sulphide [the lethal time to 50% mortality (LT₅₀) was 22.5 h when exposed to 500 M sulphide] but, whenever possible, it attempts to avoid the presence of sulphide in its immediate vicinity. Any sulphide entering the animal is oxidized to thiosulphate, which accumulates in the haemolymph and in the tissues. During exposure to low concentrations of sulphide, the rate of oxygen consumption is maintained or even enhanced even though the lobsters become quiescent. The apparent increase in oxygen consumption is probably due to the oxidation of sulphide to thiosulphate. At higher concentrations, oxygen consumption decreases, perhaps because of the inhibitory effect of sulphide on electron transport, and N. norvegicus resorts to anaerobic metabolism as indicated by the accumulation of lactate in the haemolymph and in the tissues.Communicated by L. Hagerman, Helsingør     

Fécondité de Nephrops norvegicus dans le golfe de Gascogne

Females of the Norway lobster Nephrops norvegicus (L.) incubate their eggs under the abdomen for a period of several months. We estimated the individual fecundity of N. norvegicus (1) before spawning, and (2) at the end of the incubation period. From these two relationships, we then calculated the average egg loss. Egg loss seems to be relatively higher for small females than for large ones. The mean egg loss estimated for the Bay of Biscay (45%) differs from the only other existing value (75%) calculated by other authors for Portuguese waters. Egg loss may regulate stock-density, and may be a mechanism intervening in stock-recruitment. In order to estimate the fecundity per recruit, fecundity at the end of the incubation period must be used: this takes into account the rate of egg loss as a function of female length.

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Contribution No. 721, du Centre Océanologique de Bretagne     

Effect of aerial exposure on concentrations of selected metabolites in blood of the Norwegian lobsterNephrops norvegicus (Crustacea: Nephropidae)

The Norwegian lobsterNephrops norvegicus (L.) collected from Firth of Clyde, Scotland between December 1987 and March 1988, was unable to survive longer than 18 h experimental emersion at 10°C. During this time the partial pressure of oxygen (P O₂) in the venous blood decreased rapidly and the lobster supplemented cellular energy requirements by anaerobic metabolism. This was indicated by the rapid accumulation ofL-lactate in the blood. Although the survival rate increased (to ca 36 to 48 h) if lobsters were kept on ice, the accumulation ofL-lactate in the blood was not significantly different from lobsters at 10°C, despite the temperature difference. There was no indication thatN. norvegicus was able to further metabolize circulatingL-lactate during emersion. On emersion there was also a marked hyperglycemia in the blood due to the stress of handling and asphyxiation. There was fairly good agreement between results obtained during laboratory studies and simulated fishing activity in the Firth of Clyde. Both sets of results are discussed in the context of adaptation to air breathing within the Crustacea and an assesment of post-harvest treatment of lobsters.     

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.     

Effects of hypoxia on the respiratory and circulatory regulation of Nephrops norvegicus

The burrowing decapod Nephrops norvegicus (L.) was kept under various degrees of hypoxia in order to measure respiration, heart rate, scaphognathite rate, haemolymph oxygen content and pH. An emergence reaction to hypoxia occurred only in dim light (<10^-2 m-c) or darkness, but after 10 d of moderate hypoxia the decapods showed no emergence response at all. The weight specific respiration of quiescent individuals was relatively low and increased only slightly in hypoxia (P_wO₂=40 torr). Heart rate, about 50 beats min^-1, changed little during hypoxia, down to P_wO₂=40 torr, whereas scaphognathite rates rose from about 60 beats min^-1 at normoxia to peak at 120 beats min^-1 at P_wO₂=40 torr. The oxygen extraction efficiency (E) remained at 20 to 30% during the first hour of hypoxia then rose gradually to maximum values of 30 to 40%. A small respiratory alkalosis of the blood became evident only after 4h of hypoxia (P_wO₂=50 torr). Normoxic postbranchial O₂ tensions (P_aO₂) were low (25–30 torr) and showed only a small decline during hypoxia. Over 10 to 13 d in moderate hypoxia an effective biosynthesis of 0.024 mM haemocyanin individual^-1 d^-1 occurred in fed decapods, whereas controls (normoxic) showed no significant change in pigment levels. A linear relationship between oxygen carrying capacity and haemocyanin concentration was found. It is contended that N. norvegicus is better able to cope with periodic exposure to hypoxia when food of sufficient quantity and quality is available.     

Swimming performance and endurance of the Norway lobster Nephrops norvegicus

The swimming behaviour of the Norway lobster, Nephrops norvegicus (L.), was studied in the laboratory. The lobsters were stimulated to swim repeatedly until they failed to respond to a single tactile stimulus. The position of the initiating stimulus on the body determined the height of the escape path above the bottom of the tank. A stimulus to the rostrum resulted in a low swimming trajectory, rarely exceeding a height of 0.1 m. A similar tactile stimulus to the abdomen produced higher mean swimming trajectories up to 0.5 m. Several parameters of swimming performance and endurance were measured for each swimming sequence, including distance, duration, mean and maximum velocities and the frequency and number of tail beats. Results showed no significant differences in the average swimming performance and endurance of males and females, but there was a sex difference in the relationship between swimming performance and N. norvegicus size.     

Ecological investigations of the marine nematode Metoncholaimus scissus

Significant concentrations of the large oncholaimid Metoncholaimus scissus Wieser and Hopper (1967) (>7000 animals/24 cm² area or 2.68x10⁶ specimens/m²) have been noted in soft-bottom turtle grass communities. Activities of the nematode frequently show an indirect relationship to blooms of the benthic diatom Pleurosigma balticum. Analysis of biomass data for M. scissus (as much as 28 g wet weight/m²) demonstrate the importance of microsite activity and sensitivity of the species to small significant changes in the benthic environment.This work was supported by grant GM 12824 from the National Institutes of Health to the Institute of Marine and Atmospheric Sciences, University of Miami. Contribution No. 1188 from the Institute of Marine and Atmospheric Sciences, University of Miami and from the Entomology Research Institute, Research Branch, Canada Department of Agriculture, Ottawa.     

Factors affecting the burrowing behaviour of Helice tridens (Grapsidae) and Macrophthalmus japonicus (Ocypodidae) in an estuary of northeast Japan

Helice tridens (De Haan) (Grapsidae) and Macrophthalmus japonicus (De Haan) (Ocypodidae) build separate burrows in reed marshes and muddy tidal flats, respectively, in the brackish-water estuaries of northeast Japan (38°11N, 141°48E). Habitat segregation between burrows of these two species was analysed by comparing the density of burrows of both species in their natural habitats, and on tidal flats subjected to various types of artificial treatment, in the summer of 1981 and 1982. Baskets, containing many stones, were placed on the ground in the reed marsh, tidal flat and creek and attracted many individuals of H. tridens, but not M. japonicus. When stones were placed on an area of tidal flat, H. tridens frequently formed burrows at the border between the stones and mud, suggesting that burrowing of H. tridens was related to the presence of solid substances lying on the ground, such as stones and shoots of the reed Phragmites australis (Cav.) Trin. In closed systems on the tidal flat and reed marsh, H. tridens and M. japonicus were able to construct burrows in both substrata regardless of high or low frequency. Moreover, it was recognized that H. tridens prevented the burrowing of M. japonicus in these closed systems. These results suggested that habitat segregation between H. tridens and M. japonicus burrows was caused primarily by an exclusive interaction between individuals of these two species.     

Home range and social behavior of three species of European Pitymys (Mammalia,Rodentia)

Summary Habits, home range, and social behavior of three species of voles of the genus Pitymys, P. multiplex, P. subterraneus, and P. savii, were studied in Tessin (Switzerland) by radioactive tagging. P. multiplex and P. savii are fossorial and often use the burrow systems of moles (Talpa); P. subterraneus moves around on the surface under dense vegetation. Males and females of P. multiplex were territorial and their home ranges averaged 342 and 229 m² respectively. The mating system appeared to be monogamous. P. savii male and female home ranges averaged 445 and 298 m² respectively. Groups of up to one to three adult females, one or more adult males, and several young, occupied exclusive communal territories. The mating system appeared to be flexible, including monogamy, polygyny, and polyandry. P. subterraneus females occupied exclusive territories (mean 256 m²) overlapped by the home ranges of one or more males (1025 m²). The mating system appeared to be promiscuous. Individuals of the three species occupied one to seven nests. In P. savii all family members utilized three or more communal nests at the same time or alternatively. Co-nesting between males and females of P. multiplex or P. subterraneus were observed less frequently.     

Distribution of Trypetesa lampas (Cirripedia,Acrothoracica) in various gastropod shells

The distribution of Trypetesa lampas (Hanoock) and its rates of infection of different species of gastropod shells in the waters of Anglesey were studied. T. lampas was found in depths from 3 to at least 25 fathoms (5 to 46 m). It infected dead shells, mainly of Buccinum undatum L., Colus gracilis (da Costa), Neptunea antiqua (L.) and Natica alderi (Forbes); it was absent from shells occupied by hermit crabs in the intertidal zone. Significantly more B. undatum shells were infected than shells of N. antiqua and Natica alderi, and more of both C. gracilis and N. antiqua than of Natica alderi. Larger shells of B. undatum were more likely to be infected than were the smaller ones. The number of females per shell varied with the shell length raised to the power 2.39 instead of as expected with the square of the length.     

NADP+-dependent glutamate dehydrogenase from Acropora formosa: purification and properties

As an initial step in our study of nitrogen metabolism in the coral/algal symbiosis we have purified glutamate dehydrogenase (EC 1.4.1.4) to homogeneity from polyp tissue of the staghorn coral Acropora formosa collected from Magnetic Island (North Queensland) in 1985–1986. The purified enzyme had a specific activity of 78 U mg^-1. The native enzyme had a relative molecular weight, M _r, of 360 000 (±20 000), and appears to be a hexamer with subunits of M _r=56000 (±3 000). Like the enzyme from other coelenterates, the coral glutamate dehydrogenase (GDH) was absolutely specific with respect to the coenzyme substrate (NADP⁺/NADPH), and was insensitive to allosteric regulation by nucleotides; unlike other coelenterate GDHs, the coral enzyme was absorlutely specific for ammonium as amino group donor in the reductive amination reaction, and major differences in kinetic properties were apparent. Linear Michaelis-Menten kinetics were observed for the substrates a-ketoglutarate, NADPH and NADP⁺, the K _m values being 0.93, 0.11 and 0.03 mM, respectively. However glutamate dehydrogenase displayed biphasic kinetics with respect to l-glutamate and ammonium, indicating two apparent K _m values (18 and 81 mM for l-glutamate and 9.2 and 416 mM for ammonium). The enzyme also exhibits Scatchard plots, Hill coefficients and cooperativity indices characteristic of enzymes displaying negative cooperativity.