Respiration,energy balance and development during growth and starvation of Carcinus maenas L. larvae (Decapoda:Portunidae)

In all larval stages of Carcinus maenas L. oxygen consumption was measured at three temperatures (12,18,25 °C). Values increased during development and were in the range of 0.037 ± 0.01 (zoea-1, 12°C, $\text{x}\text{?}\text{± 95\% CL}$) to 0.734 ± 0.047 μl O₂ · h^?1 · ind^?1 (megalopa, 25 °C). Growing larvae showed temperature dependent trends in weight specific respiration rates (referred to dry wt; DW), with values between ≈2.4 and 9.4 μl O₂· h^?1·mg DW^?1. Increase in oxygen consumption of megalops did not differ much at temperatures between 18 and 25 °C. This points to an exceptional physiological position of this stage. Fed zoea-1 of C. maenas (18 °C) revealed growth rates in terms of 40% DW, 20% carbon (C), 30% nitrogen (N) and 65% hydrogen (H). At the same time larvae gained individual energy by 13% (J · ind^?1), while weight specific energy dropped by ≈ 19% (J · mg DW^?1) during the first day and remained constant until the moult. Starved zoea-1 of C. maenas (18 ° C) gained ≈ 20 % in DW through the first day, probably caused by inorganic salts which enter the organism after the moult of the prezoea. DW dropped to ≈ 25 % of initial value, when starvation continued. Single components decreased by ≈50% (C), 54% (N), 57% (J · ind^?1). Weight specific energy (J · mg DW^?1) decreased by 40% during the first 4 days of starvation, remaining constant thereafter. Individual respiration rate (R) dropped by 61 %, weight specific respiration rate (QO₂) by 55 %. Individual energy loss in starved zoea-1 was 0.077 J over a period of 11 days. In this period ≈ 9.3 μl O₂·ind^?1 were consumed. Thus effective oxygen capacity was lower than in growing larvae. It dropped to 5.3 J·mlO₂^?1 after 4 days and remained constant if starvation continued, i.e. 65 % of possible energy loss occurred during the first 4 days. Decrease in requirement for oxygen and its effective capacity were both recognized as independent components of survival during starvation. Partitioning of energy through individual larval development of C. maenas was investigated for all five larval stages. The cumulative budget could be calculated: consumption (C) = 28.23 J, growth (G) = 0.92 J, exoskeleton (Ex) = 0.20 J, metabolism (M) = 5.30 J, egestion and excretion (E) = 21.82 J. Mean gross and net growth efficiency were, K₁ = 3.3% and K₂ = 14.8%, respectively.     

The incidence of autotomy in Carcinus maenas (L.)

A total of 1023 individuals of the common shore crab, Carcinus maenas (L.), were obtained from the environs of Whitby Harbour and Robin Hood's Bay. The bulk of the sample was taken from the sublittoral zone, but crabs from a sheltered shore were included. Each crab was measured, sexed and examined for the incidence of autotomized and regenerated limbs.There was a positive correlation between the incidence of autotomy and carapace width for crabs in both the sublittoral zone and the exposed intertidal zone. In sublittoral crabs the males suffered a higher incidence of autotomy than did the females and the incidence of cheliped autotomy was higher than for the walking limbs. The discrepancy between the loss of the chelipeds and walking limbs was reflected in a similar disparity between the occurrence of the regenerated chelipeds and walking limbs.Crabs on an exposed shore suffered a higher incidence of autotomy than did those from a sheltered shore. There were more crabs with cumulative limb losses than would be expected which suggests that once a crab enters into a state of autotomy it becomes increasingly vulnerable to the loss of another limb. Only 1 % of the population of sublittoral crabs would be expected to carry the loss of five limbs at once. No crabs were found with six or more limbs missing.     

Influence of starvation on the larval development ofHyas araneus (Decapoda,Majidae)

The influence of starvation on larval development of the spider crabHyas araneus (L.) was studied in laboratory experiments. No larval stage suffering from continual lack of food had sufficient energy reserves to reach the next instar. Maximal survival times were observed at four different constant temperatures (2°, 6°, 12° and 18 °C). In general, starvation resistance decreased as temperatures increased: from 72 to 12days in the zoea-1, from 48 to 18 days in the zoea-2, and from 48 to 15 days in the megalopa stage. The length of maximal survival is of the same order of magnitude as the duration of each instar at a given temperature. Sublethal limits of early starvation periods were investigated at 12 °C: Zoea larvae must feed right from the beginning of their stage (at high food concentration) and for more than one fifth, approximately, of that stage to have at least some chance of surviving to the next instar, independent of further prey availability. The minimum time in which enough reserves are accumulated for successfully completing the instar without food is called point-of-reserve-saturation (PRS). If only this minimum period of essential initial feeding precedes starvation, development in both zoeal stages is delayed and mortality is greater, when compared to the fed control. Starvation periods beginning right after hatching of the first zoea cause a prolongation of this instar and, surprisingly, a slight shortening of the second stage. The delay in the zoea-1 increases proportionally to the length of the initial fasting period. If more than approximately 70 % of the maximum possible survival time has elapsed without food supply, the larvae become unable to recover and to moult to the second stage even when re-fed (point-of-no-return, PNR). The conclusion, based on own observations and on literature data, is that initial feeding is of paramount importance in the early development of planktotrophic decapod larvae. Taking into account hormonal and other developmental processes during the first moult cycle, a general hypothesis is proposed to explain the key role of first food uptake as well as the response pattern of the zoea-1 stage to differential starvation periods.Contribution to research project Experimentelle Marine Ökosystemanalyse sponsored by Bundesministerium für Forschung und Technologie, Bonn (Grant No. MFU-0328/1).     

Heart and scaphognathite activity in the shore crab Carcinus maenas (L.)

Simultaneous recordings of the activity of the heart and both scaphognathites of Carcinus maenas (L.) were investigated during a wide repertoire of normal activities. The scope of activity for the heart and scaphognathites could usefully be subdivided into three smaller ranges or levels which, although arbitrary, were recognizable on the basis of frequency and arrhythmic phenomena.Alterations of heart and scaphognathite activity induced by a variety of laboratory treatments were investigated in order to evaluate and aid the interpretation of such data for comparative purposes. Providing the recent manipulative history of the animal is known, the behaviours of heart and scaphognathites have been shown to be predictable and useful as sensitive sources of comparative data.The observed behaviour of the heart and scaphognathites has been discussed in terms of their nervous control systems and of their implications in a variable system of blood circulation and gill ventilation.     

The respiratory responses of Carcinus maenas (L.) to changes in environmental salinity

Exposure of Carcinus to a reduction in salinity is accompanied by an increase in oxygen consumption. The highest rates of oxygen consumption are found during the first 2–3 h after a reduction in salinity, after which oxygen consumption then declines, but to a higher than ‘normal’ level which is maintained throughout the period of exposure to low salinity. This augmented rate of oxygen consumption persists even after 3–4 days in water of reduced salinity and indicates that the respiratory rate of Carcinus does not acclimate rapidly to reduced salinity. The increased oxygen consumption is associated with an increase in ventilation volume resulting from an increase in the rate of beating of the scaphognathites, while oxygen utilization remains at a low level. Since the oxygen-transporting properties of the blood of Carcinus show little change under conditions of reduced salinity, the increased oxygen demand of the tissues is met by a rise in cardiac output resulting mainly from an increase in heart rate.     

Temperature and growth in Carcinus maenas L. (Decapoda: Portunidae) larvae reared in the laboratory from hatching through metamorphosis

Larvae of Carcinus maenas L. were reared in the laboratory from hatching through metamorphosis at 9, 12, and 18°C. Dry weight (DW) and elemental contents of carbon (C), nitrogen (N), and hydrogen (H) were analysed at short intervals through successive larval moulting cycles (four zoea-stages, megalopa), and newly metamorphosed crabs. C. maenas larvae grew significantly during all instars, at all temperatures tested. Biomass (DW, C, N, H) and energy (Joule) slightly declined shortly before ecdysis in zoea stages. This terminal decrease was more distinct in the megalopa stage, where ≈39 and 83% of the maximum energy attained, was lost at 12 and 18°C, respectively. Changes of biomass and energy in successive moult cycles showed best fits to quadratic equations, whereas their maximum in successive larval instars formed exponential sequences with time. Due to parabolic growth curves, biomass and energy accumulation within single instars were discussed as maximum (MG) and effective growth (EG), considering gain both at times of maximum biomass, and shortly before ecdysis. Metamorphosing larvae achieved EG with 1137% (DW), 1195% (C), 1108% (N), 1395% (H), 1339% (Joule) at 12°C, and 1140% (DW), 1099% (C), 1133% (N), 1225% (H), 1107% (Joule) at 18°C, relative to newly hatched zoea-1. Ash content and inorganic C in newly hatched zoea-1, were 29.4% and 5.5% ash, respectively. The stoichiometric C H N method of Gnaiger & Bitterlich was used to assess protein, lipid, and carbohydrate compositions. Obviously proteins formed the major part of larval biomass (>50% DW). C: N ratios indicate that more lipid than protein was built up shortly after moulting, but relatively more protein was subsequently accumulated. Temperature effects on larval growth (MG, EG), growth rates (GR), and gross growth efficiencies (K₁) were discussed. C. maenas zoea stages accumulated energy and biomass with higher efficiencies at 18 than at 12°C. Megalopa growth seemed to be limited at 18°C, showing lower K₁ values than at 12°C. N was accumulated with higher efficiencies than C in all larval stages. Characteristic variations in larval K₁ values between premoult and ecdysis were discussed. Cumulative gross growth efficiencies (MG-related) were calculated as ≈11 and 10%, at 12 and 18°C, respectively.     

Patterns of morphological and genetic variability in UK populations of the shore crab, Carcinus maenas Linnaeus, 1758 (Crustacea: Decapoda: Brachyura)

Previous research has identified extensive inter-population variability in the morphology of the shore crab (Carcinus maenas L.). To determine the source of this variation (genetic or environmental), morphological and genetic data were analysed from crabs collected from eight sites around the coast of the UK. Ten morphometric traits were measured from over 800 crabs and the degree of morphological similarity among sites was calculated using multivariate techniques. Allozyme electrophoresis was used to investigate patterns of genetic similarity. Extensive morphological variability was detected: eight out of the ten morphometric traits analysed were useful when discriminating between crabs from each site. Discriminant function analysis revealed that over 35% of individuals could be classified to their site of origin on the basis of their morphology. In contrast, the allozyme analysis revealed low levels of genetic variability, both within the meta-population and among the crab population at each site. Pairwise comparisons revealed a moderate correlation between the degree of morphological and genetic similarity of crabs at each site, which suggests that the observed phenotypic variability has a genetic component. However, only around 20% of the phenotypic variability detected was associated with the patterns of genetic similarity. This means that patterns of morphological variability in this species are largely determined by the local environmental conditions: local factors could have a within-generation selective influence on mean trait values or C. maenas may exhibit phenotypic plasticity.     

Ventilatory responses to ambient oxygen in stressed and fully settled shore crabs Carcinus maenas (L.)

Scaphognathite activity and ventilatory responses to changes of ambient oxygen have been studied in stressed and fully settled crabs, Carcinus maenas (L.). Compared with resting, fully settled animals, resting stressed crabs exhibited very high values of scaphognathite beat frequency, negative hydrostatic pressure in the gill chamber and amount of time spent ventilating, suggesting a much elevated ventilatory flow rate. Unlike fully settled animals, stressed crabs did not show significant ventilatory responses to changes of ambient oxygenation.     

Reconstitution of Carcinus maenas (crustacea) hemocyanin,in the presence of nonionic detergent

Carcinus maenas hemocyanin was reconstituted from the apo-protein using copper(I) chloride in presence of Triton X-100 at physiological pH. In these conditions Cu⁺ is stable and permitted a kinetic study of the reconstitution reaction to be made.     

Size,temperature, and scaphognathite frequency-dependent variations of ventilation volumes in Carcinus maenas (L.)

There is a direct, linear relationship between scaphognathie beat frequency and absolute ventilation volume in Carcinus maenas (L). The absolute ventilation volume is directly proportional to the size of the animals. At all scaphognathie frequencies small animals have a greater relative ventilation volume than large animals. The absolute stroke volume (ml/beat) is greater in large animals than in small animals. The majority of the available evidence suggests that, whilst the force of the scaphognathie stroke may be variable, the amplitude of its excursion remains constant at all beat frequencies.     

Fine structure and secretion of the capsule enclosing the embryo in a crab (Carcinus maenas (L.))

Carcinus maenas oocyte is encased, at the end of viteilogenesis phase, by two superposed vitelline envelopes. If the outermost (envelope la) is relatively thin, the innermost (envelope 1b) is however five times much thicker. The envelope 1b, secreted mostly by the oocyte, contains proteinaceous material and a substance which strongly binds WGA. The 1b material deposition is completed when the oocyte nuclear apparatus still has the characteristics of a germinal vesicle. After the secretion phase and always at the ovarian level, envelope 1b material undergoes remarkable structural modifications. These last events occur while the oocyte nuclear apparatus is in the first meiotic metaphase stage. Later, after egg-laying, a new material appears below the two superimposed vitelline envelopes 1a and 1b. This new innermost material (envelope 2), observed as a very thick and homogeneous secretion, is secreted by the newly laid egg probably after fecondation. Envelope 2 material is wholly deposited in few hours and arises from vesicles of an oocytic origin. These three superimposed envelopes make up the embryo capsule of which the main part is constituted by envelope 2. This tripartite capsule will stay, surrounding the developmental stages, superimposed to newly secreted successive embryonic envelopes, up to the end of embryogenesis.     

Effects of juvenile non-indigenous Carcinus maenas on the growth and condition of juvenile Cancer irroratus

The Atlantic rock crab, Cancer irroratus, is a commercially fished species and a critical prey item for the American lobster, Homarus americanus, in Atlantic Canada. The recent invasion of European green crab, Carcinus maenas, may have significant effects on the growth and condition of native C. irroratus, because both species overlap spatially and temporally and have similar habitat and dietary requirements. To examine such potential effects, we measured the growth of juvenile C. irroratus in the presence of juvenile C. maenas over a period of 4 months (growing season), under the following species combinations: (1) one C. irroratus (10-25 mm CW); (2) two C. irroratus (10-25 mm CW); (3) one C. irroratus (10-25 mm CW) and one C. maenas (10-15 mm CW). Morphological measurements included pre- and post-molt carapace width, chela height, abdomen width (mm), weight (g), and estimates of molt increment (%) and intermolt duration (days). Analysis of the hepatopancreas for % lipid content at the end of the experiment provided an estimate of physiological condition. The effect of the presence of C. maenas on the growth of C. irroratus shifted from negative to positive, when C. irroratus reached CW of 19-22 mm and gained a presumably significant size advantage over C. maenas. The positive effect resulted from increased energy intake through crab consumption. In the absence of crab consumption, the presence of a second crab (conspecific or C. maenas) had no effect on growth. C. irroratus consumed crabs more frequently when the second individual was a green crab than a conspecific. Consumption of C. maenas had a pronounced effect on the growth rate of C. irroratus, resulting in shorter intermolt periods and larger percent molt increments than in the presence of a conspecific. Therefore, the presence of juvenile C. maenas does not appear to have a prolonged negative effect on the growth of C. irroratus; rather, it may provide an additional food item as rock crabs grow, as long as encounters between the two species occur at high enough rates.     

The geography of crushing: Variation in claw performance of the invasive crab Carcinus maenas

The major claws of predatory, durophagous decapods are specialized structures that are routinely used to crush the armor of their prey. This task requires the generation of extremely strong forces, among the strongest forces measured for any animal in any activity. Laboratory studies have shown that claw strength in crabs can respond plastically to, and thereby potentially match, the strength of their prey's defensive armor. These results suggest that claw strength may be variable among natural populations of crabs. However, very few studies have investigated spatial variation in claw strength and related morphometric traits in crabs. Using three geographically separate populations of the invasive green crab in the Gulf of Maine, we demonstrate, for the first time, geographic variation in directly measured claw crushing forces in a brachyuran. Despite variation in mean claw strength however, the scaling of claw crushing force with claw size was consistent among populations. We found that measurements of crushing force were obtained with low error and were highly repeatable for individual crabs. We also show that claw mass, independent of a linear measure of claw size, and carapace color, which is an indicator of time spent in the intermoult, were important predictors of claw crushing force.     

Temperature and the larval development of Hyasaraneus L. (Decapoda : Majidae); extrapolation of laboratory data to field conditions

The larval development of the spider crab Hyas araneus L. was studied in the laboratory at different constant temperatures (2, 6, 12, and 18°C). Linear relationships between log temperature and log stage duration were described by means of regression equations. They were used in a simple simulation model predicting larval moulting and metamorphosis at different temperatures. Experiments were also carried out under temperature regimes which simulated field conditions. Correspondence between predicted data and those observed in these experiments was generally good. Based on these observations, a number of predictions was made on larval occurrence in the plankton, on changes in the composition of larval populations, and on the period and intensity of settlement near Helgoland (North Sea). The extent and quality of the effects of deviations from the average course of temperature were also estimated. The most important predictions were: (1) settlement of H. araneus takes place mainly from late May to mid-June; (2) assuming wide annual fluctuations in the average spring temperature (±2°C), a total settlement period ranging from late April to early July can be expected; (3) delay of larval development due to a decrease in temperature is stronger than the acceleration caused by an equally great increase; and (4) both delay and acceleration effects become weaker during the hatching season and from stage to stage.     

Physiological and metabolic variations in Carcinus mediterraneus (Crustacea: Decapoda) parasitized by Thelohania maenadis (Microspora: Microsporida): An ecophysiopathological approach

This paper examines the influence of Thelohania maenadis (Protozoa: Microsporida), a muscle microsporidian, on certain features of the metabolism of Carcinus mediterraneus (Crustacea: Decapoda). Comparison of biochemical parameters reveals that only hemolymph total protein and glucose levels are reduced significantly by this parasitic infection. Muscular effort results in a diminution of muscular glycogen and hemolymph glucose levels, while hepatopancreatic glycogen remains unaffected; lactate accumulates in the muscle tissue. Experimental variations in the environmental water temperature and salinity affect protein levels and metabolic indicators of carbohydrate metabolism. Proteinemia varies with temperature and salinity, but parasitic infection accentuates these variations. Glucose and lactate concentrations in the hemolymph are increased in healthy specimens after muscular effort, under conditions of increased temperature or hypoosmotic shock; microsporidiosis provokes the reverse of these tendencies, on a less pronounced scale. Hyperosmotic shock at high temperature leads to a slight reduction in glucose and lactate concentrations. The paper concludes with a discussion of the physiopathological significance of the results obtained as an expression of the effects of parasitism infestation.     

Genetic differentiation in the mud crab Scylla serrata (Decapoda: Portunidae) within the Indian Ocean

Scylla serrata (Decapoda: Portunidae) is a swimming crab that is widespread in the Indo-Pacific region and commonly found in estuarine and mangrove waters. An extended planktonic larval phase suggests high dispersal potential and the possibility of extensive gene flow between conspecific populations at least on a geographic mesoscale (tens to hundreds of kilometres).Intraspecific variation of the mitochondrial DNA cytochrome oxidase subunit I (mtDNA COI) gene was investigated in 77 individuals from four representative mangrove swamps of the African tropics (Kenya and Zanzibar) by means of DNA sequencing. We examined 535 base pairs (bp) and identified 24 different haplotypes. Each population sample is characterised by a single most frequent haplotype, shared among all four populations, and a small number of rare ones, typically present in only one or two individuals and representative of a specific population.Analysis of molecular variance (AMOVA), F_ST statistics and χ² contingency analysis of spatial distribution of mtDNA haplotype frequencies revealed in toto a significant genetic differentiation among populations. These results could indicate that gene flow might be reduced, even between geographically close sites, despite the high potential for dispersal; anyway, at the recorded level of divergence and owing to the abundance of rare haplotypes and singletons in our data set, repeated sampling over time is necessary to establish whether the recorded pattern of genetic differentiation is stable and biologically significant.Finally, integration of our data with those reported by Gopurenko et al. [Mar. Biol. 134 (1999) 227] on S. serrata from South Africa, Red Sea and Mauritius Islands allowed to infer S. serrata population structure within a larger area of the Indian Ocean region.     

Predatory impact of the green crab (Carcinus maenas Linnaeus) on post-settlement winter flounder (Pseudopleuronectes americanus Walbaum) as revealed by immunological dietary analysis

Predation on flatfish during the early juvenile stage is an important factor regulating year-class strength and recruitment. In this study, immunological dietary analysis was performed on green crabs (Carcinus maenas) collected from the Niantic River, Connecticut, in an effort to evaluate the predatory impact of this species on post-settlement winter flounder (Pseudopleuronectes americanus). Through the use of species-specific antiserum, winter flounder proteins were identified in 4.8% of the green crab stomachs analyzed (n = 313, size range = 14-74 mm carapace width, CW), revealing that crabs ≥ 29 mm CW are predators of post-settlement winter flounder in natural populations. The most significant factor underlying the predator-prey interaction was the relative size relationship between species, such that the incidence of winter flounder remains in the stomach contents of green crabs was positively correlated with predator-to-prey size ratio. Results from dietary analysis were incorporated into a deterministic model to estimate the average daily instantaneous mortality and cumulative mortality of winter flounder owing to green crab predation. Accordingly, green crabs may account for 0.4% to 7.7% (mean = 2.2%) of the daily mortality of winter flounder and consume 1.1% to 32.3% (mean = 10.2%) of the flounder year-class. Model simulations further indicate that variations in green crab abundance and size-structure account for the greatest variability in winter flounder mortality. Relative to other macro-crustacean predators, however, predation by green crabs has a minimal effect on winter flounder survival, due in large part to the low densities of these crabs in temperate estuaries.     

Agonistic interactions between the invasive green crab, Carcinus maenas (Linnaeus) and juvenile American lobster, Homarus americanus (Milne Edwards)

Invasive organisms have the potential for competition with native organisms. In the Southern Gulf of St. Lawrence, juvenile American lobsters have a potential spatial overlap with adult green crabs. Crustaceans use agonistic behaviour to settle disputes, with the larger organism often winning contests for limited resources such as food and shelter. Two experiments were carried out using adult green crabs (53-76 mm carapace width) and juvenile American lobsters (28-57 mm carapace length). The first experiment used a limited food resource. We found that green crabs were the first to the food in all trials, fed in significantly more trials than lobsters and spent a significantly greater proportion of time with the food. The lobsters were only able to displace the green crabs from the food in 2 of 65 attempts. The second experiment was designed to examine shelter competition; unexpectedly some predation by green crabs on lobsters occurred, which allowed us to test hypotheses about how relative size and shelter use affect predation. Green crabs captured and consumed juvenile lobsters in 6 of 11 trials. The lobsters that survived spent significantly more time in shelter. There was no clear relationship between shelter use and size of lobster. The lobsters that were larger in relation to the green crabs suffered a higher rate of predation, which we believe was due to more conspicuous activity and less use of shelter. It appears that green crabs have the potential to negatively impact native juvenile lobster.     

Agonistic interactions between invasive green crabs, Carcinus maenas (Linnaeus), and sub-adult American lobsters, Homarus americanus (Milne Edwards)

The invasive green crab, Carcinus maenas, has recently expanded its range into the Southern Gulf of St. Lawrence, where there is potential for substantial niche overlap with juvenile American lobsters, Homarus americanus. We used two experiments to elicit, record and analyze the agonistic interactions of adult green crabs (carapace width of 63-75 mm) and sub-adult (carapace length of 55-70 mm) lobsters. The first experiment gave each animal equal access to a limited food resource. The green crabs were first to the food in significantly more trials, spent a significantly greater proportion of time with the food, and were able to successfully defend the food from attacks by the heavier lobsters. In the second experiment, we allowed the lobsters to gain possession and initiate feeding on the food before releasing the green crabs. In these trials, the lobsters spent significantly more time with the food, and were able to defend the food from the green crabs. The results of both experiments are discussed in the context of game theory. The different behaviour of the crustaceans in the two experiments is consistent with the “bourgeois” strategy in a hawk and dove game simulation. With this strategy, an animal acts like a hawk if in possession of a resource, but acts like a dove if the other animal is in possession of the resource. The fact that the green crabs were able to physically compete with, and in many cases dominate the larger, heavier lobsters supports the potential for competitive impacts of green crabs on sub-adult lobsters.