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.     

Effect of bilateral eyestalk ablation on ovarian development and moulting in early and late intermoult stages of female spiny lobster Panulirus homarus (Linnaeus, 1758)

Abstract

The present study describes the effect of bilateral eyestalk ablation (BESA) on reproduction and moulting of spiny lobster Panulirus homarus females in their early and late intermoult stages. The lobsters obtained from the wild were conditioned for the experiment. The experiments were conducted at the Calicut research centre of Central Marine Fisheries Research Institute, India. Eyestalk ablation of females was done by ligation. The responses of the bilaterally eyestalk-ablated lobsters were statistically analysed. BESA conducted on the early and late intermoult stages resulted in the simultaneous acceleration of the somatic growth and reproductive processes with higher emphasis for oogenesis in lobsters ablated in the early intermoult phase and comparatively lower activity in those ablated in the late intermoult phase. Ablation in late intermoult phase resulted in faster entry into the premoult stage compared to the control.     

The influence of temperature and light on larval pre-settlement metamorphosis: a study of the effects of environmental factors on pre-settlement metamorphosis of the solitary ascidian Styela canopus

This study investigated the effects of two environmental factors, temperature and light, on larval settlement and metamorphosis in the solitary ascidian Styela canopus. The results revealed that larval settlement rates decreased with increasing temperature in the range 12–30°C. We also demonstrated for the first time that pre-settlement metamorphosis of ascidian larvae can occur as a function of temperature. We suggest this could be an adaptation to avoid the greater energetic cost of active larval swimming, presumably resulting from the increasing temperature. They are able to metamorphose into passive drifting post-larvae and to continue planktonic life. This finding has implications for larval dispersal, especially under conditions of ocean warming. In addition, the effect of light intensity on larval settlement and metamorphosis was significantly different between photoperiods of 24 L : 0 D and 12 L : 12 D. These results provide some insight into the complex cues affecting settlement and metamorphosis of ascidian larvae and ascidian distribution in nature.     

The evolution of armament strength: evidence for a constraint on the biting performance of claws of durophagous decapods

Performance data for the claws of six sympatric species of Cancer crabs confirmed a puzzling pattern reported previously for two other decapod crustaceans (stone crabs, Menippe mercenaria, and lobsters, Homarus americanus): Although biting forces increased, maximum muscle stresses (force per unit area) declined with increasing claw size. The negative allometry of muscle stress and the stress at a given claw size were fairly consistent within and among Cancer species despite significant differences in adult body size and relative claw size, but were not consistent among decapod genera. Therefore, claw height can be used as a reliable predictor of maximum biting force for the genus Cancer, but must be used with caution as a predictor of maximum biting force in wider evolutionary and biogeographical comparisons of decapods. The decline in maximum muscle stress with increasing claw size in Cancer crabs contrasts with the pattern in several other claw traits. Significantly, three traits that affect maximal biting force increased intraspecifically with increasing claw size: relative claw size, mechanical advantage, and sarcomere length of the closer muscle. Closer apodeme area and angle of pinnation of the closer muscle fibers varied isometrically with claw size. The concordant behavior of these traits suggests selection for higher biting forces in larger crabs. The contrast between the size dependence of muscle stress (negative allometry) and the remaining claw traits (isometry or positive allometry) strongly suggests that an as yet unidentified constraint impairs muscle performance in larger claws. The negative allometry of muscle stress in two distantly related taxa (stone crabs and lobsters) further suggests this constraint may be widespread in decapod crustaceans. The implications of this performance constraint for the evolution of claw size and the "arms-race" between decapod predators and their hard-shelled prey is discussed.     

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.     

Effects of seasonality and moult cycle on the proliferation of nerve cells and on the labelling of ecdysone receptors in an estuarine crab

Decapod crustaceans show proliferation of the nerve cells in the olfactory lobe throughout their lives. However, the regulation of this process is still poorly understood, since it may vary with endogenous and exogenous factors. The objective of the present investigation was to quantify the proliferation of nerve cells and number of nerve cells with ecdysone receptors in the clusters of the central olfactory system in Neohelice granulata, according to moult stages and in different seasons (summer and winter). Three injections of bromodeoxyuridine (BrdU) were administered to the crabs. Brains were sectioned by microtome and fixed on slides for immunohistochemistry with anti-BrdU and anti-EcR antibodies. The proliferation of nerve cells was higher in winter than in summer, probably because in winter the crabs do not breed and the premoult and postmoult periods are longer. Crabs in postmoult exhibited more BrdU-labelled cells than crabs in premoult or intermoult in winter, because of a greater number of mitoses related to an increase in body size and addition of olfactory receptor neurons. The number of EcR-labelled cells was higher in premoult than in postmoult or intermoult in winter. The proliferation of nerve cells is regulated seasonally and according to moult stages.     

Dietary composition of the blue swimmer crab Portunus pelagicus L. Does it vary with body size and shell state and between estuaries?

The aim of this study was to determine the diets of Portunus pelagicus in the large Peel-Harvey and Leschenault estuaries in south-western Australia in order to ascertain whether the dietary composition of this crab changes with body size during two different moult stages and differs between the two estuaries. Portunus pelagicus, ranging in age and carapace width from ca. 2 months and 12 mm to ca. 112 years and 159 mm, were collected from the shallow basins of the Peel-Harvey and Leschenault estuaries. Examination of the cardiac stomachs of these crabs showed that P. pelagicus does not feed just before or immediately after moulting and that the stomachs of recently-moulted crabs contained significantly more food than those of intermoult crabs. Although the volumetric contribution made by calcareous material to the stomach contents was similarly high in all size classes of recently-moulted crabs, i.e. 47 to 55%, the volumetric contributions made by small bivalves decreased with body size, whereas the reverse occurred with shell fragments of large decapods and, to a lesser extent, polychaetes. The dietary compositions of intermoult crabs were shown by classification and multi-dimensional scaling ordination to differ markedly from those of recently-moulted crabs and to undergo similar progressive ontogenetic changes in both the Peel-Harvey and Leschenault estuaries. Thus, the contribution made by small benthic and epibenthic crustaceans, such as amphipods and tanaids, declined with increasing body size, whereas the reverse occurred with larger prey, such as nereid polychaetes, small decapods and teleosts. The dietary composition of P. pelagicus was influenced more by moult stage, i.e. recently moulted vs. intermoult, than by body size. Although the dietary compositions of P. pelagicus in the two estuaries were not significantly different, they did differ from those recorded from coastal marine waters in the same region, thereby reflecting differences in the potential prey in those two environments.     

The moulting cycle and changes in body density in larvae of the snow crab Chionoecetes opilio (Brachyura: Majoidea) under laboratory conditions

The moulting cycle and the time course of changes in body density from hatching to the end of the megalopal stage in snow crab (Chionoecetes opilio) larvae were investigated in laboratory-reared specimens. Morphological changes in the epidermis and cuticle were photographically documented to characterize the moult-cycle stages: A–B (postmoult), C (intermoult), D (premoult) and E (ecdysis). Moult-stage characteristics were based on a microscopical examination of integumental modifications, particularly of the telson. During stages A–C, the larval cuticle changed from a spongy structure to become conspicuously thicker and more solid in appearance. In stage D, the epidermis retracted from the cuticle and new setae and appendages were formed. The body densities of larval snow crabs were lowest just after moulting; they increased greatly during stage C, and then gradually increased to reach a plateau at 1.0897–1.0931 g cm^?3 during stage D. Over the whole larval period, they have a density greater than that of seawater. These observations will assist in understanding of larval distribution and transport in snow crabs in their natural habitat, and provide a useful tool to determine the developmental stages of larvae sampled from the plankton and from larval cultures.     

Some physiological and biochemical considerations of larval development in the American lobster,Homarus Americanus Milne Edwards

The weight-specific respiration rates of fed and starved lobsters and the ammonia excretion rates of fed lobsters increased with each larval stage (I through IV) and decreased with the first postlarval stage (V). The rate of change in metabolic rates was greater than the rate of change of body size of the larval stages, indicating an increased energy demand of the later larval stages. There was no significant difference in the O: N ratio for the first three larval stages but a reduction was observed in stage IV and V lobsters, reflecting an increased dependence on protein catabolism for energy.Protein was the principal biochemical constituent of all lobster stages. Significant decreases in lipid content and increases in ash and chitin content of the last larval (IV) and first postlarval (V) stages were detected.     

Intraspecific morphological variation related to the moult-cycle in colour forms of the shore crab Carcinus maenas

The common shore crab Carcinus maenas exhibits a range of carapace colours from green through orange to red, green forms showing some differences of distribution from red/orange forms. To test the hypothesis that colour differences were moult-related, large numbers of Carcinus were collected intertidally and subtidally in summer when moulting is most prevalent, and their moult stages determined. Red and orange coloration was found only to occur in the larger size classes in crabs in prolonged intermoult, but not solely in the largest crabs in terminal anecdysis. Red crabs were characterized by a heavier load of epibionts and a strong, thicker carapace. In contrast, green crabs were found over the entire size range and appeared to be actively moulting forms; they carried fewer epibionts and had a thinner integument than red forms. The significance of morphological differences between red/orange and green forms of Carcinus maenas is considered in relation to previously reported behavioural, physiological and ecological differences between the colour forms.     

Untersuchungen zur Häutungsphysiologie der dekapoden Krebse am Beispiel der StrandkrabbeCarcinus maenas

Under constant laboratory conditions, juvenile shore crabs moult at fixed intervals which depend upon their body size. During one moult every crab exhibits increases of the same relative amounts, independent of its absolute size. Basing on the predictable duration of the intermoult period, the morphological changes in the structure of the cuticle and the development of limb-buds, the intermoult period could be divided into 21 different stages. After studying the moulting rhythm in constant milieu, the influence of the following exogenous and endogenous factors upon the moulting rhythm and growth of normal and of eye-stalkless individuals was investigated: temperature, photoperiod, loss of pereiopods, feeding, and presence of larger specimens. From these investigations it became evident that the moulting rhythm is regulated by growth. The crabs are able to moult only after achieving a minimum of tissue growth. So long as this minimum growth is not achieved, a moult-inhibiting hormone is secreted and moulting is prevented. If the moult-inhibiting hormone is absent, moulting hormone is secreted and initiates a moult. Under dangerous conditions, the crabs are able to delay the next moult. Under unfavourable conditions they consume less food than normal. Therefore, the amount of tissue growth which is the necessary prerequisite for moulting is delayed, and continued release of moult-inhibiting hormone prevents the moult. Under conditions favourable for moulting, or demanding moult (e. g. after loss of many pereiopods) the crabs accelerate the moult. Temperature influences the moulting rhythm by indirect effects on the metabolic rate. During further investigations, the variation of the following parameters were determined quantitatively: content of moulting hormone in whole crabs; content of aminoacids, protein, glucose, Na⁺, K⁺, Mg⁺⁺ and Ca⁺⁺ in the hemolymph; pH and osmotic pressure in the hemolymph; and Ca⁺⁺ content in skeleton and whole crabs. All parameters mentioned — excepting pH and K⁺ content of the hemolymph — vary characteristically during the intermoult period. The titre of moulting hormone has 4 different maxima. Of all parameters, only the content of animoacids and protein in the hemolymph vary in the same way as the titre of the hormone. From these results the following conclusions are drawn: The moulting hormone not only initiates the moulting process, but controls it at several stages. Only protein metabolism seems to be under direct control of the moulting hormone which stimulates protein-synthesis. Chitin formation, regeneration, apolysis and ecdysis are indirectly controlled by the moulting hormone through protein metabolism. As in most of the other processes mentioned, the calcification of the new cuticle is not under the direct influence of the moulting hormone. The conclusion ofDigby (1966) that calcification in crabs is an electrochemical process, is confirmed.     

Reproductive biology and genetic diversity of the Sea Anemone Aulactinia stella (Verrill, 1864)

Artificial reef areas can be used for management and conservation of commercially exploited crustacean decapods but their behaviour in these environments is poorly characterised. Acoustic tags were used to study the behaviour of 3 adult spiny lobsters (Palinurus mauritanicus) and 3 adult spider crabs (Maja squinado) over a period of 64 days in summer, evaluating the use of artificial reef areas as suitable sites for re-stocking of overfished decapods. For this purpose, animals were released in a western Mediterranean artificial reef located at 20 m depth, close to a cabled seafloor observatory, which simultaneously recorded temperature, salinity, current direction, current speed and light intensity over the study period. Spiny lobsters lingered in the reef area, whereas the spider crabs left the area 21–45 h after release. These behavioural differences suggested that artificial reefs might be a good area to deploy adult lobsters for re-stocking purposes. The movements displayed by the lobsters during this experiment were not influenced by any of the measured environmental factors, whereas spider crab movements occurred against major current direction, when current speed was intense.     

The brighter side of climate change: How local oceanography amplified a lobster boom in the Gulf of Maine

Ocean warming can drive poleward shifts of commercially important species with potentially significant economic impacts. Nowhere are those impacts greater than in the Gulf of Maine where North America's most valuable marine species, the American lobster (Homarus americanus Milne Edwards), has thrived for decades. However, there are growing concerns that regional maritime economies will suffer as monitored shallow water young‐of‐year lobsters decline and landings shift to the northeast. We examine how the interplay of ocean warming, tidal mixing, and larval behavior results in a brighter side of climate change. Since the 1980s lobster stocks have increased fivefold. We suggest that this increase resulted from a complex interplay between lobster larvae settlement behavior, climate change, and local oceanographic conditions. Specifically, postlarval sounding behavior is confined to a thermal envelope above 12°C and below 20°C. Summer thermally stratified surface waters in southwestern regions have historically been well within the settlement thermal envelope. Although surface layers are warming fastest in this region, the steep depth‐wise temperature gradient caused thermally suitable areas for larval settlement to expand only modestly. This contrasts with the northeast where strong tidal mixing prevents thermal stratification and recent ocean warming has made an expansive area of seabed more favorable for larval settlement. Recent declines in lobster settlement densities observed at shallow monitoring sites correlate with the expanded area of thermally suitable habitat associated with warmer summers. This leads us to hypothesize that the expanded area of suitable habitat may help explain strong lobster population increases in this region over the last decade and offset potential future declines. It also suggests that the fate of fisheries in a changing climate requires understanding local interaction between life stage‐specific biological thresholds and finer scale oceanographic processes.     

Review of Panulirus argus virus 1--a decade after its discovery

In 2000, a pathogenic virus was discovered in juvenile Caribbean spiny lobsters Panulirus argus from the Florida Keys, U.S.A. Panulirus argus virus 1 (PaV1) is the first naturally occurring pathogenic virus reported from lobsters, and it profoundly affects their ecology and physiology. PaV1 is widespread in the Caribbean with infections reported in Florida (U.S.A.), St. Croix, St. Kitts, Yucatan (Mexico), Belize, and Cuba. It is most prevalent and nearly always lethal in the smallest juvenile lobsters, but this declines with increasing lobster size; adults harbor the virus, but do not present the characteristic signs of the disease. No other PaV1 hosts are known. The prevalence of PaV1 in juvenile lobsters from the Florida Keys has been stable since 1999, but has risen to nearly 11% in the eastern Yucatan since 2001. Heavily infected lobsters become sedentary, cease feeding, and die of metabolic exhaustion. Experimental routes of viral transmission include ingestion, contact, and for newly settled juveniles, free virus particles in seawater. Prior to infectiousness, healthy lobsters tend to avoid diseased lobsters and so infected juvenile lobsters mostly dwell alone, which appears to reduce disease transmission. However, avoidance of diseased individuals may result in increased shelter competition between healthy and diseased lobsters, and greater predation on infected lobsters. Little is known about PaV1 outside of Mexico and the USA, but it poses a potential threat to P. argus fisheries throughout the Caribbean.     

Sex-specific mediation of foraging in the shore crab, Carcinus maenas

Experiments were conducted to investigate the sex-specific differences to feeding responses of the shore crab Carcinus maenas throughout the year. Results demonstrate that female shore crabs exhibit stronger feeding responses than males throughout the year with a significantly reduced feeding response in males during the summer months' reproductive season. We also studied the possible function(s) of the moulting hormone, 20-hydroxyecdysone (Crustecdysone) that has been described as a potential female-produced sex pheromone to initiate male reproductive behaviour in a number of crustaceans. We recently presented evidence that for shore crabs this is not the case and now show that the steroid is instead functioning as a sex-specific feeding deterrent protecting the moulting 'soft' female crabs. Whilst male shore crabs were deterred from prey (Mytilus edulis) and synthetic feeding stimulants glycine and taurine when these feeding stimulants were spiked with crustecdysone, intermoult female crabs were significantly less affected and rarely deterred from feeding. This sex specificity of the moulting hormone, in combination with the female sex pheromone, which has no anti-feeding properties, ensures that male crabs mate with soft-shelled, moulted females rather than engage in cannibalism, such as found frequently in cases when soft-shelled females are exposed to intermoult females.     

Die Schlei,ein Modell für die Verbreitung der StrandkrabbeCarcinus maenas

ACarcinus maenas population inhabiting the Schlei, a glacial fjord of the Baltic Sea, was studied during a three-year period of at least monthly sampling. Due to slightly higher water temperatures in the Schlei (c. 1 °C higher than in the neighbouring waters of the western Baltic Sea) annual larval development starts there one month earlier. When in some years salinities are unfavourable (<13), larval development may be almost completely prevented. Juveniles and adults tolerate changing salinities, even though females prefer staying in deep waters and juveniles in shallow waters of high salinities. During winter all crabs move to deeper waters and stay huddled together in crevices and holes until March or April. Females usually moult after being fertilized, which takes place after the breeding season in August. Males moult between May and June; juveniles continue to moult during the warm season. Moulting for growth lasts until puberty is reached in the second year. From then on intermoult periods are more extended, but males moult more frequently than females, attaining ultimately a larger size. Under favourable environmental conditions, the maximum lifespan ofC. maenas in the Schlei amounts to five years. During this period, five larval moults and about fifteen moults for growth occur.     

Variability in water temperature affects trait-mediated survival of a newly settled coral reef fish

As animals with complex life cycles metamorphose from one stage to the next, carry-over effects from earlier stages can affect future mortality. To examine the relationship between early life history traits and survival, seven monthly cohorts of newly-settled bluehead wrasse Thalassoma bifasciatum were collected immediately after settlement and over sequential 3-day periods. Otolith analysis was used to quantify mean larval and juvenile growth rates, pelagic larval duration (PLD), and settlement size and condition of different age classes to identify the traits most important for survival. Overall, survivors tended to have shorter PLDs, to settle at smaller sizes and higher condition levels, and to exhibit faster early juvenile growth. Water temperature contributed to among-cohort variability in traits as warmer water led to faster larval and juvenile growth and shorter PLDs. Trait-specific fitness functions demonstrated that temperature can influence fitness by changing the nature of selection on each trait. Estimates of selection intensity revealed that settlement condition contributed the most to variation in fitness across cohorts, followed by juvenile growth. Frequent loss of low settlement condition individuals and occasional loss of the very highest condition fish suggest that particularly high settlement condition during the warmest temperatures may be detrimental. Not only does the quality of settlers vary over time, but selective loss of individuals with particular phenotypic traits is not pervasive and can vary with environmental conditions such as temperature.     

Role of habitat in mediating mortality during the post-settlement transition phase of temperate marine fishes

The transition phase describes a distinct post-settlement stage associated with the recruitment to benthic habitats by pelagic life stages. The habitat shift is often accompanied by feeding shifts and metamorphosis from larval to juvenile phases. Density-dependent settlement, growth and mortality are often the major factors controlling recruitment success of this phase. Habitat use also becomes more pronounced after settlement. The role of habitat-mediated post-settlement mortality is elucidated by focusing on the early life history of Atlantic cod ( Gadus morhua ) and cunner ( Tautogolabrus adspersus ) in the north-west Atlantic. In these species, settlement can occur over all bottom types, but habitat-specific differences in post-settlement mortality rates combined with size and priority at settlement effects on growth and survival determine recruitment and eventual year-class strength. These results and those from other temperate marine fish species along with work on tropical reef species emphasize the generality of habitat-based density-dependent mortality during the transition phase and its potential for population regulation. These results have implications for fisheries management and can be used to outline a procedure to assist managers in identifying and managing essential transitional habitats including the potential role of marine protected areas in habitat conservation.     

Temperature and salinity effects on post-marsupial growth of Neomysis integer (Crustacea: Mysidacea)

There has been an increasing interest in using the brackish water mysid Neomysis integer as a toxicological test species for Western European estuarine systems. In this respect, more data on growth, moulting and development in this species is needed. The influence of prevailing environmental variables (e.g. temperature, salinity) and age on these processes as well as their optimal range have to be known in order to develop optimal laboratory cultures and to differentiate between chemically induced variability and natural variability in toxicity testing. Individual post-marsupial growth (size, intermoult period, growth factor) was studied from first day neonates until adulthood at eight environmentally relevant temperature-salinity conditions. Three salinities (5, 15 and 30 psu) were tested at 15 and 20 °C, and two more extreme temperatures (8 and 25 °C) were tested at a salinity of 5 psu.Survival and growth of N. integer were detected within the whole range tested, but sexual maturation was only possible in the narrower range of 15-25 °C and 5-15 psu. The size at maturity of N. integer increased with decreasing temperature and increasing salinity. Salinity seems to have a stronger effect than temperature on the duration of maturation. The sigmoid von Bertalanffy growth model was fitted to the individual and pooled data, except for the 8 °C experiment where growth was linear. Estimates from pooled data were comparable with individually based estimates, but generally underestimated the asymptotic length. Temperature was negatively correlated with the asymptotic length and positively correlated with the growth constant K. Higher temperatures caused smaller intermoult periods but had no effect on the growth increment, while salinity effects were less straightforward and dependent on the water temperature. A tool is provided to estimate the age, moult number, intermoult period, growth factor and growth rate from the body standard length of N. integer. Experimentally derived von Bertalanffy parameter estimates resulted in a higher growth performance index compared with field-based estimates for the Schelde estuary and Galgenweel populations of N. integer.