Phenotype-environment matching in the shore crab (Carcinus maenas)

The shore crab (Carcinus maenas) exhibits a range of carapace pattern polymorphisms, but little is known regarding their function or maintenance. If patterns represent some form of crypsis, then associations between carapace colouration and substrate are expected; to determine whether such relationships exist, frequency of crab morphs and quantity of substrate type were measured from fifteen 10×40 m² quadrats at each of three sites along the southern shore of the Firth of Forth, Scotland. Five thousand one hundred and thirty-seven crabs and 3.6 km of line intercept transect data were collected during a 9-week period. Crab abundance, relative frequency of morphs and substrate type varied significantly among the three sites. Plain crabs were strongly associated with macro-algal substrates whereas patterned crabs were associated with mussel beds. This pronounced phenotype-environment matching, as well as various characteristics of the carapace patterns themselves, suggests that patterned crabs are cryptic on polychromatic backgrounds. The frequency of patterned crabs and the percentage of white pigment on the carapace both declined significantly with carapace width. The loss of pattern coincides with an ontogenetic shift in habitat use and we present evidence to suggest that individual crabs lose their pigment, rather than larger patterned crabs being preferentially removed from the population by predators. Throughout their life history, shore crabs encounter high variation in predation, food supply, and physical habitat; to survive they have evolved a strategy that includes elements of pattern polymorphism, crypsis, ontogenetic shifts, and plastic responses.     

Epibiotic community of the horseshoe crab Tachypleus gigas

Horseshoe crabs act as moving substrata for simple to complex communities of small marine organisms. Amplexed adult pairs migrate for breeding once every 2 weeks from deep waters towards nearshore waters during highest high tide. Female horseshoe crabs bury themselves to the level of the lateral eyes to deposit eggs while the male crabs fertilize them. Subsequently eggs are buried by the female. Tachypleus gigas (Müller) is the most abundant horseshoe crab species above available along the Orissa coast (India). Adults reach terminal anecdysis once sexually mature and live with their carapace for 4 to 9 years. In spite of this, epibiosis is limited. In the current investigation, differences in the epibiotic community (diatoms and macro-epibionts) present on horseshoe crabs, according to gender, were evaluated, and the macro-epibiont population from different regions of the carapace was mapped. In general, female horseshoe crabs harbored fewer epibionts than the males. Among the diatoms, Navicula spp., Nitzschia spp. and Skeletonema sp. were dominant in both sexes. However, the abundance and diversity of diatoms was greater on the carapaces of male crabs. Among the macro-epibionts, the acorn barnacle (Balanus amphitrite Darwin) and encrusting bryozoan (Membranipora sp.) were the most dominant forms. Barnacles and bryozoans were greater in abundance in the “rough” zone (cardiopthalmic region and anterior region of the opisthosoma). Mapping of the macro-epibionts from different regions of the carapace revealed differential distribution in males and females. Such differentiated distribution of the macro-epibionts can be related to factors such as changing habitat by the horseshoe crabs during breeding, mechanical abrasion and surface availability during mating and nesting periods, requirements of epizootic larvae and surface properties of the carapace (wettability and roughness). In the case of females, mechanical abrasion and surface availability played an important role in the epibiotic community structure and distribution patterns. The surface wettability measurements indicated male carapace to be slightly more hydrophobic than the female carapace. Scanning electron microscopy revealed that the male carapace was comparatively rough compared to the smooth carapace of females. A comparison of surface properties of the carapace indicated that the male carapace is more conducive for epibiosis. Received: 23 August 1999 / Accepted: 25 January 2000     

Habitat selectivity of megalopae and juvenile mud crabs (<Emphasis Type="Italic">Scylla serrata</Emphasis>): implications for recruitment mechanism

Megalopae of several crab species exhibit active habitat selection when settling. These megalopae usually select structurally complex habitats which can provide refuge and food. The portunid mud crab, Scylla serrata, is commonly found within the muddy estuaries of the Indo-West Pacific after attaining a carapace width >40 mm. Despite substantial efforts, the recruitment mechanism of juvenile mud crabs to estuaries is not understood because their megalopae and early stage crablets (carapace width <30 mm) are rarely found. We used laboratory experiments to determine whether megalopae and early stage crablets are selective among three estuarine habitats which commonly occur in Queensland, Australia. These animals were placed in arenas where they had a choice of habitats: seagrass, mud or sand, and arenas where they had no choice. Contrary to the associations exhibited by other portunid crab megalopae, S. serrata megalopae were not selective among these estuarine habitats, suggesting that they tend not to encounter these habitats, or, gain no advantage by selecting one over the others. The crablets, however, strongly selected seagrass, suggesting that residing within seagrass is beneficial to the crablets and likely increases survival. This supports the model that for S. serrata, crablets and not megalopae tend to colonise estuaries, since a selective behaviour has evolved within crablets but not megalopae.     

Relationships between sponges and crabs: patterns of epibiosis on Inachus aguiarii (Decapoda: Majidae)

The association between sponges and the crab Inachus aguiarii Brito Capello, 1876 was studied by analysing the relationships between sponge distributional patterns on the crab carapaces and several morphological and biological crab parameters. Juveniles, mature females and mature males were differentiated on the basis of sex dimorphism and terminal pubertary moult. All three groups were fouled to different degrees by sponges. Percent sponge cover was related to carapace size only in mature males whereas mature females, all but one of which were ovigerous, were extensively covered regard-less of their size. It is proposed that some behavioural patterns unique to females, such as long resting periods in sponge-rich microhabitats, are responsible for these high sponge covers in females. Sponges showed two trends in the colonization of the carapace, leading to either a monopolizing or a sharing of the available carapace surface. The sponge species found on the carapaces studied are not obligatory epibionts of crabs, but are believed to reflect the sponge population characterizing the crab home range.     

Weapon size versus body size as a predictor of winning in fights between shore crabs, Carcinus maenas (L.)

Relative body size (carapace width) and weapon size (chela length) were used as indicators of resource holding potential (RHP) in the agonistic behaviour of male shore crabs, Carcinus maenas (L.). Weapon size was found to be a more reliable predictor of the outcome of pairwise fights than body size. Crabs with longer chelae than their opponents were more likely to win fights than crabs with relatively larger bodies. Body size had less influence on the outcome of fights. Relative body and weapon size did not influence initiation of contests but did affect the likelihood of winning; however, this was significant only for weapon size. Winning crabs had heavier claws with greater surface area than losing crabs. There was no relationship between relative size and fight duration. The frequency of cheliped display increased with chela length and win- ners performed significantly more displays than losers. Received: 5 February 1997 / Accepted after revision: 20 May 1997     

Rhythmic activity responses of the fiddler crab Uca crenulata to artificial tides and artificial light

The fiddler crab Uca crenulata, from California (USA), was exposed to artificial tides in order to differentiate between the influence of a tidal cycle from that of a light-dark cycle on its locomotor activity. Most crabs could be well synchronized by tides, but the activity patterns of other crabs was merely exogenously reinforced. Under constant conditions, after tidal treatment, crabs showed bimodal or unimodal activity patterns. When a 24 h light-dark cycle and a 12.35 h tidal cycle acted as concurrent stimuli, their effectiveness in synchronizing the crabs' activity was variable, depending on the undividual. It is likely that crabs which tend to exhibit a unimodal activity pattern are more sensitive to a light stimulus, whereas crabs with a bimodal activity pattern respond preferably to the tides. Thus, responses of endogenous activity of U. crenulata to tides and light-dark cycles are similar, suggesting that the basic oscillarory mechanism for circadian and tidal activity may be the same.     

Influence of cadmium accumulation and dietary status on fatty acid composition in two colour forms of shore crabs, Carcinus maenas

 The effects of cadmium exposure and dietary status on cadmium accumulation, fatty acid (FA) content and profiles were investigated in two colour forms of the shore crab Carcinus maenas. Groups of shore crabs were either starved or fed with blue mussels, Mytilus edulis, during a 40 d exposure period to 2 or 6 μM Cd²⁺ (as CdCl₂). Starved green individuals accumulated more cadmium in haemolymph and hepatopancreas than did red crabs and green crabs fed during the experiments. In the red colour form, no difference in cadmium accumulation was observed between starved and fed individuals. In both colour forms, hepatopancreas contained more FA than gills and muscle. The FAs often present in the largest amounts in the tissues were 16:0, 16:1ω7, 18:1ω7, 18:1ω9, 20:4ω6, 20:5ω3 and 22:6ω3. However, saturated (SAFA) and mono-unsaturated fatty acids (MUFAs) were dominant in hepatopancreas, whereas poly-unsaturated fatty acids (PUFAs) were dominant in gills and muscles. At the beginning of the experiment, the total FA content in the hepatopancreas was 111.6 mg g⁻¹ (dry weight) for red crabs and 78.4 mg g⁻¹ for green shore crabs. During the experiment, however, the FA content decreased in red crabs. This decrease was more pronounced for starved individuals than for fed individuals. Also, the decrease in FA content was more pronounced in crabs exposed to 6 μM cadmium compared to crabs exposed to 2 μM or crabs not exposed to cadmium. No change in FA content was observed in green shore crabs, irrespective of diet and cadmium exposure. For both colour forms, no change in FA content was observed for gills and muscle. In red crabs, a decrease was observed for all FAs in the hepatopancreas. This decrease, however, was more pronounced for SAFAs and MUFAs than for PUFAs, indicating that the metabolism of FAs during starvation and cadmium exposure is selective. The experiments indicate that green colour forms of shore crabs are more tolerant of natural stress such as starvation and anthropogenic stress, e.g. cadmium exposure, than are red colour forms of shore crabs. Received: 23 September 1999 / Accepted: 29 April 2000     

Seasonal variations of ATPase activity and antioxidant defenses in gills of the mud crab <Emphasis Type="Italic">Scylla serrata</Emphasis> (Crustacea,Decapoda)

The mud crab Scylla serrata is an important commercial crustacean being widely distributed along the southeastern coast of China. This crab and particularly its gills are subjected to seasonal fluctuations of environmental factors due to direct exposure to seawater. The investigation of seasonal variations of ATPase and antioxidant defenses in gills could be helpful for the understanding of physiological regulation mechanism of seasonal adaptations. In this study, mud crabs were collected from the subtropical waters near Xiamen island, Southeast China (24°26′46′′N, 118°04′04′′E), in August and November, 2002, and February and May, 2003, respectively, being considered as specimens from summer, autumn, winter and spring, respectively. Only healthy intermoult male crabs without carapace or appendage damage were used, having a carapace width of 7.05 ± 0.52 cm and a wet weight of 130 ± 20 g. The activities of four ATPases and three antioxidation enzymes (superoxide dismutase, catalase, glutathione peroxidase) and the content of malondialdehyde in gills were measured. The results showed that the activities of Na⁺, K⁺-ATPase, Mg²⁺-ATPase, Ca²⁺-ATPase and Ca²⁺, Mg²⁺-ATPase reached maximum levels similarly in summer and decreased in winter. The activities of superoxide dismutase and catalase were higher in summer and lower in winter, with a significant decrease in winter compared to the other seasons (p < 0.01). Glutathione peroxidase activity was higher in summer, and lower in autumn, being significantly higher in summer or winter than in spring or autumn (p < 0.01). Malondialdehyde content was higher in summer and lower in spring with significant differences among the different seasons (p < 0.01). In summary, the obvious seasonal activity variations of four ATPases and antioxidant enzymes and the content of malondialdehyde reflect a seasonal regulation of the physiological metabolising enzyme and the antioxidant capacity to cope with seasonal alterations of environment factors such as fluctuating salinities and temperatures.     

The biology of the blue manna crab Portunus pelagicus in an Australian estuary

Various aspects of the biology of the blue manna crab Portunus pelagicus have been investigated in the large Peel-Harvey estuarine system of Western Australia, using samples collected regularly by beach seine, gill net and otter trawl between February 1980 and July 1981. Whereas crabs were widely dispersed throughout Peel Inlet, Harvey Estuary and the saline regions of tributary rivers during the summer and autumn, they were found mainly near the estuary mouth in the winter and spring. Since our data suggest that P. pelagicus has a preference for salinities of 30 to 40, the above changes in distribution are apparently related to the marked seasonal variation in salinity which results from the very seasonal pattern of rainfall. The number of ovigerous crabs in the estuary were greatest in January and February. The mean carapace width and number of eggs of ovigerous females were 110 mm (range 85 to 157 mm) and 509 433 (range 270 183 to 847 980), respectively. P. pelagicus started to reach the minimum legal size for capture (carapace width 127 mm) in the summer when they were approximately I yr old, and left the system in large numbers in the following winter when they were 15 to 20 mo old. These features explain why the fishery for P. pelagicus is highly seasonal, with the vast majority of crabs being taken between January and May. As crabs approached the end of their first year of life, the ratio of females began to exceed that of males, apparently as a result of the movement of males out of the system and legislation against the capture of ovigerous females.     

Interindividual variability in metal status in the shore crab<Emphasis Type="Italic"> Carcinus maenas</Emphasis>: the role of physiological condition and genetic variation

The relations between genetic variation, physiological condition, size and metal status in the shore crab Carcinus maenas (L.) were investigated. Shore crabs were collected at three sites around the island of Funen, Denmark, and carapace width, colour, moulting stage and major ions and protein in the haemolymph were determined. Concentrations of water, cadmium, copper and zinc in midgut glands, muscles and gills were measured, and allozyme variation at 15 loci was studied. Generally, tissue water content and copper, zinc and cadmium concentrations decreased from the moulting stages C₃ over C₄ to D crabs, whereas the opposite was true for copper and zinc concentrations in gills and muscle. However, the water content of the tissues increased with the size of the crabs. Since tissue water contents changed consistently over moulting stages, conclusions on changes in metal concentrations sometimes diverged when expressed on a dry versus wet weight basis. Regarding allozymes, esterase, malate dehydrogenase and phosphoglucomutase were polymorphic. The crabs showed a low level of genetic variability for both polymorphism (0.14) and for heterozygosity (0.07) at the three sites. Allelic frequencies showed very low levels of differentiation among the samples, no evidence of inbreeding and no population subdivision. Multilocus heterozygosity was positively correlated with size. Concentrations of copper and zinc in midgut gland, muscle tissue and gill were not correlated with genotype, whereas a weak correlation between phosphoglucomutase genotypes and concentrations of cadmium in the midgut gland was found.Communicated by M. Kühl, Helsingør     

Significance of mangrove–mudflat boundaries as nursery grounds for the mud crab, Scylla paramamosain

Fisheries managers frequently try to protect juveniles in order to preserve stocks. Juveniles can be protected by either implementing changes designed to avoid catching immature animals (e.g. increasing mesh size or altering fishing techniques) or protecting nursery grounds. To prevent the capture of immature animals, an estimate of size at maturity is required as well as a knowledge of both fishing methods and the exact location of the nursery grounds. Strong demand for juvenile mud crabs to stock aquaculture ponds has resulted in development of fisheries targeting crabs of all sizes from instar 1 to mature individuals. Using five different fishing methods, different stages in the life cycle of Scylla paramamosain were followed for a period of 16 months in an estuarine population in the Mekong Delta, Vietnam. Mangrove habitat utilisation begins when crabs settle out from the plankton at instar 1 [modal internal carapace width (ICW), 0.5 cm] amongst the pneumatophores at the mangrove fringe. Increasingly larger crabs were found deeper into the mangrove but they were still living on the surface (modal ICW size class, 1.5 cm). As their size increases, the crabs either dig burrows (modal ICW size class, 4.5 cm) or they live in the sub-tidal zone, migrating into the mangrove with each tide to feed (modal ICW size class, 4.5 cm). Larger crabs were caught offshore (modal ICW size class, 12.5 cm) where females accounted for 60% of the catch although of these, only 63% were mature. Recruitment of early instars was continuous but peaked in December to February. Subsequent peaks in the catch rates of larger size classes indicated the development of a single cohort with an estimated growth rate of 2.0 cm ICW per month. On the basis of abdominal width, females were estimated to mature at 10.2 cm ICW although at 9.7 cm ICW, 50% of females had disengaged abdomens. Abdominal disengagement occurred in males at the slightly smaller size of 9.1 cm ICW. Allometric relationships between chela height and carapace width suggested 50% of males acquire mature chelae at 10.2 cm ICW. These results demonstrate the close linkage between early life stages of S. paramamosain and certain specific niches within mangrove habitats, with the main adult population found to be living sub-tidally at some distance from the mouth of the estuary. The study also highlights the special importance of the mangrove fringe, the border between the mangrove forest and the sea, an area which is particularly vulnerable to physical and anthropogenic impacts.     

Na-K-ATPase generates an active transport potential in the gills of the hyperregulating shore crab Carcinus maenas

Perfusing and bathing isolated gills of shore crabs Carcinus maenas with artificial saline or sea (brackish) water enabled us to determine potential differences (PDs) between ambient bathing medium and perfusion solution. Establishment of diffusional PDs was avoided by employment of the same solution on the internal and external side. The PDs measured were therefore of an active nature. We compared the properties of the PDs with the well-known properties of the Na-K-ATPase: dependence on biological energy (ATP), on salinity and sodium concentration, susceptibility of PD to depletion of internal K and to the addition of 5 mM internal ouabain. Considering also the magnitude of the PDs measured, the results obtained indicate that it is the Na-K-ATPase that generates an active transport potential for Na in the gills of shore crabs. This PD represents the driving force for the active uptake of Na in crabs that hyperregulate their body fluids when they inhabit environments of reduced salinity regimes or fluctuating salinities in tidal estuaries. This process counteracts diffusional losses of Na in crabs exposed to dilute media.     

Foraging behavior of juvenile Carcinus maenas (L.) and Cancer pagurus L.

Information concerning the way juvenile crabs choose their diet from a variety of prey types can be useful for a better understanding of community dynamics, as well as for the adequate management of natural resources. Prey size and species selection by juvenile Carcinus maenas (15-35 mm carapace width, CW) and Cancer pagurus (20-40 mm CW) feeding on four bivalves of contrasting shell morphology were investigated. When offered a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas, and Cerastoderma edule presented individually, crabs generally showed evidence of size-selective predation. Cancer pagurus selected larger mussels relative to the size of their chelae (relative prey size, RPS) than did Carcinus maenas of similar and even larger carapace width. However, the RPS of selected O. edulis and Cerastoderma edule were similar for all crabs, suggesting that certain prey features constitute effective barriers even to the powerful chelae of Cancer pagurus. When offered a wide size range of mussels and oysters simultaneously, all crabs consistently selected mussels. When offered O. edulis and Crassostrea gigas, crabs consumed both these oyster species in similar numbers. Carcinus maenas consumed similar numbers of mussels and cockles; Cancer pagurus, however, showed no preference for either prey in the smaller size classes but selected more mussels than cockles as prey increased in size. Although previous studies report that adult Carcinus maenas select prey species according to their profitability (amount of food ingested per unit of handling time, milligrams per second), consumption rates of the size classes of prey selected by juvenile shore crabs did not always parallel prey value. Although variations in crab strength can account for many of the differences between the foraging strategy of juvenile and adult C. maenas, our results suggest that juvenile crabs are less species selective than adults as a result of the restrictions imposed on small individuals that have limited access to larger prey.     

Intra-specific variability in the temporal organisation of foraging activity in the limpet<Emphasis Type="Italic"> Patella vulgata</Emphasis>

Patella vulgata is one of the most common limpet species on rocky shores throughout the north-eastern Atlantic. Foraging activity of this species has been described during various different diel and tidal phases and several factors have been suggested that might influence the timing of its activity. Among these, inclination of rock surfaces seems to have a clear role, with limpets on horizontal rocks being active during daylight at high water, while limpets on vertical rocks were active during night-time low water. However, by comparing the results of previous investigations it is difficult to separate clearly the effect of any single factor from the effects of their interactions. In the present paper we investigated the simultaneous influence of height on the shore, slope of rock surface and date of sampling (August–September and March–April) on the behaviour of a population of P. vulgata on the Isle of Man (UK). The study was carried out using modern telemetric methods, allowing the activity patterns of limpets to be recorded for long periods of time. The results showed that limpets concentrated their activity during two distinct temporal windows—daytime tide-in and night-time tide-out—while no significant activity was observed during daytime tide-out or night-time tide-in. The observed behaviour appeared to be driven by complex interactions among the three factors examined, and the influence of rock slope alone was not so strong as to override the effect of other factors completely. In addition, a key finding of this study was the conspicuous variability in the individual behaviour of P. vulgata. The greatest level of variability was observed in limpets on vertical surfaces on the upper shore, which showed an extreme switch in behavioural patterns, according to date of sampling (daylight/tide-in foraging during the spring and night-time/tide-out foraging during late summer). In addition, individuals apparently sharing the same set of conditions (inclination, height on the shore and sampling date) differed in their behaviour, and a given behavioural pattern was not always shared by all individuals from a given group. Finally, variability was also observed at the intra-individual level since the same individual could be active during different tidal and diel phases. In conclusion, although the existence of endogenous rhythms has previously been demonstrated in this species, our data clearly show that P. vulgata was able to modulate its behavioural patterns in relation to a variety of exogenous determinants.Communicated by R. Cattaneo-Vietti, Genova     

Feeding habits of the dungeness crab Cancer magister as determined by the index of relative importance

Specimens of the Dungeness crab Cancer magister, were collected subtidally and intertidally from an estuary in Washington State, USA in June and September 1980, and January, April, May and July, 1981. Gut contens of freshly collected crabs were analyzed by the Index of Relative Importance; for each prey taxon, this method measured frequency of occurrence, percentage of total biomass, and percentage of total numbers consumed. The most important higher taxon eaten was fish; however, the most important prey genus was the shrimp Crangon spp. There was greater predation on Crangon spp. at night at the intertidal site, and during winter and spring when the shrimp were most abundant there. Feeding activity, as indicated by a weight-specific gut-fullness index, showed no consistent diel pattern. There were significant ontogenetic changes in feeding patterns: first-year crabs preyed primarily on very small bivalves or small crustaceans including their conspecifics; second-year crabs preferred Crangon spp. and fish, and third-year crabs preyed less on Crangon spp. and more on fish. Such changes in feeding habits with ege could be purely due to mechanics of food handling, but might also reduce competition among age groups of crabs, possibly partitioning resources within the estuary. Findings are discussed in terms of optimal foraging and compared to other similar studies.Contribution No. 599, School of Fisheries, University of Washington, Seattle, Washington 98195, USA     

Population ecology of the mud crab <Emphasis Type="Italic">Scylla paramamosain</Emphasis> (Estampador) in an estuarine mangrove system; a mark-recapture study

Mud crabs of the genus Scylla are commercially important mangrove residents that are extensively fished throughout their range in the Indo-West Pacific. Despite this high level of exploitation very little is known about the population dynamics of any of the Scylla species. The present study concentrated on an exploited population of Scylla paramamosain in a natural estuarine mangrove on Can Coc Island in the mouth of the Hau River, Mekong Delta, Vietnam. A total of 6,114 juvenile crabs captured on the seaward mangrove fringe were internally tagged by injection of coded microwire tags and released over a period of 29 days. Recaptures were monitored over the following 144 days. There was little migratory movement within the estuary; of 285 recaptured crabs, 93% were recovered within the island mangrove study area and only two individuals were recovered from mangroves lining the opposite mainland riverbanks. A von Bertalanffy growth function was fitted to growth increment data from recaptured crabs, with L _max = 150 mm to derive a growth constant, k = 2.39 year⁻¹. The theoretical age at which carapace width is zero (t ₀) was derived from known size at recruitment at instar 1, giving a value of −0.0095 years. Previous studies of the same population have shown that female S. paramamosain reach maturity at a mean size of 102 mm carapace width. The present study indicates that they attain this size at around 160 days from first settlement in the mangrove fringe. Abundance of juvenile crabs in the study area was estimated by the Petersen method as 1,101,500 (95% CI 4,17,300–1,785,800, representing 1,102 crabs ha⁻¹ of mangrove. Estimation of mortality from tag returns and from the age-catch curves during a period of constant recruitment were comparable (Z = 1.11 and 1.04 month⁻¹, respectively). Fishing only accounted for 14% of total mortality, suggesting that at the time exploitation was not at a critical level despite the apparent high level of fishing activity.     

Effect of acanthocephalan parasites on the behaviour and coloration of the mud crab Macrophthalmus hirtipes (Brachyura: Ocypodidae)

In the field, the numbers of cystacanths of the parasitic acanthocephalan Profilicollis spp. harboured by crabs are relatively high and correlate with carapace width. In a field experiment, the responses of crabs to the simulated approach of a bird predator (the parasite's definitive host) was not influenced by the number of acanthocephalans they harboured. Crabs that were exposed at the surface of the sediments during receding high tide, however, tended to harbour more parasites than nearby crabs hidden in burrows. An analysis of colour patterns on the carapace of crabs showed that infection levels did not influence carapace pigmentation, and thus did not affect the conspicuousness of a crab relative to the background environment. However, the likelihood of a male crab winning a ritualized fight against a conspecific in the field was associated with its infection level, but in a way that suggests that this finding is a consequence of pathology rather than an adaptation of the parasite to increase its transmission rate. Although only weak evidence was found indicating that Profilicollis manipulates the behaviour or colour of its host to its own benefit, the high infection levels observed suggest that the crab population acts as a major reservoir for larval stages of this parasite that are infective to birds.     

Temporal and spatial settlement patterns of sympatric hermit crabs and the influence of shell resource availability

Larvae of marine organisms often need specific resources or environments at settlement, and their success at settlement might be strongly influenced by the abundance and distribution of such specific resources. The larvae of hermit crabs need small shells to settle, so it is thought that the distribution and abundance of small shells influence the settlement pattern of hermit crabs. To investigate the influence of small shell distribution on the settlement of pagurid hermit crab larvae, we conducted a field experiment at an intertidal rocky shore in Hakodate Bay, Japan. From the line-transect sampling in the field, we found that Pagurus middendorffii settled extensively in the offshore side of the intertidal zone while P. nigrofascia settled in the uppermost area of the intertidal zone. Small shells were most abundant in a narrow shallow trough, slightly offshore from the uppermost area of the intertidal zone. For both species, settler abundance was high where adults were abundant, but settler abundance did not appear to be related to shells abundance. An experiment to clarify settlement patterns showed that larval recruits tended to be similar to those in the line-transect sampling of settlers. Thus shells may not be a primary factor affecting settlement patterns at relatively large scale within the intertidal flat. However, when we analyzed the relationship of settlers and shells separately within each transect, the distribution of settlers was well explained by shell resource availability. Therefore on a smaller scale, shell availability may influence the number of settlers. Settlement periods of P. middendorffii and P. nigrofascia fully overlapped, so their larvae probably were affected by similar transport factors, such as current and tidal movement. Nevertheless they showed different spatial patterns of settlement.Communicated by T. Ikeda, Hakodate     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

Field and experimental studies on cadmium in the edible crab Cancer pagurus

The distribution of cadmium within captive crabs (Cancer pagurus) exposed experimentally to cadmium-contaminated food and water is described and illustrated by triangular diagrams. Crabs from the Orkney Islands (Scotland) are known to contain relatively high levels of cadmium (up to 62 g g^-1 wet wt) in the hepatopancreas. The distribution of cadmium between the hepatopancreas, gonad, gill, carapace, claw muscle, heart, and haemolymph, is described in crabs collected during 1978, and compared with similar data from crabs exposed to cadmium for ca. 300 d (September 1978 – June 1979) experimentally. It is concluded that the dominant uptake route of cadmium to Orkney crabs is through their diet.