Dynamics of shell disease in the edible crab Cancer pagurus: a comparative study between two sites on the Gower Peninsula,South Wales,UK

Shell disease syndrome is a degradative condition of the crustacean exoskeleton which results in the formation of black-spot lesions. Field surveys on the edible crab Cancer pagurus (L.) population at 2 sites in the Gower Peninsula of South Wales, UK, namely Langland Bay and Rhossili causeway, revealed similarities in the prevalence of shell disease but differences in the severity. Male crabs from Langland Bay displayed significantly greater severities of the disease on the dorsal carapace and ventral surfaces than those in Rhossili. The greatest severities of the disease in Langland were attributable to sand abrasion injuries during back-burrowing behaviour and prolonged exposure to high numbers of chitinolytic bacteria. Lower severities of shell disease in males from Rhossili were a reflection of the inability to back-burrow due to the nature of the onshore sediment, and hence the crabs were only exposed to low levels of chitinolytic bacteria in the water column. On areas of the causeway that permitted back-burrowing, the chitinase enzyme activities in sediments were significantly lower than those in Langland Bay. It is concluded that the potential for crustacean shell degradation is higher at Langland than Rhossili, but whether the differences in chitinolytic abilities of the sediments at the 2 sites are a result of differences in pollution levels or simply of different substratum types, remains to be determined.     

A histological study of shell disease syndrome in the edible crab Cancer pagurus.

Shell disease syndrome is characterised by the external manifestation of black spot lesions in the exoskeletons of crustaceans. In the present study, gills, hepatopancreas and hearts from healthy (<0.05% black spot coverage) and diseased (5 to 15% coverage) edible crabs, Cancer pagurus, were examined histologically to determine whether this disease can cause internal damage to such crabs. There was clear evidence of cuticular damage in the gills of diseased crabs leading to the formation of haemocyte plugs termed nodules. Nephrocytes found within the branchial septa of the gills showed an increase in the accumulation of dark material in their vacuoles in response to disease. In the hepatopancreas, various stages of tubular degradation were apparent that correlated with the severity of external disease. Similarly, there was a positive correlation between the number of viable bacteria in the haemolymph and the degree of shell disease severity. Approximately 21% of the haemolymph-isolated bacteria displayed chitinolytic activity. Overall, these findings suggest that shell disease syndrome should not be considered as a disease of the cuticle alone. Furthermore, it shows that in wild populations of crabs shell perforations may lead to limited septicaemia potentially resulting in damage of internal tissues. Whether such natural infections lead to significant fatalities in crabs is still uncertain.     

Epidemiology of bitter crab disease (Hematodinium sp.) in snow crabs Chionoecetes opilio from Newfoundland, Canada

The parasitic dinoflagellate Hematodinium sp. causes a condition known as bitter crab disease (BCD) in snow crabs Chionoecetes opilio and Tanner crabs C. bairdi. As the name of the condition implies, crabs infected with BCD are unmarketable due to their bitter flavor. We surveyed the distribution of BCD in 3 regions within the snow crab fishery of Newfoundland from 1997 to 2003. Over time, the disease has become firmly established in Conception and Bonavista Bays and persists at low levels on the Avalon fishing grounds. An epizootic occurred within Bonavista and Conception Bays in 1999 and persisted in Conception Bay in 2000, reaching prevalences of over 2% to 9% in trapped and trawled male crabs and from 19 to 26% in trawled and trapped female crabs, respectively. Infections were highest in females and small males, i.e. the unfished and pre-recruit portions of the fishery. In a mortality study, all of the naturally infected crabs died and 50% of the experimentally inoculated crabs died. Patterns in the molting cycle and prevalence of infection indicate that transmission occurs during the post-molt condition, and that overt infections probably develop 2 to 4 mo later with mortalities occurring at least 3 to 4 mo thereafter. The hydrography of this bay may have contributed to the epizootic as infections were centered within the deeper confines of the bay. Analysis of various abiotic factors uncovered a significant positive association between prevalence, depth and mud/sand substrates; the nature of this relationship was not apparent but may be related to diet or alternate hosts. Lastly, given the increase in BCD in snow crabs in Newfoundland, we recommend that fishery management programs for Chionoecetes fisheries employ non-selective gear to monitor for Hematodinium infections in female and juvenile crabs because these under-sampled members of the population may forewarn of impending recruitment declines that might otherwise remain unexplained.     

Bacteria associated with crabs from cold waters with emphasis on the occurrence of potential human pathogens.

A diverse array of bacterial species, including several potential human pathogens, was isolated from edible crabs collected in cold waters. Crabs collected near Kodiak Island, Alaska, contained higher levels of bacteria than crabs collected away from regions of human habitation. The bacteria associated with the crabs collected near Kodiak included Yersinia enterocolitica, Klebsiella pneumoniae, and coagulase-negative Staphylococcus species; the pathogenicity of these isolates was demonstrated in mice. Although coliforms were not found, the bacterial species associated with the tissues of crabs collected near Kodiak indicate possible fecal contamination that may have occurred through contact with sewage. Compared with surrounding waters and sediments, the crab tissues contained much higher proportions of gram-positive cocci. As revealed by indirect plate counts and direct scanning electron microscopic observations, muscle and hemolymph tissues contained much lower levels of bacteria than shell and gill tissues. After the death of a crab, however, the numbers of bacteria associated with hemolymph and muscle tissues increased significantly. Microcosm studies showed that certain bacterial populations, e.g., Vibrio cholerae, can be bioaccumulated in crab gill tissues. The results of this study indicate the need for careful review of waste disposal practices where edible crabs may be contaminated with microorganisms that are potential human pathogens and the need for surveillance of shellfish for pathogenic microorganisms that naturally occur in marine ecosystems.     

Newcastle disease virus in double-crested cormorants in Alabama, Florida, and Mississippi.

In order to understand the epidemiology of Newcastle disease (ND) outbreaks in double-crested cormorants (Phalacrocorax auritus), a study was conducted on wintering migratory cormorants (P. a. auritus) in Alabama and Mississippi (USA) and non-migratory cormorants (P. a. floridanus) that breed in Florida (USA). Antibodies against ND virus were detected by the hemagglutination-inhibition method in sera from 86 of 183 (47%) migratory cormorants over-wintering in eight roosting sites in Alabama and Mississippi between November, 1997 and April, 1999. Titers ranged from 5 to 40. Antibody prevalences in sera collected from females in early winter (November and December) (26%) and late winter (February and March) (56%) were significantly different (P = 0.0007). None of 45 serum samples from 1- to 7-wk-old nestlings from 11 colonies in Florida during the 1997-98 and 1998-99 breeding seasons was positive. However, antibodies were detected in yolk samples from 98 of 126 (78%) eggs collected in these same colonies. Titers ranged from 4 to 256. The prevalence of antibodies in eggs collected from fresh-water colonies (63% prevalence, n = 30) and salt-water colonies (82% prevalence, n = 96) was significantly different (P = 0.041). ND virus was not isolated from tissues of 18 cormorants and cloacal and tracheal swabs from 202 cormorants collected in Alabama and Mississippi; virus was also not isolated from cloacal and tracheal swabs from 51 nestlings from Florida.     

The bitter crab syndrome in commercial crabs in the Western part of the Bering and Chukchi Seas

We studied the distribution of the “bitter-crab” syndrome, a disease caused by the parasitic dinoflagellate Hematodinium sp., in eight commercial species of crabs in the west of the Bering and Chukchi seas. The crabs (25 388 individuals) were sampled during bottom trawl surveys of July?September 2010 and October?November 2012. The disease was first identified visually by a color change of the exoskeleton and the hemolymph of the animals and then using microscope analysis of hemolymph samples. Infestation was detected in crabs of three species, Chionoecetes opilio, C. bairdi, and Paralithodes platypus. The prevalence of the disease (the percent of infected individuals relative to all of those examined) in C. bairdi and P. platypus was very low, 0.1 and 0.3%, respectively. Infestation was widespread among C. opilio, its peak in the Bering Sea was in the fall. The average prevalence of the crab disease in different areas of the Bering Sea ranged from 0.8 to 10.8%. A high rate of crab infection was recorded in the Korfa Bay. In the Chukchi Sea, the average prevalence was 2%. Infestation by Hematodinium sp. was not revealed in the deep-sea snow crabs Chionoecetes tanneri and C. angulatus, and in three species of lithodid crabs, Paralithodes camtschaticus, P. brevipes, and Lithodes couesi. This can be explained by the small sample volume and/or ecology of these species, since the disease was registered in other areas in four of them.     

Predation by blue crabs, Callinectes sapidus, on rapa whelks, Rapana venosa: possible natural controls for an invasive species?

Blue crabs Callinectes sapidus are voracious predators in Chesapeake Bay and other estuarine habitats. The rapa whelk Rapana venosa is native to Asian waters but was discovered in Chesapeake Bay in 1998. This predatory gastropod grows to large terminal sizes (in excess of 150 mm shell length (SL)) and has a thick shell that may contribute to an ontogenetic predation refuge. However, juvenile rapa whelks in Chesapeake Bay may be vulnerable to predation by the blue crab given probable habitat overlap, relative lack of whelk shell architectural defenses, and the relatively large size of potential crab predators. Feeding experiments using three size classes of blue crab predators in relation to a size range of rapa whelks of two different ages (Age 1 and Age 2) were conducted. Blue crabs of all sizes tested consumed Age 1 rapa whelks; 58% of all Age 1 whelks offered were eaten. Age 2 rapa whelks were consumed by medium (67% of whelks offered were eaten) and large (70% of whelks offered were eaten) blue crabs but not by small crabs. The attack methods of medium and large crabs changed with whelk age and related shell weight. Age 1 whelks were typically crushed by blue crabs while Age 2 whelk shells were chipped or left intact by predators removing prey. Rapa whelks less than approximately 35 mm SL are vulnerable to predation by all sizes of blue crabs tested. Rapa whelk critical size may be greater than 55 mm SL in the presence of large blue crabs indicating that a size refugia from crab predation may not be achieved by rapa whelks in Chesapeake Bay until at least Age 2 or Age 3. Predation by blue crabs on young rapa whelks may offer a natural control strategy for rapa whelks in Chesapeake Bay and other estuarine habitats along the North American Atlantic coast.     

Shell utilization,predation pressure,and thermal stress in panamanian hermit crabs: An interoceanic comparison

Shell preference patterns of two common hermit crabs from hard bottom reef flats on the Caribbean coast of Panama are examined in relation to the predation pressures and physical stresses of their habitat. Clibanarius antillensis Stimpson lives in the high intertidal habitat and minimizes exposure to predators by seeking refuge during high tides. It prefers high-spired shells which maximize protection from thermal stress. Calcinus tibicen Herbst avoids tidal emersion and prefers low-spired shells which enhance resistance to the predators common on Caribbean reef flats.The results are compared with similar results from the tropical eastern Pacific Bay of Panama. Shell-crushing predation on Caribbean hermit crabs is suggested to differ quantitatively and qualitatively from predation on hermit crabs in the Bay of Panama. Predation on hermit crabs in the Bay of Panama is more intense and effects larger individuals than predation on Caribbean reef flat hermit crabs. In addition, shell-crushing predation on hermit crabs in the Bay of Panama is primarily from teleost fish predators (Diodon spp.), while predation on Caribbean hermit crabs is primarily by bottom-dwelling crustaceans.Differences in predation pressures and tidal regimes between the Caribbean and Pacific coasts of Panama are reflected in the shell preferences and behavior of hermit crabs from the two areas.     

三疣梭子蟹蜕壳周期肝胰腺、外壳和鳃中钙含量的变化

研究测定了三疣梭子蟹蜕壳周期肝胰腺、鳃和外壳中的钙含量。结果显示,在不同时间,鳃中钙含量变化较小,各时间点差异不显著;蜕壳前期至蜕壳刚完成阶段三疣梭子蟹(软壳蟹)肝胰腺中钙含量最高,蜕壳后第4 d肝胰腺中钙含量最低。三疣梭子蟹外壳中钙含量随着外壳的硬化(钙化)逐渐增加,完全硬化后外壳中钙含量达到最高。研究表明,蜕壳后三疣梭子蟹肝胰腺中钙含量的降低与外壳的钙化之间具有关联性,肝胰腺在三疣梭子蟹蜕壳后钙的储存中起着重要作用,而鳃在钙离子储存中所起的作用可能小于钙转移的作用。     

Recent settlement trends in Panulirus argius (Decapoda: Palinuridae) pueruli around St. Thomas, U.S. Virgin Islands

Puerulus settlement of the western Atlantic spiny lobster, Panulirus argus, was monitored using modified Witham collectors from December 1996 to March 1998 at seven sites around St. Thomas, U.S. Virgin Islands. A total of 605 pueruli were collected from 553 samples for a catch per unit effort (CPUE) for all sites of 1.09 pueruli. The greatest settlement occurred on sites within a recently declared marine reserve, which had an overall CPUE of 1.77 pueruli. Settlement in non-reserve sites was much lower with an overall CPUE of 0.31 pueruli. Pueruli recruitment declined 67% at inshore sites and 53% at offshore sites between July 1992 - April 1994 and February 1997 - March 1998. Also, only 10% of months sampled in 1997-98 had a CPUE > 0.5 compared to 55% in a previous study in 1992 - 1993. Despite the decline in pueruli CPUE in 1997-98 compared to 1992-94, the commercial lobster catch in the 2000-01 fishing season, and by inference the adult lobster population (legal lobster size in the US Virgin Islands is > or = 3.5 cm carapace length), remained stable.     

Enumeration of Pseudomonas species and Pseudomonas aeruginosa bacteriophages in domestic sewage

Domestic sewage in Kuwait is mainly treated by an activated sludge process. Pseudomonas species were enumerated at all steps of sewage treatment. About 98-99% reduction in the number of these bacterial species were found in the treated effluent compared with raw sewage, which indicates a rather efficient removal of Pseudomonas from sewage. Spherical tail-less phages of Pseudomonas aeruginosa were found in all sewage samples. About 25-85% of the total phages encountered with the raw sewage were retained in the treated effluents. Seasonal variations of Pseudomonas spp and P. aeruginosa phages in two treatment stations are reported.     

Effect of extracellular products of Pseudoalteromonas atlantica on the edible crab Cancer pagurus

Previous studies have shown that injection of extracellular products (ECP) of Pseudoalteromononas atlantica isolated from shell disease-infected edible crabs (Cancer pagurus) into healthy crabs causes rapid death. In this study we examined the nature of the active lethal factor(s) in ECP. Injection of ECP into crabs caused a rapid decline in the total number of circulating hemocytes (blood cells), and the crabs died within 60 to 90 min. The individuals that died showed eyestalk retraction, limb paralysis, and lack of antennal sensitivity, suggesting that the active factor(s) targeted the nervous system. Histopathological investigations showed that affected crabs had large aggregates of hemocytes in the gills, and there was destruction of the tubules in the hepatopancreas. The active factor in ECP was not sensitive to heat treatment (100 degrees C for 30 min) and proteinase K digestion. As lipopolysaccharide (LPS) was a potential candidate for the lethal factor, it was purified from whole P. atlantica bacteria or ECP and subsequently injected into crabs. These crabs had all of the external symptoms observed previously with ECP, such as limb paralysis and eyestalk retraction, and they died within 90 min after challenge, although no significant decline in the number of circulating hemocytes was observed. Similarly, in vitro incubation of hemocytes with purified LPS (1 to 20 microg) from P. atlantica did not result in the clumping reaction observed with ECP but did result in a degranulation reaction and eventual cell lysis. Injection of crabs with Escherichia coli or Pseudomonas aeruginosa LPS (1 microg g of body weight(-1)) did not cause any of the characteristic symptoms observed following exposure to P. atlantica LPS. No mortality of crabs followed the injection of E. coli LPS, but P. aeruginosa LPS caused ca. 80% mortality at 2 h after injection. Overall, these results show that the main virulence factor of P. atlantica for edible crabs is LPS either alone or in combination with other heat-stable factors.     

The effect of a new sewage treatment plant on faecal indicator numbers, campylobacters and bathing water compliance in Morecambe Bay

Until recently, sewage from Morecambe was macerated, but otherwise untreated, and discharged at high water via a short outfall pipe into Morecambe Bay adjacent to a recognized bathing water. In March 1997, a new biological sewage treatment plant came on-line and the effluent was discharged via a longer outfall pipe into the Bay south of Heysham. The effect of the new sewage treatment on the quality of the bathing waters was monitored by testing sea water collected from the three EU designated bathing waters on Morecambe Bay: Morecambe North, Morecambe South and Heysham. After sewage treatment came on-line, the numbers of faecal coliforms and faecal streptococci were lower at Morecambe North and Morecambe South but higher at Heysham. Although the changes in numbers were not always statistically significant, they were sufficient to affect compliance with the EU Bathing Water Directive Imperative (mandatory) standards. Compliance improved markedly at Morecambe North and South but declined at Heysham, the closest bathing site to the new outfall. Numbers of thermophilic campylobacters were similar in both years, which is suggestive of their sources being different from those of the indicator bacteria. Campylobacter lari and urease-positive thermophilic campylobacters (UPTC) were the only species of Campylobacter isolated from Morecambe's bathing waters. Very low numbers of Salmonella were found, with Salm, arizonae the only species isolated.     

Nematode infection in Mugil incilis (Lisa) from Cartagena Bay and Totumo Marsh, north of Colombia

Nematode infection indices were recorded in Lisa, Mugil incilis, from Cartagena Bay and Totumo Marsh, north of Colombia, during an 8-mo period (February to September 2002). Parasite prevalence (74.67% vs. 53.48%), intensity (9.73 +/- 2.98 vs. 2.73 +/- 1.07), and abundance (7.49 +/- 2.21 vs. 2.04 +/- 1.17) were significantly greater in Cartagena Bay, an estuary polluted by domestic sewage and industrial discharge, compared with those of Totumo Marsh, a body of water with low levels of contamination. Parasites were found below the spine, within the liver and the intestinal mesenteries, and a small percentage in muscle. Morphological analysis of nematodes indicated the presence of the third larval stage of several species belonging to the Anisakidae. Size correlated moderately and significantly with parasite intensity in fish collected from Totumo Marsh (R = 0.336; P < 0.001); in Cartagena Bay the correlation was also significant, but low and negative (R = -0.212; P = 0.003), clearly showing differences in host-parasite ecology. Fish health status, as represented by condition factor and hepatosomatic index, did not show any correlation with the parasite prevalence in fishes collected in either sampling areas. These results suggest, for the first time, that the consumption of Lisa from the Atlantic coast of Colombia could represent a risk for human infection.     

滨鹬马蹄吸虫(复殖目:微茎科)的尾蚴和囊蚴期的季节动态

对科威特湾微茎科滨鹬马蹄吸虫幼虫期的中间宿主双带盾桑椹螺(Clypeomorus bifasciata)及小相手蟹(Nanosesarma minutum)的季节动态进行了研究。调查期超过一年,在检查的1 600只螺和415只蟹中, 11.8 %的螺感染了8种马蹄属线虫中的一种,且以滨鹬马蹄吸虫的感染占优势(9.9 %螺感染) ; 80 %的蟹感染滨鹬马蹄吸虫囊蚴。虽然一年四季两种宿主都会感染,但吸虫的流行和尾蚴(指成熟期感染)在夏季呈现高蜂。从螺体排出的尾蚴具有明显季节性,在此海湾必须要超过最低温度20℃。总的感染率在较大(较老)的螺里有所下降,显示吸虫影响宿主生存并随之影响宿主群体结构。囊蚴的感染丰度与蟹的个体大小有明显相关性;较大的蟹感染较多的囊蚴,显示宿主能耐受更多的吸虫。调查显示,囊蚴的感染率与蟹的大小或性别无相关性。囊蚴体外脱囊以及产卵吸虫的释放证明,成熟虫体终年存在于所有大小和性别不同的蟹里,显示从蟹到鸟的持续感染是可能的。总的来说,滨鹬马蹄吸虫在海湾的传播动态是由这两种无脊椎动物宿主来协调,并似乎是被一系列依赖于温度的活动控制,这些活动影响易感宿主种群及感染性幼虫期尾蚴和囊蚴的存在。     

Comparative analysis of growth of edible mussel Mytilus edulis from different White Sea regions

Growth properties were studied in edible mussel Mytilus edulis from different biotopes of the Kandalaksha Bay in the White Sea.Growth curves of the mussel shell were approximated by the von Bertalanffy equation. The highest growth rate, primarily dependent on hydrological conditions, was observed in mussels from the Chupa Bay (Levin Navolok fishery), while the lowest rate was observed in mussels from the Umba region (Turii Cape, Murmansk Region). Allometric relationships of the mussel shell were determined. Analysis of the relationship between the maximum width/convexity and their length demonstrated that this relationship is described by a single allometric equation for mussels from the Umba region, which indicates the homogeneity of this population. For mussels from the Chupa Bay, this relationship cannot be described by a single equation, which points to the population heterogeneity. For 90% mussels from this region, the shell width/convexity ratio ranged from 0.5 to 0.9; while for the remaining 10%, it ranged from 0.9 to 1.15.     

Fouling community of the red king crab, <Emphasis Type="Italic">Paralithodes camtschaticus</Emphasis> (Tilesius 1815), in a subarctic fjord of the Barents sea

We examined the species composition of red king crab (Paralithodes camtschaticus) fouling communities in Dolgaya Bay, a small fjord of the Barents Sea, in August 2005 and 2006. In total, there were 13 species observed on 301 crabs collected from water depths of 5–90 m. Barnacles (Balanus crenatus; prevalence 42.9%) and blue mussels (Mytilus edulis; 11.6%) were the most common epibionts, while amphipods (Ischyrocerus commensalis) were the most common symbionts (28.6%). Infestation rates in Dolgaya Bay were different from those in an “open” area of the Barents Sea (Dalnezelenetskaya Bay), probably due to differences in hydrodynamic conditions. Differences in infestation prevalence and intensity were detected neither between male and female crabs nor between crabs collected at 5–35 m versus 90 m depths. Prevalence of common fouling species increased with host size. Amphipods I. commensalis colonized the carapace and limbs in Dolgaya Bay less frequently than in Dalnezelenetskaya Bay, probably due to interspecific competition with barnacles occupying the dorsal parts of the host. Juvenile barnacles and mussels dominated the fouling communities on the crabs. The age of barnacles did not exceed 2–4 months. However, the presence of 4-year-old mussels suggests that these older mollusks have been directly transferred from mussel beds to the hosts. Our results indicate that colonization by epibionts and symbionts is generally not disadvantageous for the crab hosts, except for some possible negative impacts of amphipods occupying the gills.     

Vegetative propagation and spore-based recruitment in the carrageenophyte Chondracanthus chamissoi (Gigartinales, Rhodophyta) in northern Chile

In the present study we compared vegetative and spore‐based propagation for Chondracanthus chamissoi (C. Agardh) Kützing. Monthly field observations were made over a 1‐year period at Puerto Aldea, Tongoy Bay, Chile. Data were collected both outside and within a bed of C. chamissoi. Vegetative propagation was assessed via attachment of drifting fronds to shell‐encrusted concrete blocks at both sites. Germination of spores was recorded on the same shell substrates. Substrate re‐adhesion varied seasonally between sectors. Highest averages occurred within the algal bed between January 1997 and March 1997. The number of sporelings showed two peaks of maximal recruitment in spring and summer months (January‐March 1997 and September 1997‐January 1998). Spore‐based propagation is an important mechanism of seasonal regeneration of biomass in the C. chamissoi bed; however, re‐attachment of fronds may have been important in maintaining production of the bed during the period of maximum biomass accumulation.     

First report of the aberrant association of branchiobdellidans (Annelida: Clitellata) on blue crabs (Crustacea: Decapoda) in Chesapeake Bay, Maryland, USA

Abstract. This is the first article of branchiobdellidans on blue crabs, Callinectes sapidus , in Chesapeake Bay, MD, USA. The ectosymbionts consisted of sympatric populations of Cambarincola mesochoreus and Cambarincola pamelae living on the ventral body surface and in the gill chambers of blue crabs. The occurrence of branchiobdellidans in the upper bay region was documented at ten sites during August and late September 2003. Branchiobdellidans were found on blue crabs following a rainy summer when salinity levels fell below 3 ppt. The blue crabs move into the freshwater estuaries during the summer, where they enter the habitats of crayfishes; however, the mechanism of transfer from one crustacean host to another is not known.     

THE RELATIONSHIP BETWEEN SHELL INJURIES, SHELL THICKNESS AND HABITAT CHARACTERISTICS OF THE INTERTIDAL SNAIL LITTORINA RUDIS MATON

The hypothesis that risk of shell injury from crab predationand crushing by boulders and stones is important in the selectionof shell thickness in the intertidal gastropod Littorina rudisMaton was tested using data from 24 populations from Britain.Although there was an obvious relationship between the incidenceof shell injuries and both the distribution of crabs and substratummobility, there was no correlation between risk of shell injuryand shell thickness. This is probably accounted for by between-habitatvariations in energy levels which may vary independently ofcrab density and substratum mobility. ^*Present address: Department of Biology, University of Essex,Wivenhoe Park, Colchester C04 3SQ (Received 23 May 1977;