Predation of cockles (Cerastoderma edule) by the whelk (Buccinum undatum) under laboratory conditions

The feeding rate and behaviour of whelks (Buccinum undatum)offered cockles (Cerastoderma edule) in laboratory experimentswere examined. When presented with cockles in a range of sizes(10–40 mm), 14 B. undatum (34.6–88.3 mm),held individually in aquaria, consumed a wide size range ofcockles. Small whelks (<40 mm) consumed cockles (<23 mm),whereas large whelks, (>60 mm) ate a greater numberof larger cockles (>30 mm) and a wider size range ofcockles (12–40 mm) than smaller whelks. The majority(90%) of the shells of the predated cockles were undamaged andthe few (<10%) that were damaged showed only slight abrasionsto the anterior and posterior shell margin. Filmed observationsof B. undatum feeding on C. edule showed a method of attackthat has not previously been reported and involved the use ofthe whelk's foot to asphyxiate the cockle or to pull the shellvalves apart. No filmed evidence was found for the previouslyreported shell ‘wedging’ technique for prising openthe closed shell valves of C. edule, although 10% of the shellsof consumed cockles in feeding experiments had damaged shellmargins. (Received 4 April 2007; accepted 30 June 2007)     

VARIATIONS IN THE PHYSIOLOGICAL STATE OF THE COMMON COCKLE (CERASTODERMA EDULE (L.)) IN THE LABORATORY AND IN SOUTHAMPTON WATER

Various physiological indices (i.e. body condition index, scopefor growth and net growth efficiency) have been used to examinethe metabolic state of the common cockle (Cerastoderma eduleL.) at different sites in Southampton Water, U.K. The resultsshow that there is a measurable difference in the conditionindex of cockles collected from different localities. Thesedifferences can be broadly related to the level of contaminationof the site with metals and hydrocarbons. All the indices suggestthat cockles inhabiting the eastern, less polluted side of SouthamptonWater are in a better metabolic state than in those from somesites on the western side close to industrial areas. Growthand longevity also appear to be site related. (Received 25 September 1989; accepted 25 March 1990)     

Calorimetric Studies of Behavior, Metabolism and Energetics of Sessile Intertidal Animals

SYNOPSIS: Behaviors to conserve water during intertidal exposureat the same time impair respiratory gas exchange, so that observedresponses to emersion may reflect compromises between theseincompatible needs. Behavioral isolation of the tissues fromair results in the complete or partial reliance on anoxic energymetabolism, which is most reliably measured directly as heatdissipation. Combined direct calorimetry and indirect calorimetry(respirometry) enable the partitioning of total metabolic heatdissipation into its aerobic and anoxic components, which mayvary according to physical and biological factors. The musselMytilus edulis is tolerant of anoxia and saves water and energyduring aerial exposure in its rocky intertidal habitat by closingits shell valves and becoming largely anoxic. Like most suspensionfeeders in this habitat, its compensation for reduced feedingtime involves energy conservation; there is little evidencefor energy supplementation such as increases in feeding rateor absorption efficiency. Ammonia production continues duringaerial exposure and is involved in acid-base balance in thehemolymph and mantle cavity fluid. Infaunal cockles (Cardiumedule) and mussels (Geukensia demissa) gape their shell valves,remain largely aerobic and have high rates of heat dissipationduring intertidal exposure, a response which appears relatedto the lower desiccation potential and exploitation of richertrophic resources in their soft-sediment habitats. The variableexpansion of the symbiotic sea anemone Anthopleura elegantissimareflects interaction among the responses to desiccation, irradianceand continued photosynthesis by its zooxanthellae during exposureto air.     

SHELL GROWTH OF THE COCKLE CERASTODERMA EDULE IN THE MUNDACA ESTUARY (NORTH SPAIN)

In order to establish a growth curve for the life-span, agedetermination based on shell-ring analysis was undertaken ina population of the cockle Cerastoderma edule located in theMundaca Estuary (Basque Country, North Spain). Mean values ofheight at shell rings showed non-significant differences betweengenerations, meaning absence of interannual variations in growthrate. Frequent sampling and shell measurements over a 20 monthperiod allowed determination of the seasonal pattern of shellgrowth, which was subsequently incorporated into a growth curve. An attempt has been made to relate growth rates to latitudefor different populations of C. edule, using both data fromthe literature and the results of this study. The highly significantcorrelation found (P<0.01) confims the existence of a latitudinaltrend, with growth rates increasing southwards. Low rates ofgrowth recorded for cockles from Mundaca, by camparison withpupulations of similar latitude, are interpreted in terms ofnutritional restrictions associated with both the high tidalposition of the population and poor productivity conditionswithin this particular estuary. (Received 24 March 1989; accepted 20 June 1989)     

Über Schalenabschliffe beiCardium edule aus der Königsbucht bei List auf Sylt

Two communities of the recent bivalveCardium edule L. were collected in the tidal flats near List on the island of Sylt (Southern North Sea). The cockles were examined for abrasion marks on the surfaces of their valves arising from various causes. One important group of abrasions reveals traction marks restricted to the beak region, caused exclusively by valve movements. Another group is characterized by marks due to shell abrasions on the sea bed. The various types of abrasion marks have been studied in living cockles, both in the sea and in laboratory containers, and the results compared with those obtained on dead ones still connected by the ligament. In the sea, livingCardium edule were studied via Scuba-observations. Traction marks on the umbo support the conclusion that living cockles sometimes rest on their dorsal shell area. Holes in the convex umbo, umbonal facets, originate after death in situ or through shiftings to and fro on the sea bed, and also as a result of chemical or biological processes. The method applied and the criteria of shell abrasions employed are considered acceptable approaches to the interpretation of functional morphology and behaviour in fossil molluscs.     

Stereotypic and size-selective predation in Polinices pulchellus (Gastropoda: Naticidae) Risso 1826

Polinices pulchellus were size-selective in their choice of Cerastoderma edule. Large predators (12-15.9 mm shell length) selected both larger and a wider size range of cockles than smaller individuals (4-11.9 mm shell length). Considerable overlap occurred in the sizes of cockles frequently drilled by different size classes of snails, indicating that certain sizes of cockles may be most profitable to a wide range of predator sizes. Consumption rates were highest during July and August and were closely related to seawater temperature. Inner and outer drill hole diameters were both correlated with predator size, and the morphology of the drill hole was geometrically similar across a range of predator sizes. Polinices pulchellus showed no preference for either the left or right valve and drilled most cockles in the centre of the shell valve. The relationship between the distance of the drill hole from the umbo and prey size was unaffected by predator size, such that predators of different sizes were not found to drill cockles in different positions. When disturbed during drilling, incomplete drill holes were abandoned and, when drilling resumed, it occurred in new locations on the surface of the shell valve. The findings of this study highlight the stereotyped nature of drilling behaviour seen in the family Naticidae.     

The use of Cerastoderma glaucum as a sentinel and bioindicator species: Take-home message

Bivalves are frequently used to assess environmental contamination, and are often considered good sentinel and/or bioindicator species. For that reason the bioaccumulation and toxicity induced by metals and As in the cockle Cerastoderma glaucum, collected from areas with different contamination levels along the Óbidos lagoon (Portugal), were used to evaluate the use of this species as sentinel and/or bioindicator. The results showed that areas in the middle of the lagoon presented lower metals and As concentrations, lower total organic matter content and lower percentage of fine particles than areas in the Bom Sucesso arm. In all areas Cr, Pb and Cu were the most abundant elements, while Ni, As, Cd and Hg were less abundant. Results also showed a moderate correlation between total elements concentrations found in C. glaucum and in sediment, and thus caution should be taken when considering this species as a good sentinel species. The present study also revealed that, in general, C. glaucum from areas in the middle of the lagoon accumulated higher concentrations of metals and As (Biota-Sediment Accumulation Factor >1) than cockles from the most polluted areas located in the Bom Sucesso arm. However, in all areas, the majority of metals (Cu, Cr, Pb) were found in cockles insoluble fraction which may explain low cellular damage and reduced oxidative stress responses observed. Therefore, our results may further alert for caution when identifying C. glaucum as a good bioindicator species. Thus, our findings highlight the fact that studies should be cautious when selecting species for environmental monitoring, since good sentinels or bioindicators in highly polluted systems may not act in the same way in low or moderately contaminated areas. Furthermore, our study warns for the misclassification of cockles in different ecosystems.     

Parasite Induced Summer Mortality in the Cockle Cerastoderma edule by the Trematode Gymnophallus choledochus

In late summer 2004, a conspicuous cockle (Cerastoderma edule) mortality event was observed on a tidal flat in the northern Wadden Sea (North Sea, Germany) with many fresh valves and still living cockles lying on the sediment surface. To investigate whether trematode parasites utilizing the cockle as first or second intermediate host were involved in this mortality, buried and surfaced cockles were sampled and analyzed, and a laboratory experiment conducted. The field survey showed no statistical difference in intensity of parasites encysted in the foot of cockles. Three species of Himasthla utilizing the cockle as second intermediate host and known to impair the cockle’s burrowing ability were found in buried cockles with 148.4±111.1 metacercariae/foot and in surfaced cockles with 164.2±84.4. There was also no difference in infection levels of parasites utilizing the cockles as second intermediate host in other cockle tissues between buried and surfaced cockles. In contrast, surfaced cockles showed a ten times higher prevalence (71.0%) than buried cockles (7.4%) of the trematode Gymnophallus choledochus – a parasite utilizing the cockle as first (and second) intermediate host – filling almost the entire body cavity and eliminating gonad structures. In an aquarium experiment of 14 days, all cockles found buried on the tidal flat survived compared to only 23.3% found on the surface. This suggests G. choledochus to be a castrating agent and a serious mortality factor in adult cockle populations.     

REPRODUCTIVE CYCLES OF CERASTODERMA GLAUCUM(BRUGUIERE) AND C. EDULE (L.) WITH SPECIAL REFERENCE TO THE EFFECTS OF THE 1981-82 SEVERE WINTER

Gonadal changes in two species of Cerastoderma occurring indiscrete populations in South Wales were monitored by stereologicalanalysis during the breeding seasons of 1981 and 1982. Fecundityand spawning efficiencies were determined quantitatively. Therelationship between reproductive activity and changes in meatcondition are discussed. Both species are opportunistic breeders. Their normal patternsof reproduction are similar, consisting of minimum activityin winter and peak activity during spring/early summer. Thetwo species were affected differently by the severe winter of1981–82—C. glau-cum exhibited a single, ‘epidemic’spawning and resumed gametogenesis after a 4-month resting period.C. edule adopted a ‘polycyclic’ pattern withouta resting period. Both of these unusual reproductive strategies,augmented by reduced predation resulted in heavy spatfalls. (Received 20 March 1985; revised 20 March 1985;     

Karyotypes of Cerastoderma glaucum (Bivalvia) from Baltic and Mediterranean populations

Chromosomes of C. glaucum from Baltic (Gdansk Bay) and Mediterranean (Thau lagoon, Sète) populations were studied using karyometric analysis and silver-staining. The karyotype of the Gdansk population consists of three metacentric, ten submetacentric and six subtelocentric chromosome pairs. The karyotype of the Thau population shows four metacentric, nine submetacentric and six subtelocentric pairs. Nucleolus organizer regions were found terminally on the second largest submetacentric chromosome pair of the Gdansk population and on the second largest metacentric chromosome pair of the Thau population. This suggests a cytotaxomomic difference between these two geographically isolated populations. Striking differences exist between C. glaucum and C. edule karyotypes, probably resulting from a long divergence time.     

Metapopulation structure in the lagoon cockle <Emphasis Type="Italic">Cerastoderma lamarcki</Emphasis> in the northern Wadden Sea

Benthic invertebrates in discontinuous inshore habitats and with short or no pelagic larval dispersal are likely to exhibit regional metapopulation dynamics with partially isolated local populations. Near the island of Sylt, the bivalve Cerastoderma (Cardium) lamarcki (Reeve, 1844, syn. in part with C. glaucum Bruguiére, 1789) was widespread in intertidal seagrass beds, coexisting with the sibling species C. edule (Linné, 1758). However, the last C. lamarcki in this habitat was found in 1980. At present the lagoon cockle is restricted to disjunct ditches, creeks and ponds within island salt marshes. There it differs in year-class structure between localities. Successful recruitment events did not coincide. At one locality, a period with regular recruitment was followed by 5 years of recruitment failure, resulting in an overaged population probably at the rim of extinction. In a nearby brackish pond, extinction was followed by recolonization 3 years later. Other lagoonal habitats which seem to be suitable are without cockles. It is speculated that small and isolated habitats occasionally receive colonizers by eggs and juveniles adhering to avian vectors. Electronic Publication     

Detection of Infectious Cryptosporidium parvum Oocysts in Mussels (Mytilus galloprovincialis) and Cockles (Cerastoderma edule)

Infective Cryptosporidium parvum oocysts were detected in mussels (Mytilus galloprovincialis) and cockles (Cerastoderma edule) from a shellfish-producing region (Gallaecia, northwest Spain, bounded by the Atlantic Ocean) that accounts for the majority of European shellfish production. Shellfish were collected from bay sites with different degrees of organic pollution. Shellfish harboring C. parvum oocysts were recovered only from areas located near the mouths of rivers with a high density of grazing ruminants on their banks. An approximation of the parasite load of shellfish collected in positive sites indicated that each shellfish transported more than 10³ oocysts. Recovered oocysts were infectious for neonatal mice, and PCR-restriction fragment length polymorphism analysis demonstrated a profile similar to that described for genotype C or 2 of the parasite. These results demonstrate that mussels and cockles could act as a reservoir of C. parvum infection for humans. Moreover, estuarine shellfish could be used as an indicator of river water contamination.     

The effect of size and temperature on the predation of cockles Cerastoderma edule (L.) by the shore crab Carcinus maenas (L.)

Laboratory studies demonstrated that shore crabs Carcinus maenas (L.) can consume <40 cockles Cerastoderma (= Cardium) edule (L.) per individual · day⁻¹. Various predation techniques used by the crabs are reported. The time required to open and consume individual cockles increased exponentially with prey size. Small (<l5-mm shell height), easily broken cockles appeared to be the most profitable in terms of energy acquisition per unit of handling time, the optimal size of prey increasing with predator size. With unlimited prey available, however, crabs selected prey of mean size smaller than these predicted optima, and much below the maximum size they were capable of opening. Feeding rates, both in terms of cockles ingested or energy intake per day, rose steeply with increasing temperature, but the size range of prey consumed remained unchanged. These data strongly suggest that Carcinus maenas is a potentially important predator of small cockles, particularly during the wanner summer months.     

SOME BEHAVIOURAL AND ELECTRICAL RESPONSES OF SCROBICULARIA PLANA (BIVALVIA:TELLINACEA) TO SIPHONAL WOUNDING

Water pumping, valve movements and heart rate have been recordedfrom Scrobicularia for short periods of normal behaviour andthen after siphonal wounding. Scrobicularia exhibits regularand repetitive pumping periods interrupted for only 2–3s after siphonal wounding, without the regularity of these periodsbeing affected. Wounding does not prevent animals from usingtheir inhalant siphons for deposit feeding. A preliminary investigationof neural responses to stimulation has shown that wounding thesiphon causes minimal disturbance to the animal, a brief (2s)burst of nerve activity occurs, the siphon is retracted, butvalve adduction does not occur. In contrast to this tactilestimulation of the mantle edge always elicits a large burstof impulses in the posterior adductor nerve, valve closure results,usually for 14–15 s. ¹Present address: Dept of Zoology, University of Cape Town,Rondebosch 7700, South Africa. (Received 2 February 1981;     

QUANTIFICATION OF THE PHYSIOLOGICAL RESPONSES OF THE EUROPEAN FLAT OYSTER OSTREA EDULIS L. TO TEMPERATURE AND SALINITY

Physiological stress was quantified in specimens of the Europeanflat oyster Ostrea edulis, acclimated to a matrix of temperature(5–25°C) and salinity (16–34%o) conditions,using the Scope for Growth (SFG) index. Variations in the indexwere interpreted from the responses of its components, absorptionefficiency, filtration rate, respiration and ammonia excretionrates, to external conditions. Mean SFG values were most influencedby acclimation temperature but significant differences werefound between animals at different salinities over the rangetested. There was a severe reduction in SFG when high temperaturewas combined with reduced salinity (     

Kinetics of Maize Leaf Elongation: I. INCREASED YIELD THRESHOLD LIMITS SHORT-TERM, STEADY-STATE ELONGATION RATES AFTER EXPOSURE TO SALINITY

The short-term responses of leaf elongation to salinity areinvestigated in this study. The kinetics of maize (Zea maysL.) leaf elongation were measured with Linear Variable DifferentialTransformers (LVDTs). After exposure to salinity (0 to 120 molm^–3 NaCl), leaf elongation rates (LER) declined rapidly.Within 4 h, LER had recovered and reached a new steady-statefor all salinity treatments. These rates were reduced by 10,20, and 60% of control rates by 40, 80 and 120 mol m^–3NaCl, respectively. Osmotic adjustment in the growing zone ofleaves was correlated with the recovery of LER after plant exposureto salinity. However, after 4 h of exposure, the osmolalityof the cell sap continued to increase without effect on steady-stateLER. Estimates of the apparent turgor in the growing zone indicatedthat turgor was no longer limiting LER of salt-stressed plantsafter 4 h. An in vivo technique was developed to apply a unidirectionalforce to intact growing leaves of maize to mimic increases inelongation force. Relative elongation rate (RER) were increasedby adding weights to the LVDT core to increase elongation force.Plots of RER as a function of elongation force gave estimatesof two growth coefficients: the yield threshold and the yieldingcoefficient, mL/(m + L), where m is the cell wall extensibilityand L is the hydraulic conductivity. RER as a function of elongationforce was determined immediately, 05, 4, and 21 h after plantswere salinized. Estimates of the growth coefficients indicatedthat the apparent yield threshold decreased immediately aftersalinization. However, when LER reached steady-state, the yieldthreshold of salt-stressed plants had increased above controlvalues and was the only limiting growth coefficient. There wereno significant effects of salinity on the yielding coefficients,cell wall extensibility or hydraulic conductivity. One of theadvantages of this in vivo technique over other methods is thatyield threshold, yielding coefficient, and cell wall extensibilitycan be determined without the confounding effects of woundingor osmotic stress. This technique may prove widely applicableto the study of other growth regulating factors. Key words: Salinity, leaf growth, Zea mays L     

MECHANICAL AND MORPHOLOGICAL PROPERTIES OF THE SHELL OF SCROBICULARIA PLANA (DA COSTA) UNDER NORMAL AND STRESS CONDITIONS

Scrobicularia plana reacts to salinity stress by closing itsvalves and respiring anaerobically. The acidic products of respirationare buffered by calcium carbonate removed from the shell. Shellmass decreases by 15% over a period of three weeks, and thereis a similar reduction in shell strength. Erosion of the shell surface, indicating buffering, takes placequickly, pits can be seen in the valves of shells at the endof six hours of normal intertidal emersion. (Received 10 February 1982;     

The glycogen content in stressed marine bivalves: The initial absence of a decrease

• 1.1. Changes in the glycogen content, condition, stomach content and acetic acid concentration of mussels Mytilus edulis and cockles Cerastoderma edule were followed during periods of up to 14 days of exposure (to air) at temperatures of 5 and 20°C.
• 2.2. In animals with a high glycogen content the glycogen is not used during the first 3 to 7 days, at high and low temperature respectively.
• 3.3. After this latent period the glycogen concentration often decreased, coinciding with a high mortality and an increase of the concentration of acetic acid.
• 4.4. In cockles with a low glycogen content, and kept at a high temperature, glycogen can be used from the beginning of the stress period.
• 5.5. Between species no clear differences were found.
• 6.6. The stomach content decreased during exposure; however, the stomach content amounted to only 0.5 to 0.7% of the body weight, and is thought to be of minor importance as an energy source during the stress period.
• 7.7. Especially at the higher temperatures glycogen finally is transformed into acetic acid.
• 8.8. It is concluded that during exposure, the animals do not die because of a lack of energy reserves, but because of a high accumulation of acids.

     

Phylogeography of the temperate marine bivalve Cerastoderma edule (Linnaeus, 1758) (Bivalvia: Cardiidae) in the Subarctic: Unique diversity and strong population structuring at different spatial scales

Using mitochondrial COI sequencing, we explored the genetic diversity and population structuring of the common cockle Cerastoderma edule (Linnaeus, 1758) in the Norwegian and Barents Seas. Phylogeographic diversity and hence the evolutionary history of C. edule on the Scandinavian and Russian coastlines were found to be richer than expected for populations of temperate species in postglacially colonized seas. A major phylogeographic break at Lofoten Islands separated a group of subarctic populations dominated by a distinct star‐shaped clade of haplotypes from those to the south, extending to the North Sea and having highest gene diversities (h). At the northeastern edge of the range of C. edule, the Russian Murman coast, populations show a mosaic structure with considerable admixture of haplotypes from the south and high local‐scale variation in haplotype diversity (ranging between 0 and 0.8). To explain this mosaic we refer to the core‐satellite metapopulation model, with Norwegian populations as core, and Murman populations as satellites. Our results contradict the conventional biogeographic paradigm implying lack of metapopulation structuring in marine broadcast spawning invertebrates. Hypotheses considered to explain the origin of the unique variation in cockles from Northern Norway involve an early postglacial colonization and establishment of these populations (10–12 ka ago), a persistent oceanographic break at Lofoten, and a mitochondrial selective sweep associated with the postglacial recolonization of the subarctic seas by the boreal C. edule.     

A QUANTITATIVE STUDY OF A CERASTODERMA -- NEPHTHYS COMMUNITY IN THE LIMFJORD, DENMARK, WITH SPECIAL REFERENCE TO PRODUCTION OF CERASTODERMA EDULE

Cerastoderma edule is the dominant member of a non-random Cerastoderma—Nephthyscommunity, occupying a physically, homogeneous, submerged sandbankin the Limfjord, Denmark. C. edule is patchily distributed withinthis community, growth rates showing a clear density-dependence.Individual production is about 35%lower in high density patchesthan outside them; at low density annual individual flesh dryweight increment is 295 mg. Seasonal variation in water, ash,lipid, protein and glycogen contents is studied, and a sizedependent(= density-dependent) relationship is shown to exist for someparameters. Population production, estimated by a new graphicalmethod, and taking into account gametic and shell protein production,attains 150 g dry flesh weight, m^–2.yr^–1 at meanpopulation density rising to 250 g dry flesh weight. m^–2.yr^–1in the high density patches. ¹Present address: Institut für Physiologische Chemie, Abt.Zellbiochemie, Universitat Hamburg, Martinistrasse 52, 2000Hamburg 20 (Received 10 March 1980;