BIOLOGY AND POPULATION DYNAMICS OF A CREVICE-DWELLING LANDSNAIL, CRISTATARIA GENEZARETHANA (CLAUSILIIDAE)

Crevice-dwelling land snails emerge only infrequently to theexposed surface, and very little is known about their naturalhistory. Here we report on the biology, life cycle, populationdynamics and longevity of one such species, Cristataria genezarethana,which we studied over a period of three years. C. genezarethana spends 95–98% of its lifetime withinrock crevices. In winter the crevice serves as an egg layingsite, and as a retreat during periods of inactivity. Throughoutsummer it serves for aestivation adults aestivate near CTeviceopenings, young apparently deeper inside. The surface of the rock serves mainly as a lichen-feeding andcopulating site, to which the snails emerge during brief periodsof activity. Both for the onset and for the continuation ofits activity season, C. genezarethana is absolutely dependentupon rain. However, activity ends before the end of the rainyseason, suggesting that aestivation is not merely a direct responseto dry climate. Though the population as a whole was active on each rainy day,no more than 15%–20% of the population were active simultaneously.An individual snail was active, on average, for only 6–12days per year. Population size of the study-rock reached 2000–2900individuals, and mean density was 150–200 snails m². Thissnail carrying capacity of the rock is broadly similar to thatof densely populated bushy habitats (when snail weight is considered).In these dense populations, low mating frequencies and growthinhibition may regulate population size. In all three seasons, the population consisted of two main large,well defined age groups: adults and young. As compared to theadults, the young were active on different parts of the rockand at different times. Although a third, intermediate groupof sub adults was found, it was always very small. A fourthgroup, of juveniles, appeared during winter; later, most ofthem disappeared. We did not observe any shift from the young to the adult sizegroup. This suggests a broadly stable population, with virtuallyno recruitment, in which final growth to adulthood is inhibitedby the adults. Growth was very slow, suggesting that maturityin nature is reached in about 11 years, and that individualslive sixteen years at least. Mortality occurred in all age groups,and about 5% of the population died each year. There was noevidence for heavy predation. Crevices might be a suitable habitat for land snails in whichthe hatching period, and period of juvenile growth, are verylong. (Received 1 February 1993; accepted 27 March 1993)     

The effects of low temperature on Lymnaea truncatula.

A group of 20 young and another of 20 adult Lymnaea truncatula were abundantly supplied with food and kept continuously under cold conditions (5 degrees C) in the laboratory for 3 months and the effects of low temperature on their behavior, growth and reproduction were studied. The results indicate that at low temperature the activity of L. truncatula was markedly reduced but complete hibernation did not occur. The snails seem to be unaffected by the low temperature itself since none (both young and adult) died during the 3 months that they were kept at 5 degrees C. Reduced feeding, even in the presence of abundant food, during the cold conditions caused an almost total inhibiton of growth. Of even more significance was the suppression of reproduction which was connected with the metabolic rate of adult snails kept at low temperature. Young snails seem to profit by exposure to low temperature. On the return to normal laboratory temperature (16-22 degrees C) the young snails became very active, fed voraciously, grew rapidly, tended to live longer and produced more offspring than the controls. Low temperature, however, appears to have an adverse after-effect on the growth and reproduction of mature snails. Relatively, fewer eggs were deposited in this case. The results indicate that under natural field conditions in England, where temperature fluctuations during the usually mild winter months are common, the greater burden of increasing the population in overwintered snails must rest on the younger members of the community.     

DIFFERENCES IN THE LIFE HISTORIES OF XEROLENTA OBVIA (MENKE, 1828) (HYGROMIIDAE) IN A COASTAL AND A MOUNTAINOUS AREA OF NORTHERN GREECE

The life cycle of Xerolenta obvia (Menke, 1828) was studiedin two areas, Paleokastro (Chalkidiki), an inland mountainousarea, and Nea Karvali (Kavala), a coastal area in northern Greece.At Paleokastro snails hatch in autumn, become adult the followingJuly, but do not lay eggs until October, after which they die.Clutch sizes are small, but eggs and hatchlings are large comparedwith those at Nea Karvali. Growth is fast in spring, and continuesuntil the end of July. The young hatchlings have dark shell-bands,but by July a quarter of snails appear unbanded. At Nea Karvali,eggs are also laid in October, and young snails emerge fromhibernation in March. Here, however, they do not mature untilApril of the following year. They thus have a 2-year life cycle,with adults dying in their second autumn. Clutches are aboutthree times the size of those at Paleokastro, but eggs and hatchlingsare significantly smaller. A little growth occurs in winter,but the rate of growth is generally much slower than at Paleokastro.Only 1–2% of this population has banded shells; the bandsare less obvious and they become invisible in some individualsas they mature. At both sites population density fluctuatedduring the two study years, but it was always higher at NeaKarvali. These results are discussed in relation to the differingclimates of the sites, and comparisons made with studies onrelated species in the region. (Received 23 July 2004; accepted 30 December 2004)     

The role of growth,predation, and habitat selection in the population distribution of the crown conch,Melongena corona Gmelin

The intertidal gastropod Melongena corona Gmelin exhibits a size gradient along the shore as a function of habitat. Small, juvenile snails were found on the sand beach and larger adults were found on hard substratum habitats (shell rubble beach and oyster bar). A transplant experiment was performed to test three proximate explanations for this pattern:

• 1.(1) differential growth rate of conchs in different habitats,
• 2.(2) differences in predation intensity between habitats and
• 3.(3) active habitat selection by snails.

Shell scars on the last shell whorl were used as an index of predation pressure. Growth and predation were not significantly different for snails of similar size in different habitats, but snails were found to return to their original habitat when displaced. The ability to home did not differ between sexes or juveniles and adults. Active habitat selection appears to be a significant proximate factor maintaining the population distribution. A number of potential ultimate causes of the size class segregation are suggested.     

Predator-limited population growth of the copepod Pseudocalanus sp.

The impact of predators on population growth of Pseudocalanussp. was investigated in Dabob Bay, Washington. Mortality ofPseudocalanus sp. was determined from stage-specific survivorship,from seasonal changes in mortality rates of adult males andfemales and from incidence of injuries to adult copepods. Theprincipal predators of adult Pseudocalanus were identified asthe predatory copepod Euchaeta elongata, the omnivorous euphausiidEuphausia pacifica and the chaetognath Sagirta elegans. Predatorattack rates - and prey mortality rates - are highly density-dependentand thus sensitive to prey dispersion in the water column, particularlyto layering in the vertical plane. Predation rates by the threeprincipal predators exceeded 100% of the recruitment rate toadult Pseudocalanus sp. beginning in early summer, thus restrictingpopulation growth. Planktivorous fish predation (by adult three-spinestickleback, Gasterosteus aculeatus, and juvenile chum salmon,Oncorhynchus keta) on Pseudocalanus sp. adults was estimatedto be two orders of magnitude lower than consumption rates bypredatory zooplankton, at a deep water station in July. Analysisof seasonal changes in prey ingested by Sagitta elegans revealedthat Pseudocalanus sp. was the major prey item of S. elegansin April (61.0% of prey) and in June (67.0% of prey), thereafterdeclining seasonally in importance. Predation by S. elegansvaried seasonally with changes in chaetognath stage structure,vertical distribution and diapause, not size structure alone.Although chaetognath recruitment and population growth appearto be directly coupled to the abundance of Pseudocalanus sp.,predation by S. elegans has little reciprocal impact on Pseudocalanussp. population growth; hence asymmetries may occur in the interaction of planktonic prey and predators.     

POPULATION DENSITY EFFECTS ON GROWTH IN CULTURE OF THE EDIBLE SNAIL HELIX ASPERSA VAR. MAXIMA

The effects of population density on the growth of H. aspersaMÜller var. maxima under controlled environmental conditionswere examined. Inhibitory effects on snail growth and maturityresulting from increased population density, between 100–800snails m^–2 of floor area, were observed for a range ofcontainer cleaning frequencies. At all population densities,enhanced snail growth was observed when the frequency of containercleaning was increased to a two-day interval. No significantdifferences were recorded, following 19 weeks growth, betweenfinal mean weights of snails from containers cleaned less frequently.The lowest snail mortality was consistently recorded at thelowest population density in the most frequently cleaned containers.At all snail population densities three phases of growth wereobserved: (a) lag (0–5 weeks), (b) rapid (6–15 weeks)and (c) stable (16 weeks and over). During the first three weeksof growth, high population density had a positive effect ondiet consumption, food conversion efficiency and snail growth.Adverse population density effects increased progressively duringphase (b), typically following 9 weeks growth. Juvenile snailstransferred from high to low population densities during phase(b) continued to exhibit slower growth rates associated withhigh population densities. Food conversion efficiency of snailsin all treatments decreased throughout the experimental periodbut with no overall effect of container cleaning frequency apparent.Inherent growth variability of sibling snails was unaffectedby population density or container cleaning frequency. The importanceof the results for intensive snail culture is discussed. (Received 23 June 1994; accepted 1 December 1994)     

Toxicite a long terme d'un insecticide organophosphore (fenthion) chez le mollusque Lymnaea stagnalis L.

A chronic intoxication with fenthion was induced in adult and young snails. The fecundity was only reduced by the concentratration of 4 ppm in adults. In the snails intoxicated by fenthion from hatching the shell growth and the fecundity were both reduced from a concentration of 2 ppm. Above 2.15 ppm the rearing of young snails was not possible.

     

The effects of changes in food availability on the breeding ecology of great skuas Catharacta skua in Shetland

Great skuas on Foula, Shetland have responded to a decline in the availability of sandeels since the late 1970s by increasing the proportion of other items in their diets. This change is correlated with the annual recruitment of sandeels in Shetland waters. Since 1983 there has been a 10-fold increase in predation by great skuas upon other seabirds, as Furness & Hislop (1981) suggested might occur in response to a low availability of sandeels. Changes in diet have been accompanied by a 50% reduction in adult territorial attendance as adults increased their foraging effort, such that between 1987 and 1989 breeding adults were probably working as hard as they were able to. Despite this, breeding success was less than 40% in 1987 and less than 15% in 1988 and 1989. The major cause of breeding failure was predation of unguarded chicks by adults from neighbouring territories. The willingness of adults to expose their chicks to high predation risk is probably maintained because of a positive correlation between chick pre-fledging growth and post-fledging survival, which is expressed up to the age of two years and which will place a strong pressure upon adults to feed their chicks as well as possible. The high expenditure of effort by adults in 1987 and 1988 did not affect the weights of those birds incubating eggs in 1988 and 1989, but there was a slight (3%) decrease in egg size between the late 1970s and the late 1980s. Changes in the age structure of the breeding population and the absence in 1989 of 28% of adults colour-ringed during incubation in 1988 suggest an increase in the rate of egress since the 1970s. These changes probably represent an increase in the long-term costs of reproduction to adults at this colony.     

Distribution in relation to life history in the directdeveloping gastropod Batillaria cumingi (Batillariidae) on two shores of contrasting substrata

Population structure and spatial distribution with growth ofthe direct-developing gastropod, Batillaria cumingi, were investigatedon two shores of differing substrata. Sand-mud shore and rockyshore populations differed in size structure; first-year snailswere ca. 7 mm in shell length (SL) in both populations, whereassecond-year snails, merging with older cohorts, measured 15–25mm SL in the sand-mud shore population and ca. 15 mm SL in therocky shore population. Egg distribution matched adult distributionin the sand-mud shore population, but was more restricted thanthat of adults in the rocky shore population. The distributionof newly-hatched juveniles (0–1 mm SL) was restricted inboth populations, but the growth stage at which snails extendedtheir distribution differed between the two populations; 1–2.5mm SL on the sand-mud shore and 5 mm SL on the rocky shore. Floatingachieved by early juveniles (ca. 2 mm SL), was commonly observedin the sand-mud population, but rarely in the rocky shore population.The sudden expansion in distribution of the 1–2.5 mm SLgrowth stage in the sand-mud shore population is consideredto have been caused by floating, while expansion of the distributionof older growth stages (>5 mm SL) in the rocky shore populationprobably occurs by crawling. (Received 13 May 1998; accepted 25 August 1998)     

Snail populations in arctic lakes: competition mediated by predation?

Summary For 2 species of snails in arctic Alaskan lakes, I studied the patterns of snail distribution with respect to habitat, distribution of predatory fish, and the potential for interspecific competition. The snails Lymnaea elodes and Valvata lewisi co-exist in these arctic lakes, either in the presence of lake trout, Salvelinus namaycush, or in the absence of predation. Intensive sediment core sampling of Toolik Lake and Lake N-2, with trout and lacking trout, respectively, showed that the smaller snail, Valvata, was abundant in Toolik but ocurred at very low densities in Lake N-2. On the open sediments of lakes containing trout, diver surveys revealed very low densities of adult Lymnaea (0.12±0.12/m²), but similar surveys in lakes without trout revealed much higher densities of adult Lymnaea (7.1±1.8/m²). A survey of 14 lakes indicated that adult Lymnaea grew to a smaller mean size in lakes with trout than in lakes which lacked trout.In laboratory and field experiments, the presence of Lymnaea lowered the fecundity of Valvata. Laboratory experiments also showed that Lymnaea fecundity was enhanced by the presence of Valvata. Enhancement was not due to predation by Lymnaea on Valvata eggs or young. The observed patterns of distribution and abundance in the absence of trout, combined with results from laboratory experiments, are consistent with the hypothesis that competitive and facilitative interactions control the population dynamics of the two snails. The distribution and abundance patterns of snails where trout are present suggest that trout predation rather than competition controls snail population dynamics in lakes containing trout.     

DISPERSAL PATTERNS OF THE GIANT AFRICAN SNAIL, ACHATINA FULICA (FERUSSAC) (STYLOMMATOPHORA: ACHATINIDAE), EQUIPPED WITH A RADIO-TRANSMITTER

The dispersal pattern of the African giant snail, Achatina fulica(Férussac) was examined by radio-tracking for six months.They showed typically nocturnal locomotory activities. Theiractivities differed among three developmental stages; juvenile,young adult, and old adult. Juveniles dispersed for the longestdistances. The most active juvenile moved 500 m in 6 months.Young adults and old adults travelled only within a narrow area,although the dispersal range for young adults was greater thanthat of old adults. (Received 15 September 1992; accepted 11 December 1992)     

The demographics of destruction: isolated populations of arboreal snails and sustained predation by rats on the island of Moloka’i 1982–2006

Populations of Partulina redfieldi, an achatinelline tree snail studied in four isolated trees, grew 100–900% between 1983 and 1995. Beginning in 1995, populations declined by 85%, and shells of rat-killed snails accumulated beneath the trees. While rat-marked shells were always present in the study area, numbers increased significantly. Despite a rat-abatement program begun in 1995, the snails continued to disappear, which we conclude was due to continued rat migration into the study area, despite baiting, and a switch in rat-food preference toward the snails. In neighboring forest where tree canopies are more continuous, snail density is lower and rat predation is not apparent. Captive-bred snails were successfully introduced to a small unoccupied tree in the same area in 1989, and this population suffered the same fate as the natural snail populations. Since 2000, P. redfieldi populations have remained low and rat predation continues.     

Constrating short-term and long-term effects of predation risk on consumer habitat use and resources

I contrasted the short-term and long-term effects of predationrisk on snail habitat use and resource dynamics. Pulmonate snails(Physella gyrina) were placed into experimental pools and exposedto four levels of predation risk while holding their densityconstant. Periphyton resources were made available in two habitats:open and covered. I hypothesized that a behavioral responseby snails to predation risk would influence periphyton standingcrop in the open and covered habitats. Snails responded to increasingpredation risk by moving into safer (covered) habitats, andthe magnitude of their response was sensitive to the actuallevel of risk: intermediate levels of risk resulted in intermediatehabitat use. However, use of the risky (open) habitat by snailswas time dependent Snails initially responded strongly to predationrisk, but they exhibited similar patterns of habitat use atall risk levels by the end of the experiment Periphyton standingcrop was positively related to predation risk. In contrast tosnail habitat use, this response was initially weak and becamestronger as the experiment progressed. Thus, the short-and long-termresponses of snail habitat use and periphyton standing cropcontrasted sharply. I suggest that the changing patterns ofsnail habitat use over time are consistent with the idea thatsnails balance predation risk against foraging gains when selectinghabitats and that the manner in which they balance foraginggains and predation risk determines the pattern of periphytonstanding crops across habitats     

COPULATION AS FEMALES AND USE OF ALLOSPERM IN THE FRESHWATER SNAIL GENUS BULINUS (GASTROPODA:PLANORBIDAE)

Each of 8 snails in 2 groups of Bulinus (Physopsis) globosus,1 group raised in isolation and 1 group raised in community,were paired for 14 consecutive days with a male-acting partnersnail. In each group, the experimental snails, which were notallowed to act as males, were able to copulate as females onapproximately 94% of the days paired. Two copulations as female,with the same male partner, occurred on 50% of the days thatthe snails were paired, in the 2 groups combined. Non-receptivefemale behaviour by the experimental snails occurred frequently,and copulation was prevented by such behaviour during 6 pairings,3 in each group. Young B. (P.) africanus first copulated as females when theywere 31–33 days old. The accessory sex glands of the femalereproductive tracts of these young female-acting snails containedmoderate to large amounts of secretion. B. (P.) africanus, which were raised in pairs, laidcross-fertilizedeggs in isolation for an average of 76 days, and 1559 eggs/snailwere deposited before cross-fertilization ceased. Cross-fertilizedeggs were produced for as long as 120 days. After 1 copulation as female, virgin B. (P.) africanus laidcross-fertilized eggs for an average of 78 days and deposited3654 eggs/snail before crossfertilization ceased. Cross-fertilizedeggs were produced for as long as 113 days. After 2 copulationsas female, 1 copulation on each of 2 consecutive days, virginB. (P.) africanus laid cross-fertilized eggs for an averageof 102 days and produced 4397 eggs/snail before cross-fertilizationceased. Cross-fertilized eggs were produced for as long as 123days. Snails which were homozygous for an allele governing mantlepigment pattern were raised with a partner which was homozygousfor a different pigment pattern. Young produced in a 4-day periodafter the snails were isolated were 100% heterozygous. The snailswere then rearranged into pairs with a partner of the same genotypefor 4 days, during which time 26% of the young produced werehomozygous. The snails were again isolated for 4 days, and 49%of the young produced during this 4-day period were homozygous.The results of this experiment strongly suggest that multipleoutcrossing occurred. In B. (P.) africanus, stored allosperm were used to fertilizeeggs after 1, 4 and 7 weeks of starvation; after 1 and 4 weeksof 15°C low temperature and 4 weeks of 15°C + 4 weeksof 10°C low temperature; and after 1 and 4 weeks of desiccation.After 8 weeks of desiccation, 2 of 3 surviving snails reproducedby self-fertilization and 1 snail did not reproduce. Too fewsnails survived 8 weeks of desiccation for a conclusion to bereached on the ability of allosperm to survive. (Received 1 June 1984;     

DEVELOPMENT OF ACCESSORY SEXUAL ORGANS IN BIOMPHALARIA GLABRATA (PLANORBIDAE) IN RELATION TO TIMING OF INFECTION BY SCHISTOSOMA MANSONI: CONSEQUENCES FOR ENERGY UTILIZATION PATTERNS BY THE PARASITE

The growth rates of ovotestis and individual accessory sexualorgans (ASO) of Biomphalaria glabrata snails were studied forcontrols and for immature and mature snails infected with Schistosomamansoni. The infection of immature B. glabrata strongly delaysgrowth of the ovotestis and inhibits the development of theaccessory sexual organs. There is no significant differenceup to 2 weeks post infection in the volume of the ovotestisand the ASO between mature infected B. glabrata and controlsnails. From 3 to 4 weeks post infection there was a reductionin the volume of the ovotestis and the ASO of infected matureB. glabrata; then growth of the ovotestis, albumen gland andfemale organs was stopped, but the effect of infection was lessconsistent for the male organs. For a parasite, immature andmature snails have to be considered as two different resourceenvironments, each having at infection time a particular patternof resource allocation, towards growth for juvenile and towardsreproduction for adult snails, changing the possible energyutilization patterns which can be used by the trematode. (Received 29 January 1993; accepted 22 April 1993)     

Predation by ciliates on a metalimnetic Cryptomonas population: feeding rates, impact and effects of vertical migration

Deep algal maxima are frequently overlayed by dense populationsof ciliates, rotifers and crustaceans. This has been interpretedas evidence of heavy predation on the algae, although the impactof this predation has never been determined experimentally.We determined the vertical and seasonal distribution of thealga Cryptomonas phaseolus and its most relevant predators,the ciliates Coleps sp. and Prorodon sp., forming metalimneticmaxima in Lake Cisó. On several dates, in situ feedingrates of ciliates were determined by three independent methods:(i) epifluorescence counts of ingested algal cells togetherwith estimates of the food turnover time of the ciliates; (ii)in situ incubations with radioactively labeled algae: (iii)HPLC determination of alloxanthine content in the predator sizefraction. Feeding rates varied between 0.07 and 0.64 Cryptomonasciliate^–1 h^–1. We then calculated integrated predationon the algae. using the functional response of the ciliatesand the vertical distribution of each population. We found thateven though the ciliates were always food saturated, their predationimpact on Cryptomonas was not very large: as an average, 5–25%of the biomass of Cryptomonas was removed daily by the ciliates.Finally, we studied the effects of the diel vertical movementsof these populations on predation impact. By migrating intothe sulfide-rich hypolimnion during the night, Cryptomonas couldreduce its predation losses by 38%. Thus, the algae were protectedfrom predation during several hours of each diel cycle and maintaineda very large biomass throughout stratification, although thisresulted in a very slow growth. Slow growth, coupled to largebiomass, seems to be a general feature of metalimnetic accumulationsof organisms.     

Interspecific Interactions Among Larval Trematode Parasites of Freshwater and Marine Snails

Freshwater and marine snails serve as intermediate hosts fornumerous species of larval trematodes. Any particular populationof snails may be infected by several species. It is commonlyobserved that mixed species infections are less frequent thanexpected by change in collections of host snails from naturalpopulations. While several mechanisms might generate such negativeassociations, laboratory studies of freshwater snail-trematodeassociations have demonstrated the presence of strong antagonisticinteractions between intramolluscan larval stages (rediae andsporocysts) of species that infect the same host individual.Both predatory and non-predatory antagonism has been observed,the former taking the form of predation by large, dominant redialforms on the sporocysts and rediae of subordinate species. Theseinteractions are largely hierarchical, although in some systemspriority effects have been observed, and in one case a sporocystspecies replaced a redial species by strong non-predatory antagonism.Several instances of positive association between larval trematodespecies have also been observed. In such cases, interferencewith host defense mechanisms by the first parasite appears toenhance superinfection by the second. My own study of the larvaltrematode guild that infects the salt marsh snail, Cerithideacalifornica, has revealed patterns of association and interactionthat are very similar to those demonstrated by laboratory studiesof freshwater systems. Ultimately, the frequency of interactionsamong larval trematodes depends on the availability, relativeto the numbers of susceptible snails, of infective eggs andmiracidial larvae transmitted from definitive hosts.     

Prey responses to pulses of risk and safety: testing the risk allocation hypothesis

The risk allocation hypothesis predicts that prey responses to predation risk should depend on the temporal pattern of risk. In systems where activity is dangerous, predicted activity levels should be ranked as follows: activity during a pulse of safety>activity during continual safety>activity during continual risk>activity during a pulse of risk. We conducted the first experimental test of the basic predictions of the risk allocation hypothesis by examining responses of freshwater physid snails, Physa gyrina, to chemical cues associated with predation on snails by predatory crayfish,Orconectes juvenilis . As predicted, the snails' pattern of activity, microhabitat use and response to risk depended on the temporal pattern of risk. Snails held in continual risk had very low activity levels, but showed an immediate, large increase in activity during a brief period of safety. In contrast, snails held in continual safety showed moderate levels of activity, but surprisingly, only a weak reduction in activity when exposed to a pulse of danger. Further studies are needed to identify general patterns for how temporal variation in risk influences antipredator behaviour.     

GROWTH, SURVIVORSHIP, AND POPULATION SIZE IN THE LAND SNAIL RHAGADA CONVICTA COX, 1870 (PULMONATA: CAMAENIDAE) FROM A SEMIARID ENVIRONMENT IN WESTERN AUSTRALIA

Basic characteristics of a population of the camaenid land snailRhagada convicta were studied in a semiarid environment nearNorth West Cape, in Western Australia. Consistent evidence frommark-recapture studies and analysis of size-frequency distributionindicates an average annual increase in shell diameter of 3.6mm, with an average period of 5 years from hatching to reachadult size. These rates are low, compared with most other landsnails studied, including cama-enids from wetter environments.Estimates of mortality rate and the rate of recruitment intothe adult population indicate that the snails live an averageof approximately 5 years after reaching adult size, which meansthat the period of turnover for this population is approximately10 years. The population density was estimated to be 0.8 adultsper m², with a total population size (± s.e.) of 875± 164. The population is very localised, with an estimateddiameter of the neighbourhood area of only 38 m. (Received 28 August 1990; accepted 12 February 1991)     

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)