An environmentally induced tidal periodicity of microgrowth increment formation in subtidal populations of the clam Ruditapes philippinarum

The periodicity of increment formation in the shell of the Manila clam Ruditapes philippinarum was investigated in the subtidal zone of the Auray River estuary (South Brittany, France). Calcein markings were performed at different periods between May and October 2007 using in situ benthic chambers tented by scuba divers. This study shows that shell microgrowth increments were well-defined and deposited with a tidal periodicity in the subtidal zone, providing the calendar base for high-resolution ecological studies and environmental reconstruction from these R. philippinarum shells. Endogenous rhythmicity in shell microgrowth increment formation and oxygen consumption was previously documented in this species from intertidal flats. Our study suggests that, in the subtidal zone, Manila clams' rhythmic activity may be controlled by such an endogenous process, synchronized by tidal cues. As in other bivalves, R. philippinarum is an osmoconformer euryhaline bivalve. The tidal rhythmicity of shell microgrowth increments in subtidal specimens of this species could be explained by a behavioral adaptation of valve closure at low tide to protect the clam from low salinities and/or to synchronize with food availability. Finally, large inter-individual variability in tidally associated growth rates and asynchronous growth breaks were observed, and could be due to genetic variability between individuals, asynchronous partial spawning events or predation.     

Oxygen and carbon isotope variations in Chamelea gallina shells: Environmental influences and vital effects

Stable isotopes in mollusc shells, together with variable growth rates and other geochemical properties, can register different environmental clues, including seawater temperature, salinity and primary productivity. However, the strict biological control over the construction of biominerals exerted by many calcifying organisms can constrain the use of these organisms for paleoenvironmental reconstructions. Biologically controlled calcification is responsible for the so called vital effects that cause a departure from isotopic equilibrium during shell formation, resulting in lower shell oxygen and carbon compared to the equilibrium value. We investigated shell oxygen and carbon isotopic composition of the bivalve Chamelea gallina in six sites along with a latitudinal gradient on the Adriatic Sea (NE Mediterranean Sea). Seawater δ¹⁸O and δ¹³C_DIC varied from North to South, reflecting variations in seawater temperature, salinity, and chlorophyll concentration among sites. Shell δ¹⁸O and δ¹³C differed among sites and exhibited a wide range of values along with the ~400 km latitudinal gradient, away from isotopic equilibrium for both isotopes. These results hampered the utilization of this bivalve as a proxy for environmental reconstructions, in spite of C. gallina showing promise as a warm temperature proxy. Rigorous calibration studies with a precise insight of environment and shell growth are crucial prior to considering this bivalve as a reliable paleoclimatic archive.     

Shell occupation as a limiting factor for Pagurus brevidactylus (Stimpson, 1859) in the Marine State Park of Laje de Santos,Brazil

Hermit crabs are soft-abdomen decapods. They depend upon gastropod shells for protection, with the latter influencing growth rate, mortality and reproduction. The present work aims to provide information about the biology and reproduction of Pagurus brevidactylus in a conservation area in Brazilian southeastern littoral. The material was sampled through active searching by SCUBA diving and refuge traps. We analyzed population structure, fecundity, reproductive output (RO), reproductive periodicity and shell utilization for P. brevidactylus. In total, 54 males, 18 non-ovigerous females, 34 ovigerous females and 10 juveniles were sampled. The females had a fecundity of 69.5 ± 65 eggs, which was lower than that compared to other populations from different localities. The RO ranged from 1.48% to 63.04%, with a mean of 26.75 ± 18.23%. There was a positive relationship between egg number and female size. The animals had continuous reproduction, but they reached smaller maximum sizes when compared with other populations. The shell occupation pattern was also different from other localities, the shells of Phrontis alba being the most occupied; this shell has a smaller size when compared to others that P. brevidactylus occupies in other localities. Therefore, P. alba could have had limited the growth and reproduction of P. brevidactylus.     

Environmental Heterogeneity and Morphological Variability in Pisidium subtruncatum (Sphaeriidae,Bivalvia)

On the example of P. subtruncatum, a widely distributed bivalve, we studied the relationship between environmental and morphological variability. The main questions addressed were: (1) are there differences in shell morphology between different populations, (2) are they dependent on sediment composition, hydrology or water chemistry, (3) are there differences in morphological variability between stable and unstable environments. Measurements of specimen from nine sampling sites showed high variability in shell shape and size. Length‐height relation, symmetry, width and dry weight of the shell all varied significantly. Shell shape was significantly correlated with sediment composition and dependent on hydrological connectivity, with higher, asymmetric shells occurring in sandy sediments and lower, rounded shells in muddy sediments. Analysis of intrapopulation morphometric variation showed that variability is higher in habitats with high temporal and spatial environmental variability and lower in habitats with low or reduced environmental heterogeneity. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The tropical giant clam <Emphasis Type="Italic">Hippopus hippopus</Emphasis> shell,a new archive of environmental conditions as revealed by sclerochronological and δ<Superscript>18</Superscript>O profiles

The use of the sclerochronology and geochemistry of a New Caledonian (South West Pacific) giant clam Hippopus hippopus shell as markers of environmental changes has been investigated. Growth increment thickness and δ¹⁸O ratios were measured on 4 years of shell growth of a modern specimen. During the last year, this giant clam was placed in a tank equipped for high-frequency environmental monitoring. Because shell is secreted in isotopic equilibrium with the seawater, the palaeo-sea surface temperature (SST) equation obtained faithfully reproduces the seasonal SST amplitudes. Growth increment thickness changes are seasonal and, for more than 50%, governed by the SST changes. The transplantation from the in situ site to the tank and reproduction events reduces the strength of growth and SST relationships. Nevertheless, growth increment thickness measurements can give information on average, minimal and maximal past SST in diagenetically altered shells. A peculiar growth event characterized by a short, drastic and significant decrease has been identified and attributed to an intense upwelling event. This study further highlights the use of giant clam shell δ¹⁸O as a SST proxy but also demonstrates that H. hippopus growth increment thickness changes provide useful information on past environmental settings and on exceptional events, for example, intense upwellings.     

Environmental and physiological controls on shell microgrowth pattern of Ruditapes philippinarum (Bivalvia: Veneridae) from Japan

The reproductive cycle and shell microgrowth patterns of thevenerid bivalve Ruditapes philippinarum from Matsukawa-ura,a small inlet facing the Pacific Ocean, northeastern Japan wereexamined. Histological examination of the gonads revealed thatspawning in this species occurred twice in 2005, once betweenlate June and early August and the other between late Septemberand early October. Comparison of the shell microgrowth patternswith the developmental stages of the gonad in each individualrevealed that in spawning individuals (mature and spawning stages),the mean lunar-day growth rates were significantly smaller thanthose in individuals which were not in spawning condition (spent,recovery and growing stages). In non-spawning individuals, themean lunar-day growth rates were positively correlated withseawater temperature up to 20°C. However, in spawning individuals,no correlation was observed between shell growth and seawatertemperatures >16°C. These facts suggest that physiologicalstress during reproduction has a negative influence on shellgrowth and results in spawning breaks. The present study indicatethat spawning breaks can be used to identify the timing of sexualmaturity and to count the number of spawning events occurringper year in extant and fossil bivalves.     

ADAPTATION FROM RESTRICTED GEOMETRIES: THE SHELL INCLINATION OF TERRESTRIAL GASTROPODS

The adaptations that occur for support and protection can be studied with regard to the optimal structure that balances these objectives with any imposed constraints. The shell inclination of terrestrial gastropods is an appropriate model to address this problem. In this study, we examined how gastropods improve shell angles to well‐balanced ones from geometrically constrained shapes. Our geometric analysis and physical analysis showed that constantly coiled shells are constrained from adopting a well‐balanced angle; the shell angle of such basic shells tends to increase as the spire index (shell height/width) increases, although the optimum angle for stability is 90° for flat shells and 0° for tall shells. Furthermore, we estimated the influences of the geometric rule and the functional demands on actual shells by measuring the shell angles of both resting and active snails. We found that terrestrial gastropods have shell angles that are suited for balance. The growth lines of the shells indicated that this adaptation depends on the deflection of the last whorl: the apertures of flat shells are deflected downward, whereas those of tall shells are deflected upward. Our observations of active snails demonstrated that the animals hold their shells at better balanced angles than inactive snails.     

Biomineralization plasticity and environmental heterogeneity predict geographical resilience patterns of foundation species to future change

Although geographical patterns of species' sensitivity to environmental changes are defined by interacting multiple stressors, little is known about compensatory processes shaping regional differences in organismal vulnerability. Here, we examine large‐scale spatial variations in biomineralization under heterogeneous environmental gradients of temperature, salinity and food availability across a 30° latitudinal range (3,334 km), to test whether plasticity in calcareous shell production and composition, from juveniles to large adults, mediates geographical patterns of resilience to climate change in critical foundation species, the mussels Mytilus edulis and M. trossulus. We find shell calcification decreased towards high latitude, with mussels producing thinner shells with a higher organic content in polar than temperate regions. Salinity was the best predictor of within‐region differences in mussel shell deposition, mineral and organic composition. In polar, subpolar, and Baltic low‐salinity environments, mussels produced thin shells with a thicker external organic layer (periostracum), and an increased proportion of calcite (prismatic layer, as opposed to aragonite) and organic matrix, providing potentially higher resistance against dissolution in more corrosive waters. Conversely, in temperate, higher salinity regimes, thicker, more calcified shells with a higher aragonite (nacreous layer) proportion were deposited, which suggests enhanced protection under increased predation pressure. Interacting effects of salinity and food availability on mussel shell composition predict the deposition of a thicker periostracum and organic‐enriched prismatic layer under forecasted future environmental conditions, suggesting a capacity for increased protection of high‐latitude populations from ocean acidification. These findings support biomineralization plasticity as a potentially advantageous compensatory mechanism conferring Mytilus species a protective capacity for quantitative and qualitative trade‐offs in shell deposition as a response to regional alterations of abiotic and biotic conditions in future environments. Our work illustrates that compensatory mechanisms, driving plastic responses to the spatial structure of multiple stressors, can define geographical patterns of unanticipated species resilience to global environmental change.     

Seasonal change in xylem growth of <Emphasis Type="Italic">Pinus densiflora</Emphasis> in central Japan

This study investigated the seasonal change in xylem growth of Japanese red pine (Pinus densiflora). Wood cores were sampled at 2-week intervals from April to November in 2012 using the microcoring method. Daily increment rates of tracheid number and tree-ring width were compared with seasonal changes in daily mean temperature and photoperiod. Xylem growth started in early to late May and stopped in late October to early November. The maximum daily increment rates of tracheid number and tree-ring width were in early July. The 95 % confidence intervals of the timing of the maximum daily increment rates included the summer solstice (23 June) with the longest photoperiod, but not the warmest day (30 July). The maximum daily increment rate of xylem growth is thought to be controlled by the photoperiod rather than by temperature. The daily mean temperature exceeded 20 °C after the summer solstice, indicating that temperature is not a limiting factor for xylem growth. This study suggests that the timing of maximum daily increment rates of xylem growth of P. densiflora is controlled by the photoperiod.     

Growth line deposition and variability in growth of two circumpolar bivalves (Serripes groenlandicus, and Clinocardium ciliatum)

Growth patterns of two common circumpolar bivalves, the Greenland cockle (Serripes groenlandicus), and the hairy cockle (Clinocardium ciliatum) have been used in previous studies to reconstruct environmental conditions in the arctic. To date, there has been no direct determination that growth lines in either species are deposited periodically, and there has been no examination of factors affecting growth. We placed calcein-marked individuals of both species on oceanographic moorings in two fjords (Rijpfjord and Kongsfjord) in the Svalbard archipelago for one and two (Kongsfjord only) years. Growth patterns were compared with concurrent in situ temperature and fluorescence data in order to assess environmental controls on growth. Dark growth lines are evident on the outer shell surface and internally in shell cross section in both S. groenlandicus and C. ciliatum, and both species deposited only one line per year, unequivocally confirming that internal lines are deposited annually. Growth line deposition in both species began in late summer to early fall, before the seasonal decline in temperature. There was no difference in growth of S. groenlandicus between the two fjords despite differences in water temperature (3°C), fluorescence (nearly threefold) and the onset and duration of the winter season. C. ciliatum, however, grew approximately 2.8 times faster in the warmer, more food-rich Kongsfjord than in Rijpfjord. Subannual lines were counted in two individuals of each species from each fjord, but deposition of these lines was not clearly related to number of growing days estimated by temperature and fluorescence.     

Mineral composition variation in the shells of freshwater mussel Anodonta woodiana at different growth stages

The levels of mineral element Na, Mg, K, Ca, Cr, Mn, Fe, Co, Ni, Cu, Zn, Se, Mo, Al, As, Ag, Cd, and Tl were quantified in the whole shells of the freshwater bivalve Anodonta woodiana at three different growth stages (i.e. J1 juveniles of 1 month old, J2 juveniles of 3.5 months old, and adults of 36 months old). The concentrations of Na and Al were different between different growth stages (p < 0.05). The highest Na concentrations (2715 ± 86 μg/g dry weight) were found in J2 juveniles. The highest Al concentrations (303.9 ± 5.95 μg/g dry weight) were found in J1 juveniles. Manganese concentrations (517.0 ± 47.98 μg/g dry weight) were significantly higher in J2 juveniles than in J1 juveniles (432.3 ± 9.87 μg/g dry weight) (p < 0.05). Copper concentrations (27.32 ± 0.15 μg/g dry weight) were significantly higher in J1 juveniles than in J2 juveniles (26.21 ± 0.86 μg/g dry weight) and adults (24.74 ± 1.43 μg/g dry weight) (p < 0.05). Burdens of Na, Ca, Mn, Fe, Co, Cu, Mo, Ag, and Tl were positively correlated with the shell length (p < 0.05). These findings can possibly contribute to an understanding of elemental requirements for shell growth and, hence, facilitate improvement of survival and growth rates during artificial mussel culture.     

Regeneration of excised shell by the invasive apple snail Pomacea canaliculata

After excision of shell from Pomacea canaliculata, survival rates, repair process, haemocyte response, and calcium content were observed and quantified for three weeks. The following experimental treatments were used: black-colored individuals with large (BL-group) or small (BS-group) shell heights, yellow-colored individuals with large (YL-group) or small (YS-group) shell heights. The survival rates in BL-, YL-, BS-, and YS-groups after making partial excisions of snail shells were, respectively, 93.59, 94.87, 92.31, and 96.15%. Freshly regenerated shell could be seen at 1 day among the four groups after induction of shell regeneration. Regeneration shell filled the wound after 5–10 days. The area of regenerated shell in yellow-colored individuals was larger than that of black-colored individuals, but the thickness of regenerated shell was not significantly different. Multiple (four times) inductions of shell regeneration significantly enhanced the thickness of newly formed shells. Two types of circulating haemocytes (hyalinocytes and granulocytes) were identified under light microscopy based on Wright’s staining. Artificial induction of shell regeneration rendered a significant increase in the number of total circulating haemocytes which was followed by a decrease to pre-wounding levels. This peak value of total circulating haemocytes was 2.86, 2.43, 1.69, and 1.71-fold higher than pre-wounding levels in group BL, YL, BS, and YS, respectively. The number of total circulating haemocytes in yellow-colored individuals was significantly higher than in black-colored. Calcium concentrations in the mantles of snails are favorable for calcium nucleation. It is postulated that calcium content increased in the mantle increase at the start of the experiment and then returned to pre-wounding levels at the end of the experiment. This study shows that apple snails have effective shell regeneration abilities and that haemocytes and calcium transportation appear to play an important role in shell growth and regeneration.     

Shell morphometrics and growth rate of the invasive bivalve mollusc Mytilus galloprovincialis in the Knysna estuarine embayment,South Africa

Shell morphometrics of the invasive mussel Mytilus galloprovincialis were compared at five sites and growth rate at four sites (in four seasons) in the Knysna estuarine embayment. Mussels from two sites (The Heads, Leisure Isle) where wave action was present had shells significantly lower for any length when compared with other more sheltered sites. There was no significant difference in shell width of mussels for any given length among sites. Mussels from The Heads had thicker shells than other sites, and those from Leisure Isle thicker shells than three other embayment sites where shells did not differ in thickness. Growth rate of mussels at two embayment sites (Thesen’s Wharf and Thesen Islands Marina) was greatest in autumn and summer whereas at The Heads and Leisure Isle there was little seasonal difference in growth rate. Growth rate of mussels at Thesen’s Wharf and Thesen Islands Marina was mainly greater in all seasons when compared with mussels at The Heads and Leisure Isle. The more rapid growth rate of mussels at the sheltered embayment sites might in part explain why M. galloprovincialis now dominates the mid- to lower intertidal on hard substrata in this region of the Knysna estuary.     

Microgrowth patterns in the shell of the Malaysian cockle Anadara granosa (L.) and their use in age determination

Marked specimens of the Malaysian cockle Anadara granosa (L.) were placed in buried cages (filled with sediment from the surrounding substratum) which were located in intertidal and subtidal environments in an area between Penang Island and mainland West Malaysia. In the majority of cockle shells examined the number of growth bands deposited was close to the number of tidal periods. Bands in the shells of intertidal animals were more strongly defined than those in shells continuously immersed. The banding pattern in shells from the subtidal environments showed narrow increments during spring tides alternating with a few wider increments during neap tides. The growth increments between bands correspond to tidal periods so they can be used to record growth rates and provide estimates of the ages of the shells. Samples of cockles collected from four sites in a commercial cockle-culture area off the coast of West Malaysia were aged. Two of the sites were located in an estuary where the cockles were periodically exposed to fluctuating salinities during the intermonsoon period. Cockles from the other two sites were situated away from the estuary and experienced full strength sea water. There was good agreement between the calculated estimates of the ages determined from the tidal bands and their known ages. Shells of cockles from the estuary had characteristically marked banding patterns which may be related to shell deposition during the intermonsoon period when the cockles were exposed to sea water of low salinity. Cockle shells collected outside the estuary did not display these patterns.     

Relative growth and reproductive cycle of the date mussel Lithophaga lithophaga (Linnaeus, 1758) sampled from the Bizerte Bay (Northern Tunisia)

Relative growth and the reproductive cycle of the date mussel Lithophaga lithophaga were studied from September 2002 to October 2003 in the Bizerte Bay (Northern Tunisia). Relationships between shell length and shell width, shell height as well as volume-related variables (shell dry weight, soft tissues dry weight and total wet weight) showed negative allometries in both sexes except for shell width in males (isometry) and shell height in both sexes (positive allometry). The sex ratio was unbalanced, within the length range of 6–48 mm, 49.43 % of the animals were males, 24.39 % females and 26.17 % sexually undifferentiated; within the length range of 49–92 mm, the respective values were 41.19, 52.7 and 6.1 %. Histological investigations as well as analyses of the condition index and the gonadosomatic index (CI and GSI) revealed the presence of a single reproductive cycle per year. Spawning occurred at the end of August and early September and was associated with a decrease in seawater temperature and salinity. A resting phase occurred in winter, coinciding with the lowest water temperatures. Histological examinations of the gonads of a total of 130 specimens revealed only two cases of hermaphroditism. The present study constitutes a useful baseline for a sustainable management of local wild stocks of L. lithophaga.     

Observations on the shell morphology of two species of cockle Cerastoderma edule and C. glaucum

Aspects of the shell morphology of the cockles Cerastoderma edule (L.) and C. glaucum (Poiret) collected from the R. Crouch and R. Roach (Essex, England) are considered with reference to interpretation of the environment of sub-fossil assemblages. Discrimination between the two cockle species is best achieved by assessment of qualitative shell characters, quantitative measurements being of rather limited value. Factors influencing shell rib-number differ in the two species of Cerastoderma; within a common environment C. glaucum has fewer ribs than C. edule . Mean rib-number of C. edule is directly related to salinity over the colonized salinity range 20–36.5%₀ whilst in C. glaucum rib-number is lowest in both low (<10%₀) and high (>35%₀) salinity environments, being maximal where salinities between 20–30%₀ occur. Internal shell pigmentation may be absent in C. glaucum from low salinity environments. Magnesium content of whole shells is similar in the two cockle species from a mixed population. In both cockles, however, the prismatic layer contained more magnesium than the nacreous shell layer. The most useful molluscan indicators of brackish water lagoon environments are C. glaucum and Hydrobia ventrosa; these species being absent from areas of only limited wave action. C. edule may be associated with marine or estuarine molluscan indicators.     

QUANTITATIVE GENETICS OF SHELL FORM OF AN INTERTIDAL SNAIL: CONSTRAINTS ON SHORT-TERM RESPONSE TO SELECTION

We investigated the genetic and environmental determinants of shell form in an intertidal snail (Prosobranchia: Littorina sp.) to identify constraints on the short-term response to selection. Our quantitative genetic parameters were estimated from a half-sib experimental design using 288 broods of snails. Each brood was divided into two treatments differing in snail population density, and therefore in grazing area per snail. Differences in population density induced marked differences in shell form. Snails in the low density treatment grew faster and had lighter shells with narrower whorls and narrower apertures than their siblings at high density. Despite this environmental plasticity in shell shape we found significant additive genetic variance for components of shell shape. We discuss two mechanisms that may maintain additive genetic variance for shell shape in intertidal snail populations: migration between environments with different selective pressures and migration between environments with different mean growth rates. We also estimated a genetic variance-covariance matrix for shell form traits and used the matrix to identify constraints on the short-term response to selection. We predict the rate of response to selection for predator-resistant morphology such as would occur upon invasion of predatory crabs. The large negative genetic correlation between relative spire height and shell weight would facilitate simultaneous selection for a lower spire and a heavier shell, both of which would increase resistance to predatory crabs.     

Relating annual increments of the endangered Blanding's turtle plastron growth to climate

This research is the first published study to report a relationship between climate variables and plastron growth increments of turtles, in this case the endangered Nova Scotia Blanding's turtle (Emydoidea blandingii). We used techniques and software common to the discipline of dendrochronology to successfully cross‐date our growth increment data series, to detrend and average our series of 80 immature Blanding's turtles into one common chronology, and to seek correlations between the chronology and environmental temperature and precipitation variables. Our cross‐dated chronology had a series intercorrelation of 0.441 (above 99% confidence interval), an average mean sensitivity of 0.293, and an average unfiltered autocorrelation of 0.377. Our master chronology represented increments from 1975 to 2007 (33 years), with index values ranging from a low of 0.688 in 2006 to a high of 1.303 in 1977. Univariate climate response function analysis on mean monthly air temperature and precipitation values revealed a positive correlation with the previous year's May temperature and current year's August temperature; a negative correlation with the previous year's October temperature; and no significant correlation with precipitation. These techniques for determining growth increment response to environmental variables should be applicable to other turtle species and merit further exploration.     

Stacking Increments: A New Model and Morphospace for the Analysis of Bivalve Shell Growth

A model employing stacking increments is introduced for the analysis of bivalve shell growth and form. The model is based on the components of shell growth that are potentially independent: the rate of mantle cell proliferation, the rate of precipitation of shell material, and the rate of translation of the pallial line, where the mantle is attached to the shell. This model is defined in terms of the following parameters: (1) the ratio of accretion of shell material at the shell margin to growth of the mantle by cell division, (2) the ratio of shell accretion at the pallial line to mantle growth, and (3) the ratio of the amount of pallial muscle translation, away from the umbo toward the shell margin, to mantle growth. In this model, the shape of a radial section through the shell is simulated by stacking of internal microgrowth increments. The mode of stacking of the increments is determined by the balance among the parameters defining growth. A theoretical morphospace defined on the basis of this model is largely consistent with the range of forms of naturally occurring bivalve shells. Analysis of the distribution of actual shell forms in relation to this morphospace suggests that the absolute rate of shell precipitation and the gradient in precipitation rate away from the shell margin along a radial cross-section are physiologically as well as geometrically constrained.     

An estimate of the sustainable rate of shell extraction from the Dutch Wadden Sea

1. Shell production by cockles Cerastoderma edule was studied to examine whether or not the present licensed rate of shell extraction in the Dutch Wadden Sea exceeds the current rate of shell addition to the exploitable stocks.
2. Long-term data on numbers of cockles and weights of their shells were used to estimate their annual production on Balgzand, a 50-km² tidal flat area in the western-most part of the Wadden Sea. During the 1969–97 period, it amounted to an average of 125 g m^–2, including 107 g m^–2 of shells large enough to be exploitable for shell-lime fishery.
3. The very irregular annual recruitment of cockles was the main cause of the wide 95% confidence limits (74 and 140 g m^–2 year^–1) of this 28-year estimate. Moreover, high mortality rates in severe winters substantially reduced production per recruit in some year classes.
4. About one-third of the estimated production does not reach exploitable stocks, because it is fragmented by birds (particularly eider ducks), permanently buried in the sediment, or removed by the fishery for live cockles.
5. During the last few decades, the estimated mean amount added annually to the exploitable stocks was 88 million kg or 132 000 m³ of large cockle shells. This amount compares favourably with the current annual level of removal of 134 000 m³ of shells, three-quarters of which are cockles.
6. Even at temporarily lower production rates, the exploitation of shell stocks at its present rate is not expected to lead to a rapid exhaustion of the existing stocks in the tidal inlets of the Dutch Wadden Sea, as these stocks will be in the order of a few million m³.