Annual variation in the growth rate of Yoldia notabilis (Bivalvia: Nuculanidae) in Otsuchi Bay,northeastern Japan,analyzed using shell microgrowth patterns

Annual variation in the growth rate of the protobranch bivalve Yoldia notabilis (collected in Otsuchi Bay, northeastern Japan between 1989 and 1991) was determined by measuring shell length at successive growth lines. Factors affecting growth rates were assessed by examining long-term environmental data. Shell cross-sections of Y. notabilis showed a clear pattern of internal growth lines which formed simultaneously with the annual external lines on the outer shell surface. Hence, they were used as an age marker. A maximum lifespan of 17 yr was determined for this species, and a sigmoidal growth curve was obtained by the internal growth line analysis. Examination of the growth lines also revealed a large annual variation in growth rate, fluctuating as much as 32-fold during the past 9 yr. The variation correlated negatively with water temperature and positively with chlorophyll a content in the water column. It is suggested that the variation in annual growth rate is dependent on food supply during the spring phytoplankton bloom which varies from year to year according to the flow of the cold Oyashio current each spring.     

Analysis of growth rate in Mya arenaria using the Von Bertalanffy equation

Field studies were conducted in Gloucester, Massachusetts, USA, to determine linear shell growth rates for Mya arenaria. These rates were then compared with those reported for the same species from other locations. Most shell deposition occurred from March through November of each year. Winter interruptions in growth were not as marked in the small clams as in the larger ones (>60.0 mm). Annual variations in growth were slight during the period 1973–1974. Growth of mature clams (>35.0 mm) slowed during the spawning season. No significant sexual dimorphism in mean annual growth rates was detected. Winter rings were shown to be a reliable method for determining age in clams from Gloucester. Age-size relationships, based on two independent measures of annual growth, winter rings and tagging experiments, were computed using the Von Bertalanffy growth equation. No well-defined latitudinal patcerns in growth could be established for M. arenaria.     

Isotopic fractionation between seawater and the shell of <Emphasis Type="Italic">Scrobicularia plana</Emphasis> (Bivalvia) and its application for age validation

This study analyzed the isotopic profiles of four aragonitic shells of Scrobicularia plana in conjunction with measured seawater temperatures and salinities. Comparison of δ¹⁸O_SHELL with expected values revealed fractionation of δ¹⁸O in near equilibrium with the ambient environment. Growth cessation occurred between November and March. Carbonate deposition stopped when temperatures were <12°C. Analysis of δ¹³C_SHELL values suggested that carbon in the shell does not reflect the DIC in ambient water, likely due to the incorporation of metabolic carbon. An ontogenetic trend of increasing δ¹³C values over time was observed, likely related to changes in metabolic activity. Annual growth patterns were inferred from δ¹⁸O_SHELL profiles and compared with internal and external growth lines. Estimations of age based on external lines were unreliable, resulting in overestimation of age and underestimation of growth rates, likely due to the disturbance lines being wrongly identified as annual. Analysis of internal lines may lead to over- or underestimation of age and was more reliable in recent portions of the shell.     

Growth in the bivalve Yoldia eightsi at Signy Island,Antarctica, determined from internal shell increments and calcium-45 incorporation

The growth rate of the infaunal nuculanid bivalve Yoldia eightsi at Factory Cove, Signy Island, South Orkney Islands (maritime Antarctica), was estimated from internal shell increments and ⁴⁵Ca incorporation of individuals collected monthly from December 1987 to April 1989. Acetate peels of etched shells revealed clear first-order increments, with less well defined, narrower, second-and third-order increments. The first-order increments were assumed to be annual, although there is no independent confirmation of this assumption. Unfortunately abrasion of the umbo region and the small thin shells of Y. eightsi meant that in no case could a complete sequence of increments be measured realiably on any individual shell. Measurements of 1043 first-order increments from 130 shells where a minimum of two consecutive increments could be detected were therefore pooled, and a population growth curve constructed from a Ford-Walford plot. This indicated a slow growth rate, with a maximum shell height of 22.3 mm (equivalent to a shell length of 35.6 mm) being reached at an age >60 yr. The size-frequency distribution of 1521 individuals pooled from winter (July to October) samples revealed a distinct lack of smaller (younger) individuals, possibly reflecting poor recruitment in areas of dense adult populations. The largest shell recovered in the samples was 33.5 mm in length, with an estimated age of 52 yr. Short-term ⁴⁵Ca-incorporation experiments indicated a mean daily rate of growth increment of 3.8 m for individuals of 12 mm shell height, which matches the proposed annual growth rate if growth is assumed to occur for about 150 d each year and the first-order increments are assumed to be annual.     

Age,growth and population structure of <Emphasis Type="Italic">Modiolus barbatus</Emphasis> from the Adriatic

Age, growth and population structure of Modiolus barbatus from Mali Ston Bay, Croatia were determined using modal size (age) classes in length frequency distributions, annual pallial line scars on the inner shell surface, internal annual growth lines in shell sections of the middle nacreous layer and Calcein marked and transplanted mussels. The length frequency distributions indicated that M. barbatus attain a length of ∼40 mm in 5–6 years indicating that a large proportion of the population in Mali Ston Bay is <5 years old. Some mussels of ∼60 mm were predicted to be 14 years old using the Von Bertalanffy growth (VBG) equation. Up to the first 6 pallial line scars were visible in young (<6 years) mussels but in older shells the first scars became obscured by nacre deposition as the mussel increased in length and age. The age of the older shells (>6 years) was determined from the middle nacreous lines in shell section, which formed annually in winter between February and March; the wider dark increments forming during summer (June to September). The oldest mussel, determined from the middle nacreous lines, was >12 years, with the majority of mussels aged between 3 and 6 years of age. The ages of mussels ascertained using the growth lines were not dissimilar to the ages predicted from the length frequency distributions. Age at length curves produced using modal size class data were not different from the data obtained using the pallial scar rings and internal growth lines. Taken together these data suggest that M. barbatus attains a length of 40 and 50 mm within 5 and 8 years, respectively. Eighty one percent of individual M. barbatus injected with a Calcein seawater solution (300 mg Calcein l⁻¹), into their mantle cavity successfully deposited a fluorescent line, which was visible in suitably prepared shell sections under ultra violet light. Incorporation of Calcein into the mussel shells was seasonally variable with the lowest frequency of incorporation in mussels marked in February and recovered in May. Seasonal shell growth was observed with significantly higher growth rates in mussels marked in May and removed in August (ANCOVA, F _3,149 = 23.11, P < 0.001). Mussels (∼18 to 22 mm) marked in May and recovered in August displayed maximal growth rates of >2.5 mm month⁻¹ compared with a mean mussel growth rate of 1.2 ± 0.6 mm month⁻¹. At other times of the year mussel shell growth ranged from immeasurable to 1.48 mm month⁻¹.     

Population structure and growth of Donax trunculus (Bivalvia: Donacidae) in the western Mediterranean

A comparison of shell growth in Donax trunculus (collected between 1988 and 1990 of Cullera, Spain) has been carried out using an analysis of cohort progression in monthly length frequency distributions, hyaline surface shell growth rings and internal microgrowth bands. In the Mediterranean there are two periods of recruitment of D. trunculus, one in the summer (July to September) and the other in winter (December to February). Clams recruited to the population in winter display a clear cessation in shell growth during the following summer which may possibly be correlated with spawning, whereas individuals of the summer recruited cohort show no growth cessation the following summer and continue to deposit shell during this period. The normally opaque shell of D.trunculus reveals the presence of translucent hyaline growth rings when the shells are backlit by a strong light source, and these have been shown to be laid down in the shell during summer months. Formation of a hyaline ring is accompanied by a narrowing of the microgrowth patterns present in shell sections. Both the hyaline rings and the length frequency distributions have been used to determine the age and growth rate of D. trunculus.     

Age and growth of the naticid gastropod <Emphasis Type="Italic">Polinices pulchellus</Emphasis> (Gastropoda: Naticidae) based on length frequency analysis and statolith growth rings

In Red Wharf Bay, UK the naticid gastropod, Polinices pulchellus, was more abundant and more highly aggregated during the summer months (June–August 2001) than during the winter (December 2000). Whilst small numbers of juvenile P. pulchellus (4–6 mm shell length) were present throughout the year the population consisted mainly of individuals of 12–14 mm shell length. Juvenile snails grew rapidly in size during the winter and early spring; growth then virtually ceased between May and June, following which there was a further period of rapid growth between August and February. Densities ranged between 57 and 4,073 ha⁻¹ and the largest individual collected during this investigation measured 16.2 mm in shell length. Statoliths from adult P. pulchellus revealed the presence of a settlement ring and two prominent growth rings (rings 1 and 2). A curvilinear relationship exists between statolith diameter and shell length in snails up to 16 mm in length. Settlement rings ranged in diameter from 19.7 to 45.2 μm (mean 29.8 μm; SE=0.41) giving an estimated shell length of the settled juvenile of 1.1 mm. The diameter of ring 1 and ring 2 were significantly correlated indicating that rapid growth during the first year is maintained during year 2. Shell lengths estimated from the diameters of the prominent statolith rings and those obtained from length frequency data analysis (LFDA), were broadly congruent strongly suggesting an annual periodicity to the statolith rings. The largest snails (>15 mm) present within this population were estimated to be between 2 and 3 years old. Von Bertallanfy seasonal growth curves obtained from the LFDA predicted values of L∞, K and t ₀ of 14.32 mm, 1.54 and −0.14 years, respectively, suggesting that P. pulchellus rapidly attains its maximum asymptotic size.     

Age and growth of the fan mussel Pinna nobilis from south-east Spanish Mediterranean seagrass (Posidonia oceanica) meadows

The calcitic and aragonitic shell of the fan mussel Pinna nobilis L. contains a record of the environmental changes experienced during its growth. Stable-isotope analyses of oxygen (¹⁸O:¹⁶O) in shell carbonate from the calcitic outer shell-layer have been used to validate the periodicity of clearly defined concentric rings on the aragonitic posterior adductor-muscle scar and to estimate the age and growth of fan mussels growing in Posidonia oceanica (L.) meadows at four locations on the south-east Spanish Mediterranean coast. The stable oxygen-isotope records obtained at intervals along a profile across the shell surface enabled seasonal changes in water temperature to be established, and hence seasonal patterns of shell growth to be inferred. Muscle-scar rings were found to be deposited annually in the shell in the spring and early summer (a period of increasing water temperatures), and represent an interruption in the migration of the posterior adductor muscle along the inner surface of the shell. In small pinnids (<25 cm) accretion of the shell is rapid during the first year, but in the second year it is distinctly slower than at the same time the previous year. This slowing down in growth during the second year coincides with the appearance of the “first” distinct muscle-scar ring, indicating that Pinna nobilis does not form a muscle-scar ring during its first year of shell growth. Maximum growth rates were recorded amongst pinnids from Carboneras, where they achieved a length of 59 cm in 8 yr, whilst those from Aguamarga were estimated to be the oldest (attaining a length of 45 cm in 13 yr). Received: 26 January 1998 / Accepted: 8 October 1998     

Combined sclerochronologic and oxygen isotope analysis of gastropod shells (Gibbula cineraria, North Sea): life-history traits and utility as a high-resolution environmental archive for kelp forests

The grey top-shell, Gibbula cineraria is a common member of temperate to cold water kelp forest communities, but its longevity and the age structure of its populations remains unresolved. Combined measurements of shell growth rates (sclerochronology) and oxygen isotope composition allow analysis of rate and timing of shell growth. Eight specimens were analyzed from the southern North Sea (near Helgoland, German Bight). Three age groups were identified but external measurements (width, height, ornamentation patterns and number of whorls) and shell weight are not adequate for ontogenetic age discrimination. Stable oxygen isotope data is consistent with shell growth during the interval from April to December in isotopic equilibrium with seawater, and growth increments exhibit strong tidal controls with fortnightly bundles well preserved. Reliable environmental proxy data (water temperature) can be extracted from the shell aragonite using conventional stable oxygen isotope analyses, with a temporal resolution of days attainable during intervals of maximum growth, but annual extremes are not always recorded in the shell. While demonstrating the utility of G. cineraria as a environmental and potential paleoenvironmental proxy for kelp forest habitats, its longevity has been significantly overestimated.     

Growth studies of the tropical intertidal limpet Acmaea antillarum

The tropical intertidal limpet Acmaea antillarum (Sowerby) was used in a series of field and aquarium growth studies, carried out in Venezuela. Both field and aquarium experiments demonstrated the formation of diurnal growth ridges and subdiurnal periodic growth striations on the shell surface. It was shown in aquarium experiments that the normal growth pattern was inhibited in constant darkness and that continous illumination stimulated the formation of extra growth ridges. Alterations of the experimental sequence of illumination influenced the formation of subdiurnal growth markings. This secondary growth pattern appears to be distinct from the basic system of diurnal ridge formation. Limpet shell lengths were related to estimated age based on diurnal growth ridge counts. It is probable that maximum shell size is attained in less than 1 year. Equations for allometric shell growth characteristics were calculated relative to shell length. Calculations for instantaneous relative growth rate were made from natural populations, experimental field populations and aquarium specimens. These were compared to show that the field and aquarium experiments demonstrated similar growth rates. The results of these observations and calculations are compared with other molluscan growth studies.     

Short-term shell growth in bivalves: Individual,regional, and age-related variations in the rhythym of deposition ofCerastoderma (=Cardium) edule

The influence of potential sources of variations (age, site, region) on the production of shell microgrowth increments was studied in the cockleCerastoderma edule (L.) from the shores of Denmark, Jersey Island and South Wales between 1978 and 1986. Microgrowth increments in the shell of the cockle were counted between annual growth marks corresponding to the second, third, and fourth yearly growth period of the cockle. The number of microgrowth increments per year showed a decrease with increasing age. Increments formed after deposition of the last annual mark in cockles sampled at different dates indicate that the rate of increment formation differed between age classes and populations. Furthermore, we observed pronounced intrapopulation differences in the number of increments for specimens of the same age, and differences in both number and clarity of increments in subtidal and intertidal populations. Our results do not support the hypothesis of a general endogenous rhythm of microgrowth that closely corresponds to tidal rhythms inCerastoderma edule. Instead, they emphasize the plasticity of deposition according to the origin and age of the experimental specimens.     

Age and growth in <Emphasis Type="Italic">Odontocymbiola magellanica</Emphasis> (Gastropoda: Volutidae) from Golfo Nuevo,Patagonia, Argentina

Growth, age and somatic production of the benthic predator Odontocymbiola magellanica were studied in Golfo Nuevo (42°S 65°W), on the South American Atlantic shelf. Stable oxygen isotope ratios confirmed semiannual formation of internal and external shell growth marks. Mean shell length (SL) of females was 115 and 112 mm for males, while population modal shell-free wet mass (SFWM) was 62.8 g. A Gompertz growth function (SL_∞ = 200 mm, K = 0.197, t ₀ = 5.486) fitted 113 pairs of size-at-age data (12 shells) best. O. magellanica is a long-lived species, reaching up to 20 years of age. The maximum individual somatic production of 29.3 g SFWM per year is attained at 145 mm SL, which corresponds to about 12 years of age. The life span of this volutid seems to be twice compared with other large gastropods. O. magellanica is a valuable and exploitable resource regarding its large size and somatic production, but on the other hand, its slow growth, late maturity and direct development makes it extremely vulnerable to overexploitation.     

Age and growth rate determinations for the Atlantic surf clam Spisula solidissima (Bivalvia: Mactracea), based on internal growth lines in shell cross-sections

Marked and recovered surf clams, Spisula solidissima Dillwyn, from Virginia (USA) deposited one internal growth line during a period of 11/2 years, probably in response to spawning during late summer. We have used these annual growth lines to make growth curves for two samples of New Jersey (USA) clams, one from inshore (1.8 km from shore, 15 m deep), the other from offshore (17.5 km from shore, 28 m deep) waters. The offshore and inshore clams grow at approximately the same rate until Age 3 years, after which time the growth rates differ, as does the ultimate lifespan; the offshore clams grow more rapidly and attain a greater age — up to 31 years. Comparison of our growth curves with other published curves revealed a close correspondence to a curve which was based on a study of growth over a 5-year period. Curves based on external growth lines probably underestimate growth rate in early life and overestimate it in later years.     

Growth of Flustra foliacea (Bryozoa)

The perennial bryozoan Flustra foliacea L. has annual growth-checks which leave lines across the fronds. These growth-checks have been used to determine the age and the pattern of growth of the colonies in terms of height and numbers of zooids. Monthly samples have been used to find the annual growth cycle. Heavy encrustations of epizoites on the F. foliacea colonies reduce growth rate. As the fronds increase in height, frontal budding of zooids thickens, and thereby strengthens, the holdfast.     

<Emphasis Type="Italic">Pinna nobilis</Emphasis> L., 1758 age determination by internal shell register

The age of the Mediterranean endemic Bivalve mollusc Pinna nobilis is estimated using the total size and the number and position of posterior adductor muscle scars (PAMS): a series of straight lines and rings observed in the dorsal nacre-lobe. Nevertheless, the position of PAMS of adult individuals inhabiting the same population and depth is highly variable. To assess the source of this variability, the shells of nine individuals were cut into radial sections along PAMS and the inner register was studied. The cuts showed clear marks of the inner register formed around the PAMS composed by a short nacre tongue introduced within the calcitic layer and two calcitic strips of differing colour (bright and dark). The three components are directed towards the posterior of the shell. The marks of the inner register coincide only with the straight lines of PAMS, but are observed in an anterior position further than the straight lines and rings, and in a posterior position further than the rings. Also, some of the most posterior marks can be observed one under the other in the oldest individuals. The existence of the inner register’s marks in a position further anterior than the straight lines and rings implies that a variable number of PAMS are obscured beneath nacreous deposits. Eventually, we advise against the utilization of PAMS to study P. nobilis age. We instead recommend the use of the inner register as it records more accurately the life history of the individuals, and data indicates that each mark is formed annually.     

Individual growth and somatic production in<Emphasis Type="Italic"> Adelomelon brasiliana</Emphasis> (Gastropoda; Volutidae) off Argentina

Growth, age and somatic production of the benthic predator Adelomelon brasiliana were studied at the southern limit of its distribution on the South American Atlantic shelf. Stable oxygen isotope ratios confirmed annual formation of internal shell growth marks. Modal shell length of the population was 140 mm, while modal shell-free wet mass was 255 g. A logistics growth function (SL=186.28 mm, K=0.185, t₀=4.601) fitted 131 pairs of size-at-age data (25 shells) best. A. brasiliana is a very long-lived species, reaching up to 20 years of age. The maximum individual somatic production of 46 g shell-free wet mass year^–1 is attained at 145 mm shell length, which corresponds to about 12 years of age.Communicated by O. Kinne, Oldendorf/Luhe     

Life history of the temperate subtidal gastropod Umbonium costatum

The life history and reproductive biology of the trochid sand snail Umbonium costatum (Kiener) were investigated on a subtidal sandy shore in Hakodate Bay, Japan between 1988 and 1991. Female U. costatum mature at 1 yr of age (shell diameter-11 mm), reproduce twice (in June-July and September-October) in successive years, grow to a maximum size (shell diameter=22 mm) at age 8 yr, increase annual fecundity with age from 2000 (age 1 yr) to 37000 (age 8 yr), and show a maximum monthly gonad somatic index of 8% which is constant among ages. In comparison to a previously studied life history of a tropical Umbonium vestiatium, temperate U. costatum shows more sustained growth and a longer life span after maturation. This could be explained by: (1) the optimal size model concerned with resource investment, in gametes (Sebens 1987); and by (2) bet hedging to compensate large variability in larval success at high latitudes. These two hypotheses are not mutually exclusive, but both are based on season-related extremes of environment at high latitudes where the period suitable for reproduction is short.     

The gastropod statolith: a tool for determining the age of<Emphasis Type="Italic"> Nassarius reticulatus</Emphasis>

The microstructure, shape and appearance of the growth rings in statoliths of Nassarius reticulatus (L.) were investigated. This species possesses two statocysts, each containing a single spherical statolith of calcium carbonate of up to 0.22 mm in diameter in the largest animals. The relationship between statolith diameter (SD) and total shell height (TSH) is exponential [ln(TSH)=26.3SD–0.842], although the function is site specific. Statoliths of the largest whelks (>29 mm) contained three or four clearly defined rings, corresponding to TSH values of ~1.1, 4.6–5.3, 12.0–13.5 and 18.5 mm, respectively. The first ring likely represents the metamorphic ring that was deposited at the time of larval metamorphosis when the post-larval whelk adopted a benthic lifestyle. The estimated size of the whelks at formation of the second, third and fourth statolith rings closely matched the TSH inferred from the shell rings. It is concluded that the patterns of growth rings present in statoliths can provide information about the age and growth of N. reticulatus.Communicated by O. Kinne, Oldendorf/Luhe     

Growth and size structure in a baltic Mytilus edulis population

Since Mytilus edulis L. has very few predators and competitors for space, it has become a biomass dominant in the Baltic proper covering hard substrates from the water surface to more than 30 m depth. In order to investigate the factors controlling size and production in a Baltic M. edulis population, growth was studied by the analysis of annual growth rings, measurements of caged individuals and the analysis of size classes in the population, and on settlement ropes. The total number of mussels in a representative mussel bed at 4 m depth varied between 36 000 and 158 000 ind · m^-2 during the year, mainly due to variations in very small mussels (<2 mm), whereas the abundance of mussels 2mm was rather constant between about 17 000 and 28 000 ind · m^-2. Maximum numbes of mussels < 2 mm, amounting to 132 000 ind · m^-2, were found after settlement in summer, but still half a year later in spring, 65 000 ind · m^-2 < 2 mm were registered, due to very strong intraspecific competition for food and space leading to the competitive suppression of small individuals and large variations in growth rates. Due to the special size-structure of the population only the analysis of annual growth rings could be used to estimate natural shell growth. From being very low in the smallest mussels, growth was linear between about 2–10 yr of age, corresponding to about 3–20 mm length, after which it decreased with a L=32 mm. Over the linear interval, growth in the populations from 3–6 m and 10–15m depth was 3.1 and 2.2 mm · yr^-1, respectively. Meat growth showed strong annual variations mainly due to gonad production. Starving mussels could, however, while utilizing energy reserves, survive losses of up to 78% of their meat biomass. This ability of M. edulis to respire away its own biomass and its apparent tolerance of weight loss has important implication. It will drastically reduce the energy flow to destruents from mussels dying naturally, which is of special significance in the Baltic, where predators and scavengers are scarce. It enables the mussels to endure bad food conditions and buffer strong seasonal variations in food abundance, maintaining the strongly food-and space-limited Baltic M. edulis population at the carrying capacity of the area.     

Resource allocation and population genetics of the bay scallop,Argopecten irradians irradians: effects of age and allozyme heterozygosity on reproductive output

The relationships among multiple-locus heterozygosity, age, reproduction and growth were examined over the ca. 2-yr lifespan of a cohort of northern bay scallops, Argopecten irradians irradians (Lamarck), from the Niantic River estuary, Connecticut, USA. Electrophoretic analyses revealed a relatively low proportion of polymorphic loci (=0.35) and low level of heterozygote deficiency ( D. across 6 loci=-0.05) in this population. Allele frequency distributions showed a high degree of temporal stability within and among cohorts. No significant correlation was found between multiple-locus heterozygosity and either somatic or reproductive growth. Our results lend support to the postulated linkage between low heterozygote deficit and lack of a heterozygosity/growth association in marine bivalves, and in the pectinid group in particular. Partial reproductive senility occurs in these scallops in their second year of life. Somatic tissue weight increased exponentially with increasing shell height, whereas pre-spawning gonad mass attained asymptotic values in 2-yr-old scallops. Thus, weight-specific reproductive effort was significantly lower in second- than first-year scallops (24 and 33% respectively). This pattern could be generated if high individual reproductive effort in the first year were correlated with accelerated (early) post-reproductive senescence. This kind of resource allocation, in which somatic growth is given precedence over reproductive growth in older individuals, has been previously reported in only one other bivalve, a pectinid (Chlamys islandica), in which the decline in reproductive effort occurs only after the scallops reach an age of ca. 20 yr.