Age determination of the soft-shell clam Mya arenaria using shell internal growth lines

Shells of known-age Mya arenaria have revealed an internal annual growth pattern in this bivalve. The number of external growth rings on the shell exterior does not correspond to the age of the clams. This study provides an alternative method to using external annual rings for age determination. The internal growth lines are visible in thin sections when the shells are cut from the umbo to the ventral margin. The lines are formed by late spring of each year prior to spawning and are probably in response to decreased winter growth. When the 7 year olds were examined it was difficult to distinguish the most recent growth line and the inner shell layer. This could result in ages being underestimated. This annual pattern of internal growth lines could be useful to biologists and paleoecologists engaged in population studies.     

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 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     

Age and growth of the lanternfish Lampanyctodes hectoris (Myctophidae) from eastern Tasmania,Australia

The age, growth and mortality of the lanternfish Lampanyctodes hectoris (Gunther) were investigated by examining the otoliths, length-frequency data, and seasonal abundance of fish collected from continental slope waters of eastern Tasmania between April 1984 and June 1985. Although L. hectoris can live to 3 yr old, it has a high annual mortality (79%) and few live past their first year. Growth, described here by a variation of the Von Bertalanffy model, takes place mainly in the first six months of life and only during late spring and summer. The hypothesis of daily deposition of primary growth increments in myctophids was supported by comparing the back-calculated birth dates of daily-aged individuals of L. hectoris with the spawning season of this species.     

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.     

Sclerochronological records of temperature and growth from shells of Mercenaria mercenaria from Narragansett Bay,Rhode Island

Annual internal growth increments in shells of hard clams (Mercenaria mercenaria) provide an accurate record of both growth history and some types of environmental change. These increments were used to determine age and growth rate of hard clams collected in 1984–1985 from ten sites in Narragansett Bay, Rhode Island, USA, and to assess geographic variation in growth within the bay. The regional comparisons were facilitated by modeling the clam growth using the von Bertalanffy equation and the parameter of Gallucci and Quinn. The optimum region for hard clam growth was near the head of the bay, with areas further north (Providence River) and south (lower bay) not as productive. The relationship between size and age was similar to that reported for hard clams from northern New Jersey. Using dendrochronological techniques, variations in growth were analyzed on a temporal (year-to-year) basis. The comparative longevity of the hard clams (individuals with 40 annual bands were encountered) demonstrated that sclerochronologies of several decades were possible. A 26 yr growth record based upon 100 individuals was established, covering the years 1959–1984. Significant temporal variations occurred in bivalve growth, and a broad trend of increasing growth indices over the last two decades of the record was noted. The yearly standardized growth index values were highly and positively correlated with mean annual water temperatures in Narragansett Bay for the same time interval. The incremental shell records of these bivalves provide a quantitative indication of marine climatic (temperature) variability and growth which can be used in hindcasting the effects of natural or anthropogenic environmental perturbations.     

Skeletal growth zones as age-markers in the sea urchinPsammechinus miliaris

Growth of the inshore sea urchinPsammechinus miliaris (Gmelin) was studied in Loch Creran, western Scotland, using the skeletal growth-marker tetracycline in order to test the validity of natural growth-banding in the coronal test plates as annual age-markers. In order to test whether tetracycline affected the growth ofP. miliaris, an injected and a control group of urchins were held in identical conditions in running sea water aquaria for 21 mo from 1989 to 1991 and measured periodically. A small but significant difference in mean size of injected compared to controls was recorded at 12 mo, but none during subsequent measurements. Size measurements during the trial were consistent with an annual growth cycle, with a maximum in spring and slowing or cessation of growth during autumn/winter. Tetracycline-labelled juveniles were recovered up to 18 mo after initial tagging in mark/recapture experiments undertaken from 1987 to 1989 at two intertidal marked quadrats in Loch Creran. Large numbers ofP. miliaris were also marked with tetracycline and held for 1 yr at 10 m depth in seabed cages in Loch Creran during 1988–1990. All of the intertidal recoveries, and about 69% of the caged specimens that had been successfully labelled, showed a consistent relationship between the position of the tetracycline tag and the pattern of natural growth zones. The remainder were mostly large, slow-growing urchins with the tag positioned near the plate margin. In these the outer growth bands were closely spaced and, particularly if major growth bands seemed to be broken up into double or multiple lines, the major bands were impossible to resolve at the margin. The results support the assumption that in wild populations the dark bands visible with reflected light (translucent in transmitted light) after charring the plate are formed when skeletal growth has stopped or slowed in winter. These lie between wider, lighter coloured (opaque in transmitted light) zones of active plate growth in spring/summer. The dark band formed beyond the tag usually was made up of several closely spaced fine lines, or sometimes of two closely spaced dark bands. The wide growth zones beyond the tag, like those formed previously, usually were broken by fine, dark lines that may represent brief discontinuities in growth. From tagging, the double dark bands can be related to growth over one year; but such anomalous bands, along with the general presence of fine, dark lines interrupting the growth zones, make it difficult reliably to estimate age from the closely spaced peripheral banding on older, slow-growing urchins.     

Interannual variability in the spring phytoplankton bloom in Auke Bay,Alaska

Data on phytoplankton primary production, biomass, and species composition were collected during a 5 yr (1985–1989) study of Auke Bay, Alaska. The data were used to examine the interannual differences in the timing, duration, and magnitude of the spring phytoplankton blooms during each year and to relate these differences to interannual variations in weather patterns. Within any given year, a pre-bloom phase was characterized by low available light, low rates of primary production, low biomass, and predominantly small (<10µm) diatoms. During the primary bloom, integrated production rates rose to 4 to 4.5 g C m^–2 d^–1, and integrated biomass levels reached 415 to 972 mg chlorophyll m^–2. Primary blooms were usually dominated by large diatoms (Thalassiosira spp.), and in a single year (1989) byChaetoceros spp. The primary blooms terminated upon nutrient depletion in the euphotic zone. Secondary blooms, triggered by nutrient resupply from below, occurred sporadically after the primary bloom and accounted for 4 to 31% of total spring production. The date of initiation and the duration of the primary bloom varied little from year to year (standard deviation 3 and 5 d, respectively). Seasonal production rates and biomass levels varied interannually by a factor of 2 to 3. In contrast, intra-annual variations of more than an order of magnitude, especially in biomass, occurred over periods as short as 10 d. These large variations over short time periods indicate the importance of synchronous timing between spring blooms and the production of larval fish and shellfish, which depend on an appropriate and adequate food supply for growth and survival. Parameters describing primary production (e.g. peak daily production, mean daily production, and total production during the primary bloom and the entire season) exhibited little interannual variation (coefficient of variation, CV = 10 to 19%), but a large degree of intra-annual variation (CV = 77 to 116%). Similarly, interannual variations in biomass (peak chlorophyll, mean chlorophyll) were also lower (CV = 20 to 33%) than intra-annual variations (CV = 85 to 120%).     

Rate of growth and longevity of a large colony of Pentapora foliacea (Bryozoa) recorded in their oxygen isotope profiles

We studied the growth patterns and the stable oxygen isotopic composition of an individual of Pentapora foliacea (Ellis and Solander 1786) collected on October 6, 1978 from the Bristol Channel, Pembrokeshire, UK, Irish Sea. The stable oxygen isotopes are in equilibrium with the ambient seawater and show a marked seasonal variation reflecting seasonal water temperature changes. The stable oxygen isotopes further suggest that regular growth patterns of less calcified growth bands, which are secreted in winter, are perennial. These winter growth check lines can easily be used to determine longevity of these bryozoa colonies and of annual growth rates. The colony analysed is at least 3-years-old with an annual growth rate of approximately 2 cm per year.     

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.     

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.     

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.     

Distribution,growth and life cycle of the mesopelagic amphipod Primno abyssalis (Hyperiidea: Phrosinidae) in the southern Japan sea

The vertical distribution, growth, maturation, brood size and life cycle of the hyperiid amphipod Primno abyssalis (formerly P. macropa) were investigated using seasonal samples collected from Toyama Bay, southern Japan Sea, during the period June 1986 to September 1992. Over four different seasons of the year, P. abyssalis was most abundant in the 200 to 350 m strata at night and the 350 to 400 m strata during the day, indicating 100 to 150 m as the general distance of diel vertical movement. Some differences in vertical migrating behavior were noted among juveniles, adult males and females. Population-structure analysis revealed the occurrence of three cohorts aged 0+, 1+ and 2+ yr. Growth as body length in this species is linear with time. Estimated time to complete one life cycle is 1.8 to 2.5 yr for females, but only 0.8 yr for males. Maximum longevity is 2.8 yr. Instar analysis based on the segment number of pleopod rami indicated that newly hatched juveniles molt ten times to reach adult male, and four more times to reach adult female. Adult instar number was found to be only 1 for males and 5 for females. Ovigerous females occurred throughout the year, but the annual peak of release of juveniles from the female's marsupium is estimated to be arly March. Brood size was not correlated with female size, a maximum brood size of 214 eggs was recorded. The dry and ash-free dry weights of instars suggested that juveniles in the female marsupium, adult males, and older adult females are less active feeding or non-feeding stages. Except for the reduced growth rate and the occurrence of small, short-lived males, most characteristics of P. abyssalis are consistent with the present view of the life modes of mesopelagic animals, including linear growth in length, aseasonal reproduction, and smaller brood size coupled with larger eggs.     

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.     

Age,growth, and mortality of the prosobranch<Emphasis Type="Italic"> Zidona dufresnei</Emphasis> (Donovan, 1823) in the Mar del Plata area,south-western Atlantic Ocean

The volutid snail Zidona dufresnei is a benthic top predator in the Mar del Plata (Argentina) shelf area where it was subjected to unregulated commercial exploitation for more than 20 years. So far there is no stock management, and hitherto even the most basic information on population dynamics of this species is missing. Annual formation of internal shell growth bands visible by x-ray was confirmed by the stable oxygen isotope record in the shell carbonate that reflects seasonal oscillations in water temperature. A Gompertz growth function ( , K=0.211 year^–1, t₀=5.496) fitted 142 pairs of size-at-age data (30 shells) best. Maximum individual production amounted to 26.8 g shell-free wet mass (SFWM) at 115 mm shell length. Based on a size-frequency distribution derived from commercial catches, annual mortality rate of Z. dufresnei was estimated to be 0.61 (±0.21) year^–1.Communicated by O. Kinne, Oldendorf/Luhe     

Growth and mortality of subtidal red sea urchins (Strongylocentrotus franciscanus) at San Nicolas Island,California, USA: problems with models

Red sea urchins, Strongylocentrotus franciscanus, were tagged with tetracycline in 1990 at subtidal sites off San Nicolas Island, California, USA. After one year in the field, the sea urchins were collected and growth increments were measured based on tetracycline marks, which indicated initial slow growth, a maximum rate, and finally a prolonged period of very slow growth. Small red sea urchins (4 cm diam) were estimated to be 3 to 4 yr old, which is much older than has previously been reported. It is estimated that about 12 yr would be required to attain 10 cm diam. Survival has previously been modeled assuming a constant rate. If the population of red sea urchins is assumed to be stable and stationary, annual survival rate was between 71 and 77% yr^-1. Census data for the two years of study have permitted annual survival to be estimated without assuming stable and stationary population structure. Under these conditions, annual survival rate was between 79 and 86% yr^-1. Analysis of transitions in the size distributions from 1990 to 1991 suggested that annual survival may have been sizespecific: 91% yr^-1 for individuals 1.1 to 4.0 cm diam, 82% yr^-1 for individuals 4.1 to 7.0 cm diam, and 63% yr^-1 for those of 7.1 to 10.0 cm diam. An alternative explanation to size-specific survival in our study is sizespecific immigration.     

Long-term changes in populations of subtidal bivalves (Abra alba and A. prismatica) from the Bay of Morlaix (Western English Channel)

Two populations of Abra alba (Wood) and one of A. prismatica (Montagu) (Mollusca: Bivalvia) were studied over a 10 yr period (1977–1987) in two muddy fine-sand subtidal communities of the Bay of Morlaix, France. The survey provided an example of long-term changes in the three Abra spp. populations, which displayed synchronized changes, with a regular annual cycle and increasing densities during 1979–1980 related to the higher concentration of organic matter resulting from the Amoco Cadiz oil spill in March 1978. A. alba rapidly adapts its demographic strategy to eutrophic conditions by increasing its reproductive potential, growth, and abundance. During times of eutrophication, A. alba has three spawning periods and three recruitments per year as opposed to two spawning periods and two recruitments per year during oligotrophic conditions. Growth of the juveniles of this species is insignificant until spring for individuals recruited in the autumn, whereas individuals which settle during spring or summer display immediate rapid growth. A. prismatica has a low capacity to adapt to eutrophic conditions. It has one annual period of sexual maturation at the end of the summer, with spawning in September–October and settlement beginning in mid-November. Growth of the juveniles after settlement is also insignificant until April. These results enable comparison of the demographic strategies of these two sympatric species.     

Annual cycles of phytoplankton community-structure and bloom dynamics in the Neuse River Estuary, North Carolina

The spatiotemporal distributions of major phytoplankton taxa were quantified to estimate the relative contribution of different microalgal groups to biomass and bloom dynamics in the eutrophic Neuse River Estuary, North Carolina, USA. Biweekly water samples and ambient physical and chemical data were examined at sites along a salinity gradient from January 1994 through December 1996. Chemosystematic photopigments (chlorophylls and carotenoids) were identified and quantified using high-performance liquid chromatography (HPLC). A recently-developed factor-analysis procedure (CHEMTAX) was used to partition the algal group-specific chlorophyll a (chl a) concentrations based on photopigment concentrations. Results were spatially and temporally integrated to determine the ecosystem-level dynamics of phytoplankton community-constituents. Seasonal patterns of phytoplankton community-composition changes were observed over the 3 yr. Dinoflagellates reached maximum abundance in the late winter to early spring (January to March), followed by a spring diatom bloom (May to July). Cyanobacteria were more prevalent during summer months and made a large contribution to phytoplankton biomass, possibly in response to nutrient-enriched freshwater discharge. Cryptomonad blooms were not associated with a particular season, and varied from year to year. Chlorophyte abundance was low, but occasional blooms occurred during spring and summer. Over the 3 yr period, the total contribution of each algal group, in terms of chl a, was evenly balanced, with each contributing nearly 20% of the total chl a. Cryptomonad, chlorophyte, and cyanobacterial dynamics did not exhibit regular seasonal bloom patterns. High dissolved inorganic-nitrogen loading during the summer months promoted major blooms of cryptomonads, chlorophytes, and cyanobacteria. Received: 12 September 1997 / Accepted: 12 December 1997     

The effect of water movement,temperature and salinity on abundance and reproductive patterns of<Emphasis Type="Italic"> Macrocystis</Emphasis> spp. (Phaeophyta) at different latitudes in Chile

This study describes the density variation and phenology of Macrocystis integrifolia and M. pyrifera populations from northern and southern Chile, respectively. Samples of both species were taken in wave-exposed and wave-protected areas. In addition, spore production, germination and early growth rate of sporophytes of each population was studied at monthly intervals under three temperature and salinity regimes. Results indicate that M. integrifolia from northern Chile presents perennial plants with a mean density of three individuals per 0.25 m² throughout the year and that it reproduces mainly during spring and winter. Although, M. pyrifera in exposed areas of southern Chile also have a perennial-type life strategy, they are able to reproduce all year round. In contrast, M. pyrifera populations in protected areas of southern Chile show a clear annual cycle, with high recruitment during late winter and fertile sporophytes in summer and autumn, although the populations become completely decimated thereafter. The effect of temperature and salinity on M. integrifolia shows that it is independent of water movement, but requires low temperatures and high salinities for the release of zoospores, germination and early sporophyte growth. This pattern differs from that of M. pyrifera in southern Chile, which has a broader tolerance range for salinity and temperature than does M. integrifolia. However, in southern Chile wave-protected populations showed higher spore release and germination at 15°C and 18°C, whereas sporophyte growth responded better at the lowest temperature tested (8°C). In general, these results are contrary to those expected, since a seasonal reproductive pattern was observed in M. integrifolia inhabiting a less seasonally variable environment. In exposed sites of southern Chile, plants showed greater tolerance and continuous reproduction throughout the year, despite the greater environmental variability. Finally, population dynamics of protected kelps in southern Chile shows an annual pattern, which is contrary to the expected perennial strategy shown by exposed populations.Communicated by P.W. Sammarco, Chauvin     

Genetic variation and covariation during larval and juvenile growth in Mercenaria mercenaria

Quantitative genetic variances and covariances were estimated for shell length of the hard clam Mercenaria mercenaria (L.) at three larval stages (prodissoconch I, 2 d and 10 d post-fertilization) in 1987 and in 1988 after ca. 9 mo of growth. At each sample interval additive genetic variance was a highly significant component of the total size variation. Narrow sense heritability estimates for shell length ranged between 0.58 (±0.10) for prodissoconch I and 1.08 (±0.29) for 2-d-old larvae. There was significant and positive genetic covariance in prodissoconch I and 2-d larval shell length which resulted in a highly significant genetic correlation (r _g=0.74) between these two traits. This covariance is not surprising since the prodissoconch I comprises the majority of the larval shell of a 2-d-old larvae. The genetic covariances between 2-d-old and 10-d-old larvae and between 10-d-old larvae and 9-mo-old juveniles were low and not significantly different from zero. These results indicate that there is substantial genetic variation for shell growth in M. mercenaria but this variation is not stable during development; the genetic variation in shell growth at one stage of development is not strongly related to the genetic variation in growth during other ontogenetic periods. In this study there were no evident constraints to natural selection for increased shell growth rate during development, which coupled with the high levels of genetic variation may suggest that in nature high rates of larval growth may not be normally subject to significant selective pressure.