Offshore marine cheilostome Bryozoa from Fort Lauderdale,Florida

Cheilostome Bryozoa were collected upon marine fouling test panels exposed offshore from Fort Lauderdale, Florida, USA between 1961 and 1967. Panels were exposed at certain depths, ranging from the surface to the bottom, in several arrays extending to a distance of 4 miles offshore. Twenty-three species were identified from the 114 panels recovered and examined. Most species were found on panels in shallow water, or on panels near the bottom in deeper, offshore water. A notable exception was a large, arborescent colony of Biflustra savartii which engulfed a panel 50 feet (15 m) deep, 4 miles from shore, in the Florida current. The lack of cheilostome Bryozoa on shallow and mid-water panels at the offshore, deepwater stations was apparently due to the low number of larvae reaching them from the distant inshore colonies.     

Mortality and growth of juvenile hard clams Mercenaria mercenaria during brown tide

Blooms of the picoplankton-sized alga Aureococcus anophagefferens Hargraves et Sieburth, also known as "brown tide," frequently recur in Great South Bay, New York, USA. A field study compared mortality and growth rates of juvenile hard clams, Mercenaria mercenaria (L.), raised at a site that experienced a bloom of A. anophagefferens (>1᎒⁶ cells ml^-1) to clams grown in a nonbloom site from May to September 2000 at two sites on the north and south shores of Long Island. High (67%) cumulative clam mortality was observed for clams raised at the brown tide site (south shore) during the bloom (June-July), and individuals exhibited no measurable growth during that period. After the bloom subsided (mid-July), clams that survived the brown tide suffered low (<5%) cumulative mortality over the remaining sampling period (July-September). These survivors exhibited rapid growth for 4 weeks shortly after the bloom's decline. Subsequently, their growth slowed to rates comparable with those observed at the nonbloom site (north shore). By contrast, a low (<5%) cumulative mortality was observed for M. mercenaria raised at the site with no brown tide, and positive growth occurred throughout the study. These data suggest that some juvenile clams not only survive brown tide but also recover following the bloom.     

Offshore distributional patterns of Hawaiian fish larvae

An analysis of ichthyoplankton samples based on relative abundance reveals pronounced inshore/offshore distributional gradients for most Hawaiian fish larvae. Larvae of pelagic bay species are found almost exclusively in semi-enclosed bays and estuaries. Larvae of pelagic neritic species are more or less uniformly distributed with distance from shore. The larvae of reef species with non-pelagic eggs are most abundant close to shore, while those of reef species with pelagic eggs are most abundant offshore. Finally, the larvae of offshore (primarily mesopelagic) species show no clear pattern but frequently occur in high numbers nearshore. Within any group, variation in pattern was often evident; for example, although Hawaiian fishes of both the families Labridae and Mullidae spawn pelagic eggs, larvae of the former had not peaked in abundance 12 km from shore while larvae of the latter had peaked between 0.5 and 2 km. Some larvae which occur offshore are highly specialized morphologically for a pelagic existence (e.g. Chaetodontidae, which is illustrated) while others are little modified (e.g. Labridae). These findings indicate ichthyoplankton surveys in tropical areas must sample offshore areas in addition to the inshore adult habitat to obtain a complete picture.Hawaii Institute of Marine Biology Contribution No. 484.     

Density-dependent growth in the Atlantic surfclam, Spisula solidissima, off the coast of the Delmarva Peninsula, USA

Spisula solidissima (Dillwyn, 1817) is a large, suspension-feeding bivalve, whose range extends from Nova Scotia to South Carolina. This species is harvested commercially. Shell length and age data were collected for this species from 1980 to 1994 during surveys of population size and structure. These data were used to examine the relationship between the growth rate of S.␣solidissima and intraspecific density. The null hypothesis was that density (represented by number of individuals per tow) would have no effect on rate of growth. A negative relationship would support the alternative hypothesis, that intraspecific competition had taken place. This analysis focused on the surfclam population offshore from the Delmarva Peninsula, USA because: (1) a major recruitment event occurred in 1977, (2) clam fishermen had reported “stunted” surfclams in that area, (3) a wide range of local densities were available to examine, and (4) the existence of a closed area within the study area set up an interesting contrast with areas left open to harvesting. Maps of surfclam abundance across the Delmarva region demonstrate that areas of highest density have generally remained in the same location through time. The results suggested that intraspecific competition has been important in structuring this population. Based on data from 1980 to 1992, shell length was significantly reduced at high density, and a significant interaction between age and density was observed. Growth modeling indicated decreased asymptotic lengths and growth rates with increasing density. In nine out of ten pairwise randomization tests, fitted von Bertalanffy growth curves, representing different densities, were significantly different from each other. High densities of clams have persisted in the area that was closed to harvesting for 11 years (1980 to 1991). In 1994, length at age was significantly less in this closed area compared to that in the surrounding area. This effect was apparent in clams from 3 to 17 years of age, and most pronounced in the cohort that recruited to the Delmarva region in high numbers in 1977. Lower growth rates within the closed area have management implications for the optimal duration of closures. Received: 16 May 1997 / Accepted: 7 October 1997     

Spatio-temporal population dynamics of the sand snail Umbonium costatum: importance of ontogenetic migration and annual recruitment variability

The spatio-temporal population dynamics of the subtidal snail Umbonium costatum (Kiener) in Hakodate Bay, northern Japan, are described over a 9-yr period (1982 to 1988, 1992). Annual variations in recruitment success not only caused the highly variable age structure of the population, but also affected its distribution pattern. In heavy recruitment years (1982, 1984 and 1988), location of the densest population differed from the other years, and the areas with 0 yr old individuals were larger. Every year, the distribution of 0 yr olds was mostly restricted to inshore (within 320 m from shore), while the distribution of adults (>0 yr) was mostly restricted to offshore (>320 m from shore). Such patterns may be generated by ontogenetic migration of cohorts to offshore areas. Ontogenetic migration also could have buffered the large annual variation in spat distribution and maintained the small variation in the distribution of adults. (Prince et al. 1988). Density-related migration has also been reported in Acmaea digitalis (Frank 1965), Patella cochlear (Branch 1974) and Patelloida alticostata (Black 1977).Dense populations of the gastropod Umbonium costatum are found in some clean subtidal sandy-bottoms in Japan, e.g. inner part of Hakodate Bay. In the bay, the age structure of U. costatum revealed predominant year-classes. The 0 yr olds were restricted to the shallowest zone (<4 m depth) and most adult (<0 yr) cohorts were restricted to 6 to 7 m depths (Noda 1991a).This paper describes the temporal variability of population structure and distributional patterns of Umbonium costatum, and examines the distribution-regulating processes.     

FADs influence on settlement and dispersal of the young-of-the-year greater amberjack (<Emphasis Type="Italic">Seriola dumerili</Emphasis>)

FADs are fish aggregating devices applied worldwide over the centuries to increase fish catch. However, the utilisation of such floating devices can influence dispersion patterns of some fish species. At present it is still not clear if FADs play a role in the shoreward migrations of juvenile fish of reef associated species, such as the greater amberjack Seriola dumerili. In this study, we hypothesised that FADs located along a distance gradient from the shore might lead the greater amberjack inshore. In such case, mean abundance of S. dumerili should decrease over time in offshore FADs and increase in inshore FADs. To test the hypothesis three FAD systems were positioned in the Gulf of Castellammare (Sicily, Italy), between July and September 2001, at increasing distances from the coast. During the study period, five visual censuses were carried out within FADs fields. During the first sampling period, no significant difference in abundance and size of juvenile S. dumerili was found between the three FAD systems. All the other sampling periods reported higher abundances and sizes in the offshore FADs than in the two inshore FADs. The findings suggest that FAD systems might be exerting two different effects on young of the year (YOY) S. dumerili distribution: (a) offshore FADs tend to retain associated fish for longer periods of time, (b) coastal FADs favour the transition of YOY S. dumerili from the pelagic to the benthic domain.     

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.     

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.     

Age structure,growth, and morphometric variations in the Atlantic surf clam,Spisula solidissima,from estuarine and inshore waters

The surf clam, Spisula solidissima (Dillwyn), population in the estuarine waters of Long Island Sound, New York, USA, was characterized in 1984 and again in 1988 by an age structure restricted to just two age-classes, and an apparent lifespan of only about 10 yr. In the inshore coastal waters off Fire Island, New York, a wide age range from 2 to 22 yr old was present. The age structure at Long Beach, New York, a third geographic region which is coastal but influenced by the Hudson River estuary, was similar to Long Island Sound. Juvenile surf clams grew at similar rates in all three geographic regions. However, adults from the Long Island Sound population grew significantly slower and reached an asymptotic maximum size which was 37% smaller than Fire Island adults. Long Beach adults had intermediate growth rates and maximum sizes. The shells of Long Island Sound clams were also 25% thinner than those from the other two regions. Density dependent effects on growth, evaluated over abundances ranging from 0.5 to 294 ind. m^–2, were present but were too small to account for observed regional differences. Results suggest that adult surf clams may be physiologically stressed by the reduced salinity and more extreme temperatures found in estuarine waters.     

Biology of the conger eel Conger oceanicus in the Mid-Atlantic Bight

In the Mid-Atlantic Bight, conger eels (Conger oceanicus) occur from the coastal portions of estuaries to the edge of the continental shelf. In deeper waters they occupy burrows of the tilefish (Lopholatilus chamaeleonticeps). Between 1972 and 1974 we examined the stomachs and intestines of conger eels from inshore New Jersey (USA) waters (n=35, with a total length: TL of 21 to 49 cm) and between 1980 and 1983 offshore (n=295, 50 to 125 cm TL)_collections. Eels from both areas fed primarily on decapod crustaceans and fish. The specific identity of prey items within these groups generally differed from inshore to offshore areas, probably reflecting the differences in prey availability. Foods of specimens collected offshore varied with size: smaller eels (<80 cm TL) fed most heavily on decapod crustaceans, whereas larger eels (>80 cm) consumed more fishes. The presence of some nocturnally active prey items in the gut, primarily the eel Lepophidium cervinum, suggests that conger eels are nocturnal feeders. This is supported by in situ observations that conger eels are present in some tilefish burrows during the day and are presumably out of burrows and foraging at night.     

Contrasting foraging tactics by northern gannets (<Emphasis Type="Italic">Sula bassana</Emphasis>) breeding in different oceanographic domains with different prey fields

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.     

Growth of the deep-sea irregular sea urchins Echinosigra phiale and Hemiaster expergitus in the Rockall Trough (N.E. Atlantic Ocean)

Growth in the deep-sea irregular sea urchins Echinosigra phiale (family Pourtalesiidae) and Hemiaster expergitus (family Hemiasteridae) was studied from deep-sea samples taken during the years 1973 to 1985 from two stations at 2900 and 2200 m depth in the Rockall Trough (N.E. Atlantic Ocean). Growth zones, similar to those described from sea urchins in shallow water, are present as a series of wide white bands separated by narrow, dark rings in the calcite stereom of the test plates after heating to 350°C. In shallow water, such growth zones seem to result from seasonally varying growth rates. In the supposedly constant conditions in the deep sea, a seasonal growth pattern is unexpected but may occur in response to recently discovered annual pulses in downward flux of detritus from the euphotic zone, providing a seasonally varying food supply for such deposit-feeding species living in the bottom sediment. On this assumption, growth curves were fitted to counts of growth zones (as representing age in years), in the larger lateral and ventral test plates of E. phiale and H. expergitus. The opportunity was also taken to fit growth curves derived from counts of growth zones in samples of the inshore spatangoids Spatangus purpureus and Echinocardium pennatifidum. Plots of counts against test length of Echinosigra phiale and H. expergitus, although scattered and not clearly asymptotic, indicate, growth to be slower than in the two inshore spatangoids, and than in the coastal species Echinocardium cordatum, for which there are good recent growth data, available.     

Growth and utilization of internal nitrogen reserves by the giant kelp Macrocystis pyrifera in a low-nitrogen environment

An adult giant kelp plant (Macrocystis pyrifera), moved from an inshore kelp forest to an offshore, low-nitrogen environment near Santa Catalina Island, California (USA), maintained growth for 2 wk on internal nitrogen reserves. Frond elongation rates decreased significantly during the third week, and plant growth rate (wet wt) dropped from an initial inshore rate of 3.6 to 0.9% d^-1. During this 3 wk period, nitrogen contents and free amino acid concentrations decreased, while mannitol and dry contents increased in frond tissues. After depletion of internal nitrogen reserves, the nitrogen content of lamina and stipe tissues averaged 1.1 and 0.7% dry wt, respectively. The experimental plant was exposed to higher ambient nitrogen concentrations during the last 2 wk. Rates of frond elongation and plant growth increased, but nitrogen content and amino acids in frond tissues remained low. Of the total nitrogen contained in frond tissue of the plant before transplantation, 58% was used to support growth in the absence of significant external nitrogen supply. Amino acids constituted a small proportion of these internal nitrogen reserves. Net movement of nitrogen occurred within large fronds, but not between different frond size classes. The nitrogen content of holdfast tissue remained relatively constant at 2.4% dry wt and accounted for 18 to 29% of the total nitrogen. Holdfast nitrogen was not used to support growth of nitrogen-depleted fronds. In comparison to Laminaria longicruris, which is adapted to long seasonal periods of low nitrogen availability, M. pyrifera has small nitrogen-storage capacity. However, internal reserves of M. pyrifera appear adequate to make nitrogen starvation uncommon in southern California kelp forests.     

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.     

Genotype-specific growth of hard clams (genus Mercenaria) in a hybrid zone: variation among habitats

Shell growth rate is an important component of fitness in bivalve molluscs. Using the parameter computed from the von Bertalanffy growth equation, we quantitatively compared rates of annual shell grwoth among the hard clams Mercenaria mercenaria, M. campechiensis, and their hybrids sampled from a variety of habitats in the Indian River lagoon, Florida, USA, a zone of species overlap and natural hybridization. Our results indicate that the classical paradigm describing hard clam growth, in which growth rate is fastest in M. campechiensis, intermediate in hybrids, and slowest in M. mercenaria is not supported in the Indian River lagoon. Instead, M. campechiensis has a growth advantage in deep-water habitats in the northern section of our study area. In the central and southern sections of our study area, hybrids have a growth advantage over M. mercenaria in shallow-water habitats, but M. mercenaria has a growth advantage over hybrids in deep-water habitats. In all other sampled habitats, either growth rate among genotype classes is equal, or M. mercenaria has a growth advantage. This complex relationship between genotype and habitat-specific growth provides a mechanism for selection to act on hard clams in the Indian River.     

Nearshore distributional gradients of larval fish (15 taxa) and planktonic crustaceans (6 Taxa) in Hawaii

Fifteen taxa of fish larvae (most identified to species) and 6 taxa of crustaceans (most identified to species) were studied in zooplankton samples collected 0.2, 0.5, 1.0 and 3.0 km off the leeward coast of Oahu, Hawaii in May and July 1975. The following distributional patterns were elucidated: inshore (2 spp); inshore-neritic (2 spp); neritic (7 spp); and offshore-neritic (7 spp). Three of these 18 species had age-related differences in behaviour that led to age-dependent distributional patterns, and 4 further species had a random distribution. All abundant types of fish larvae were either oceanic as adults or were hatches from the demersal or brooded eggs of reef species. The larvae of reef fishes with pelagic eggs were not abundant in the nearshore region sampled and were probably to be found more than 3 km offshore. The offshore-neritic distributional pattern evidently resulted from relatively passive movement with currents. In contrast, maintenance of inshore and neritic patterns probably required relatively active swimming by the animals. The current regime, including a tidal eddy and possible nearshore upwelling, probably helped maintain the inshore and neritic patterns of such animals. The limited area occupied by these inshore coastal plankton communities could make them particularly vulnerable to environmental changes, including anthrogenic ones.     

Influences of habitat and natural disturbances on contributions of massive Porites corals to reef communities

We compared densities, distributions and size frequencies of massive corals in the genus Porites on five relatively exposed, mid-shelf reefs (50 km offshore) in the central Great Barrier Reef with those on a sheltered inshore reef (10 km offshore). Data included various transect and mapping studies between 1984 and 1990, estimates of size-dependent damage from the crown-of-thorns starfish Acanthaster planci, estimated densities of herbivorous sea urchins (potential predators of juveniles), and observations of size-specific effects of tropical cyclones. Assemblages of Porites spp. on mid-shelf reefs were dominated by small colonies (2 to 10 cm diam) established either from planula larvae or from small tissue remnants that had survived A. planci predation in the early to mid-1980s. Large colonies (up to 10 m diam) were scarce, except for localized aggregations on terraces at the base of reef slopes (6 to 12 m deep). Extensive space suitable for settlement by coral larvae can be attributed to recurrent cyclones and A. planci outbreaks. Despite low sea urchin predation, the slowly growing Porites juveniles are likely to die from overgrowth by numerous, much faster growing corals. On the sheltered inshore reef, the coral community was dominated by very large (>5 m diam) Porites colonies, several centuries old; recruitment was mainly by fragmentation of large colonies; there was little space available for settlement, and probabilities of juvenile mortality from grazing urchins were high. Differences in settlement and early survival of Porites spp. are exacerbated by different regimes of storm damage. A model is proposed that links wave climate with the size and age reached by corals before dislodgement by storm waves, and which is consistent with observed densities and size-frequency distributions of Porites in sheltered and exposed areas.     

Environmental contamination using accumulation of metals in marine sponge,Sigmadocia fibulata inhabiting the coastal waters of Gulf of Mannar,India

Coastal marine ecosystems in many parts of the world are under unrelenting stress caused by urban development, pollutants and other ecological impacts such as building of infrastructure, land reclamation for port and industrial development, habitat modification, tourism and recreational activities. The present work is a first extensive field study using the marine sponge, Sigmadocia fibulata as a bioindicator to detect metal pollution between inshore and offshore environment of the ‘Gulf of Mannar’, India. Samples of S. fibulata were collected from different places of inshore (0.1–0.5?km) and offshore (3–5?kms) locations. Metal concentrations in water and bioaccumulation in sponges were determined by ICP-MS (Inductively coupled plasma-mass spectrometry). Enrichment of metal contamination was more in the inshore, when compared to offshore environment. Higher concentrations of metals Fe, Al, Ni, Mn, Cu, Cr, Co, Ba, Zn, V, Pb and Cd in inshore waters (3–6.4 times than offshore) may be due to the discharge of monsoonal rains, carrying a discharge of industrial and agricultural wastes and sewage directly into the coast. This is substantiated by a highly significant positive correlation between concentrations of metals in the water and accumulation in sponges. The bioaccumulation of metals in sponge tissue were in order of Fe?>?Al?>?Ni?>?Mn?>?Cu?> Cr?>?Co?>?Ba?>?Zn?>?V?>?Cd?>?Pb in both the inshore and offshore regions. Apparently, the macromolecule composition (sugars, proteins and lipids) was significantly reduced by the accumulation of metals in inshore sponges. The current findings indicated that the frequent monitoring is necessary to assess the eco-health of the marine environment by choosing bio-indicator species like S. fibulata, which provide accurate, reliable measurements of environmental quality.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. XI. Regeneration of ammonium in the surf environment by the Pacific razor clam Siliqua patula

A 20 month field study was conducted on ammonium excretion rates of the Pacific razor clam Siliqua patula Dixon along the beaches of Washington State, USA. Excretion rates of all those nutrients likely to be regenerated in sufficient quantity to affect surf diatom growth were measured; ammonium appeared to be the most important metabolite. Excretion of ammonium by razor clams far exceeded that by other beach fauna. Ammonium excretion rates of razor clams were positively correlated with shell length, but no correlation between ammonium excretion rate and water temperature was evident. This may be an artifact or may represent some degree of seasonal acclimation of the species to temperature. Weight-specific ammonium excretion rates were negatively related to clam size, indicating a possible large (and unknown) contribution of regenerated ammonium by smaller clams in their first year of growth; smaller clams were rarely captured during this study.Contribution No. 1048 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Assessment of intertidal growth and capacity adaptations in suspension-feeding bivalves

Curves relating instantaneous growth rate to aerial exposure were determined for six species of bivalves in laboratory and shore experiments. A dimensionless index equatable with the relative intertidal growth performance of a species was calculated by integration of the growth curve after converting both growth rate and aerial exposure to decimal fractions. Intertidal growth performance of the bivalves tested corresponded well with their natural levels of occurrence on the shore, and improved in the following order: Modiolus modiolus < Argopecten irradians < Ostrea edulis < Mytilus edulis < Crassostrea virginica. Geukensia demissa, for which an index value could not be determined, grew faster intertidally than subtidally. The relative contributions made to intertidal growth performance by energy-conserving and energy-supplementing capacity adaptations were assessed by resolving the growth curves into energy-input and energy-loss components. The rate of energy loss due to intertidal exposure was lower in the high-shore species, and also less affected by harsher subaerial conditions, than in the low-shore species. Moreover, M. edulis and C. virginica showed abilities to supplement energy input such that growth per unit immersion time was better at certain intertidal levels than subtidally. Energy conservation and supplementation in these forms made roughly equal contributions to their improved intertidal growth relative to species occurring lower on the shore.