The effects of sand movement on intertidal seaweeds and selected invertebrates at Bound Rock,New Hampshire,USA

A seasonal study of sand movement and the benthic intertidal organisms at Bound Rock, New Hampshire, USA was conducted between November, 1973 and February, 1975. The site is subjected to irregular sand fluctuations, as well as diurnal, neapspring and major summer sand intrusions. The abundance and distribution of intertidal species was interrelated with the historical sand fluctuations at the area. For example, the lower limits of Mytilus edulis, Balanus balanoides and Porphyra umbilicalis approximated the highest summer sand elevations. Highly abraded rock surfaces in the lower intertidal zone were dominated by opportunistic annuals (e.g. Enteromorpha spp.) and perennial psammophytic or sand-loving seaweeds (e.g. Ahnfeltia plicata and Sphacelaria radicans). Overall, the intertidal seaweed populations at Bound Rock showed a lower number of perennials and fewer species than adjacent rocky shores. The low species diversity of seaweeds at the study site is attributable to unstable environmental conditions and a limited number of habitats. Even so, several psammophytic microhabitats are evident at Bound Rock depending upon the extent of sand burial and abrasion. The morphological and reproductive adaptations of several psammophytic species are discussed.Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 812.Scientific Contribution No. 53 of the Jackson Estuarine Laboratory.     

Importance of biotic and abiotic regulators of abundance of the intertidal amphipod Calliopius laeviusculus (Amphipoda: Gammaridae) and assessment of the accuracy and precision of sampling methods for detecting abundance changes

We examined seasonal patterns of abundance for the intertidal amphipod Calliopius laeviusculus (Amphipoda: Gammaridae). Amphipods were sampled with an epibenthic sled during the daytime high tide period from 18 May to 8 August 1988. Amphipod density increased from May to June and reached maxima in both late June and early August. Amphipod density was unrelated to any abiotic component measured in the intertidal community. These variables included sampling location, wave height, water column height, water temperature, salinity and cloud cover. Changes in abundance were related with sampling date and with the onset of capelin (Mallotus villosus) spawning activity in the intertidal. Capelin eggs are an important food item for amphipods. The accuracy of density estimates obtained with the epibenthic sled was assessed through comparison with densities obtained with a more efficient quadrat sampler. Sled samples consistently sampled ca. 1% of the amphipod population. We found that a large portion of the amphipod population burrowed into the sediment and was not effectively sampled by the sled. Sled sampler precision was roughly equivalent to that of quadrats with D (precision) ranging from 0.26 to 0.42 for sled samples and D=0.29 for quadrat samples. Although sample collection with the epibenthic sled was achieved more quickly and under a wider range of weather conditions than was possible with the quadrat sampler, the serious underestimate of amphipod density based on sled samples alone indicates that both sled and quadrat samples, obtained in concert, are required to obtain accurate measures of daily variation in C. laeviusculus abundance.Contribution to the program of GIROQ (Groupe interuniversitaire de recherches oceanographique du Quebec)     

Influence of seasonal food abundance and quality on the feeding habits of an opportunistic feeder,the intertidal crab <Emphasis Type="Italic">Pachygrapsus marmoratus</Emphasis>

The abundance of trophic sources on the intertidal zone is discontinuous and their supply can vary both in a predictable or unpredictable way. The Mediterranean semi-terrestrial crab Pachygrapsus marmoratus, is known, as adult, to entirely rely on the intertidal trophic sources, and, consequently, it faces the fluctuations of nutritional sources and quality. To clarify the relationships between the feeding habits of an Italian population of P. marmoratus and the temporal variation of its food sources, we carried out a 2-year sampling protocol. Data on seasonal variation in composition of intertidal food item assemblages, on the average content in N and C of the commonest algae, on seasonal changes in crabs feeding habits were collected and compared using a suite of multivariate and univariate techniques. Results showed that P. marmoratus takes advantage of the recruitment phase of the most common invertebrates, affecting and controlling the abundance even of those species whose adults are out of its reach. It can act both as a herbivore, and as a carnivore that shifts between bivalves, more abundant in spring/summer, and the periwinkle Melaraphe neritoides, during winter. In conclusion, we provide evidences on the food choice of this common rocky shore species, which is very plastic and capable of relying on many trophic sources, possibly influencing the abundance and/or the population structure of a number of intertidal populations.     

Importance of predation by infaunal polychaetes in controlling the structure of a soft-bottom community in Maine,USA

Field manipulations in a Maine, USA intertidal mud flat showed that Nereis virens adults were an important factor in regulating the abundance of Corophium volutator, an amphipod which comprised 63% of the total number of individuals in the benthic community. Removal of N. virens adults resulted in an increase in C. volutator abundance, while addition of N. virens adults led to a decrease. There was a significant negative correlation between the abundance of N. virens adults and that of C. volutator. C. volutator was also negatively correlated with the density of all the other infaunal species combined. These results suggest that there is at least a 3-level interactive system (N. virens —C. volutator — other infaunal species) within the infauna in Maine; N. virens is a factor in controlling the abundance of C. volutator, and C. volutator may play a role in regulating the densities of other infaunal species. Models of generalized cropping in soft-bottom systems may be too simple if they ignore complex trophic relationships within the infauna.     

Oxygen-binding by haemocyanins from an ecological series of amphipod crustaceans

The oxygen-binding properties of haemocyanins (Hc) from three species of gammaridean amphipods, Gammarus locusta (L.) (subtidal), Echinogammarus pirloti (Sexton and Spooner), (intertidal, marine) and E. marinus (Leach) (intertidal, estuarine), one species of hyalid amphipod Hyale nilsonni Rathke (high intertidal, marine) and the talitrid amphipod Orchestia gammarellus (Pallas) (semi-terrestrial) have been studied. All the species were collected from the Firth of Clyde, Scotland, during the spring of 1992. The oxygen-carrying capacity of haemolymph from each species was low, although variable, and was correlated with the low concentration of Hc present. The Hc oxygen-affinity of native gammarid haemolymph was relatively high [partial pressure of oxygen required for half-saturation, P₅₀=4 to 5 torr (0.53 to 0.67 kPa)] at their respective in vivo pH values. At equivalent pH, however, Hc from G. locusta displayed a lower O₂-affinity than either Echinogammarus species. Gammarid Hcs had a large Bohr effect ( log P₅₀/ pH=-1.16 to-1.47). Resuspended Hc isolated from whole H. nilsonni showed similar O₂-binding properties to those of the gammaridean amphipods [P₅₀=6.3 torr (1.44 kPa) at pH=8.0; log P₅₀/pH=-1.20]. Comparable data for haemolymph from O. gammarellus showed that the Hc had a lower affinity for O₂ [P₅₀=14.1 torr (1.87 kPa) at in vivo pH] and exhibited a more moderate Bohr effect ( log P₅₀/ pH=-0.79). To eliminate the possibility that these differences were due to the different haemolymph constituents, each of the Hcs were pelleted and resuspended in physiological saline. The differences noted above persisted, demonstrating that they were due to inherent O₂-binding properties of the Hc molecules themselves. An increase in L-lactate resulted in an increase in Hc oxygen-affinity for both Echinogammarus species but not for O. gammarellus. This study has confirmed that there is a clear difference between Hcs from aquatic and semi-terrestrial amphipod genera. The results lend further support to the hypothesis that the move on to land by amphipod crustaceans is accompanied by a decrease in Hc oxygen-affinity, a decrease in the Bohr effect and a decrease in effector (in this case L-lactate) sensitivity.     

Impact of laboratory-imposed physical disturbance on the abundance of four isolates of marine gymnamoebae

Four species of gymnamoebae isolated from a sandy shore were subjected to physical disturbance within sand microcosms containing either 250 or 125 μm sand. Firstly, the effect of two different strengths of physical disturbance on amoebae populations within sand was investigated. Secondly, whether the size of sand grains occupied by amoebae populations changed their response to disturbance events was assessed. Highest abundances of amoebae were recorded in control flasks containing 125 μm sand. Increasing the strength of disturbance decreased the abundance of all four species. Abundances of the two smallest isolates (Platyamoeba sp. and a limax amoeba) were least affected by increasing disturbance. The greatest decrease in abundance was observed in populations of two larger (>15 μm) Mayorella sp. and Vannella sp. The greatest impact of physical disturbance was seen on populations within 250 μm sand. This study shows that physical disturbance may be an important community structuring force within natural marine gymnamoebae communities.     

Amphipods and isopods in the rocky intertidal: dispersal and movements during high tide

Animals in the intertidal, both mobile and sessile, generally exhibit some zonation pattern, in which each species shows a preference for, or is confined to, some height levels. The study of zonation patterns is, however, almost exclusively based on surveys made during low tide, when many animals are relatively inactive. We studied zonation patterns of amphipods and isopods on rocky shores in southwestern Iceland, both by traditional sampling at low tide as well as by sampling during high tide. The distributional patterns seen at high tide differed significantly from that at low tide. One amphipod, Anonyx sarsi, was common around baits at all levels at high tide but absent from the intertidal at low tide. Several other species were either relatively more common or tended to be recorded higher, or in one instance, lower on the shore when the tide was in than at low tide. There was also evidence of some species changing habitats within the intertidal with the tidal cycle. Many species, however, moved little away from their respective zones occupied at low tide, and for some species, including some capable of rapid swimming, very limited mobility was indicated. We conclude that low-tide surveys of the intertidal give an incomplete picture of the community structure, and even key species may be missed in such surveys.Communicated by L. Hagermann, Helsingør     

Ecological effects of intertidal uplifting from nuclear testing

Uplifting of a portion of an intertidal rock bench by the Milrow underground nuclear test at Amchitka Island, Alaska affected algal abundance, species composition, and zonation. Parts of the populations of 8 species died off, primarily in the Hedophyllum sessile zone, and mostly within the first 6 months after the disturbance. The center of abundance of these species shifted seaward. Fucus distichus, a species of the upper intertidal, appeared in the area and recolonized a portion of the H. sessile zone. The observations indicate that intertidal algal communities at Amchitka Island are very sensitive to positive elevation changes.Contribution No. 376 of the College of Fisheries, University of Washinton, USA. This publication is based on work performed under U.S. Atomic Energy Commission Contract AT(26-1)-171 for Battelle Memorial Institute, Columbus Laboratories.     

Distribution of algae on tropical rocky shores: spatial and temporal patterns of non-coralline encrusting algae in Hong Kong

Encrusting algae have been described as dominant space occupying species on rocky shores around the world. Despite their abundance, however, most studies classify species under generic names (e.g. Ralfsia sp.) or as a functional group (e.g. encrusting algae), thereby underestimating the number of species present and their ecological importance. Studies on six rocky shores of varying exposure, in Hong Kong, recorded eight common species of encrusting algae. The greatest abundance of encrusting algae was recorded on shores of intermediate exposure, where four distinct zonation bands could be identified; a cyanobacterial Kyrtuthrix-Zone in the upper midlittoral, a Bare-Zone below this, a Mixed-Zone in the lower midlittoral and a Coralline-Zone in the infralittoral fringe. Abundance declined on shores of greater and lower exposure to wave action, where bivalves and barnacles were competitively dominant. Certain species were found in greater abundance on exposed shores (e.g. Dermocarpa sp. and Hildenbrandia occidentalis), while others preferred more sheltered shores (e.g. Hildenbrandia prototypus and Kyrtuthrix maculans). With the exception of some cyanobacterial crusts, the abundance of encrusting algae was always greatest towards the low shore, an area of decreased physical stress and increased herbivore density. Zonation patterns showed seasonal variation associated with the monsoonal climate of Hong Kong. Most species increased in abundance during the cool season, while during the summer months the cover and vertical extent of encrusting algae decreased in relation to summer temperatures, although K. maculans increased in abundance during the summer. On Hong Kong shores, encrusting algae have a high species richness and exhibit within-functional group spatial and temporal variation which is mediated by herbivory and seasonal, physical stresses.     

Population and species diversity fluctuations in a rocky intertidal community relative to severe aerial exposure and sediment burial

From a series of 10 quarterly assessment between October 1975 and May 1978 (inclusive), fluctuations in abundance were determined for macroinvertebrates and macrophytes of a rocky intertidal community on Santa Cruz Island, California, USA. During afternoon low tides in late fall and winter of the first 2 yr of study, organisms of the upper and middle intertidal zones were subjected to prolonged aerial exposure. Many species there tolerated this exposure, but die-backs occurred for a barnacle (Chthamalus fissus/C. dalli) and several algae (Endocladia muricata, Pelvetia fastigiata f. gracilis, Corallina officinalis var. chilensis, Corallina vancouveriensis, Cylindrocarpus rugosus and Codium fragile). These die-backs were succeeded by blooms of Ulva californica and Porphyra perforata. In the upper and middle intertidal zones, the major cover organisms that could tolerate prolonged aerial exposure were disproportionately prevalent and appeared to be maintained by the periodic repression of species intolerant to such exposure. In February 1978, heavy rainfall caused sediment inundation of the middle and lower intertidal zones. This event was followed by declines in abundance of a barnacle (Tetraclita rubescens) and several algae (Pelvetia fastigiata f. gracilis and corallina spp.). Shannon-Wiener H species diversity fluctuated from a high in October to a low the following May during both 1975–1976 and 1976–1977 in conjunction with the period of increased daytime aerial exposure in late fall and winter. A further decline in diversity following sediment inundation in February 1978 contributed to a long-term trend of declining H species diversity (3.06 in October 1975 to 1.87 in May 1978). We hypothesize that predictable late fall to winter aerial exposure stresses, in combination with a random physical disturbance (sediment burial), exceeded an optimal intermediate level of disturbance predicted for maximal species diversity.     

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.     

Macrozoobenthic community structure in the Ría de Foz,an intertidal estuary (Galicia,Northwest Spain)

The community structure of the macrobenthic (>1 mm) fauna of an intertidal soft-bottom estuary, the Ría de Foz (Galicia, Northwest Spain) was investigated on 99 occasions during June, September and December 1984 and March 1985. The infaunal data were subjected to classification and ordination techniques. The analyses indicated that sediment characteristics (grain size, percentage silt-clay and organic-matter content) and tidal height are the most important factors governing the distribution and abundance of the intertidal communities. Three faunal assemblages were identified and examined in terms of species dominance, constancy and fidelity. In the clean sands of the outer estuary two main assemblages were distinguished; one, corresponding to the boreal LusitanianTellina community, inhabited the middle and lower tidal levels; the other, typical of a biocoenosis of sands and slightly muddy mediolittoral sands, inhabited the upper tidal level. The third benthic assemblage occupied the muddy sands and mud of the inner estuary and displayed some features in common with theCardium edule-Scrobicularia community described by Thorson.     

Variations in benthic community structure in a sub-tropical estuary

Abundances of macroinfaunal invertebrates in sand dollar (Dendraster excentricus) beds and in adjacent areas free of sand dollars were studied at 10 intertidal sites in the Pacific Northwest region of the USA and Canada. Each site was sampled once in late summer of 1977. There were no significant differences in overall diversity of species between the beds and adjacent areas; nor did cluster analysis indicate a sand dollar bed community. However, a polychaete (Armandia brevis) was significantly more abundant inside the beds than just outside them. Conversely, in comparison to the beds, adjacent regions contained significantly more specimens of the following species: a bivalve (Transennella tantilla), several tubicolous crustaceans (Corophium spp., Ampelisca agassizi, and Leptochelia savignyi) and two polychaetes (Glycinde polygnatha and Malacoceros arenicola). Sediment characteristics at each study site were not correlated with the presence or absence of sand dollars; moreover, a comparison among the 10 sites revealed no sediment characteristics typical of sand dollar beds. At most of the sites, significantly fewer tube-building invertebrates occurred inside sand dollar beds than outside. The discussion considers the possibility that mechanical disturbance by the sand dollars tends to exclude some macroinvertebrates (especially tubicolous ones) from sand dollar beds.     

Habitats and tidal accessibility of the marine foods of dabbling ducks and brant in Boundary Bay,British Columbia

The role of birds in intertidal food webs is mediated by effects of tides on the habitat, distribution, and accessibility of food organisms. We studied effects of elevation, sediment characteristics, vegetation, and tidal cycles on waterfowl feeding and food organisms in Boundary Bay, British Columbia, Canada from September to March 1990–1992. Percent dry mass of esophageal contents was mainly Zostera japonica leaves (84%) for American wigeon (Anas americana Gmelin), and leaves of Z. japonica (57%) and Z. marina (41%) for brant (Branta bernicla L.). Northern pintails (A. acuta L.) and mallards (A. platyrhynchos L.) ate mostly the leaves, seeds, and rhizomes of Z. japonica (48 and 72%, respectively) and gastropods (12 and 28%). Green-winged teal (A. crecca L.) ate mainly amphipods (68%) and saltmarsh seeds (19%). The well-sorted fine sand (median 0.125 mm) to medium sand (0.25 mm), and the low organic carbon content (1.31±0.12% dry mass), did not appear to vary sufficiently along 2.1- and 2.6-km transects to affect macroinvertebrate distributions. Three habitat zones were recognized: (1) upper intertidal (sandflats); (2) mid-intertidal (largely monotypic Z. japonica); and (3) lower intertidal (mixed Z. japonica and Z. marina). These zones, although related to elevation, had greater influence on amphipod distributions than did elevation alone or detrital biomass. Biomass and numbers of most invertebrate foods of waterfowl were greatest in the eelgrass zones, with the upper elevational limit of Z. japonica occurring at about Mean Water Level (MWL). Accordingly, wigeon, pintails, and mallards fed mostly below MWL. Because of seasonal tidal cycles, daily accessibility of areas below MWL but within reach of surfacefeeding waterfowl decreased from a mean of 10.8 h in October to 9.6 h in December. In October only 10 d and <12 h available feeding time, whereas in December and January every day had <12 h available feeding time. Seasonal effects of tides on accessibility might limit sufficient feeding by some waterfowl species in intertidal areas, increasing their reliance on alternative feeding sites.     

Comparison of mussel bed community between two intertidal mytilids Septifier virgatus and Hormomya mutabilis

Mussel bed community structure of two intertidal mytilids, Septifer virgatus (Wiegmann) and Hormomya mutabilis (Gould), whose beds were contiguous vertically on a rocky intertidal shore (Wakayama Prefecture, Japan), was compared between 1982 and 1983. In the upper S. virgatus bed, crustaceans and bivalves were dominant in terms of both number of individuals and biomass. There were three barnacle species representative of the epizoans, two isopods and one amphipod as mobile fauna, and two bivalves as infauna. The lower H. mutabilis bed supported virtually no epizoans or mobile fauna. Infaunal free-ranging polychaetes and sipunculids were dominant in terms of both number of individuals and biomass. The H. mutabilis bed contained a much greater amount of sediment than did the S. virgatus bed, and the interstices among individual H. mutabilis and among their byssal threads were filled with sediment. The biomass of six of nine species dominant in the S. virgatus bed was negatively correlated with the amount of sediment. Recruitment of these faunal assemblages into artificial mussel clumps was examined in mussel enclosure experiments, and a negative sediment effect in H. mutabilis clumps was detected for one isopod and one limpet species. Factors causing the differences between two mussel bed communities are discussed, focusing on the effects of sediment.     

Seasonal changes in the structure of an intertidal community

Quantitative (0.25 m²) samples of macrofaunal (>1.0 mm) invertebrates were taken in each season from one habitat of an intertidal sandbar in the North Inlet estuary near Georgetown, South Carolina, USA. During all seasons the community inhabiting the sample site was numerically dominated by two species of haustoriid amphipods (Acanthohaustorius millsi and Pseudohaustorius caroliniensis). Seasonal changes at the community level were clearly controlled by the population dynamics of the numerically dominant species, and qualitative information on life histories was important to the interpretation of analyses' results.This work was supported by the Environmental Technology Center of Martin Marietta Corporation and the Belle W. Baruch Institute for Marine Biology and Coastal Research. It is Contribution No. 138 of the Belle W. Baruch Institute for Marine Biology and Coastal research.     

Environmental stress,competition and dominance of Crassostrea virginica near Beaufort,North Carolina,USA

Observations and experiments were made at 2 intertidal areas near Beaufort, North Carolina, USA from May 1977 to July 1978 to determine why the oyster Crassostrea virginica dominated the community in areas protected from wave action but not in areas directly exposed to waves. Barnacles, oysters, the green alga Enteromorpha sp. and the mussel Brachiodontes exustus were the main occupiers of primary space at the mid and low intertidal levels of exposed areas. The intertidal community at the protected site consisted of a mid intertidal occupied by the barnacles Balanus amphitrite and Chthamalus fragilis, and the oyster C. virginica, and a low intertidal dominated by C. virginica. The exposed area was highly variable with high colonization and mortality for all species producing large seasonal changes in structure. The protected site remained constant throughout the year; there was no evidence of further colonization of either barnacles or oysters and mortality was very low. Recolonization experiments, selective removal of species, and growth and survival data demonstrated that C. virginica does not become dominant at exposed locations because (1) the constant wave shock at the ocean site reduces growth and increases mortality of young and adult oysters and (2) oysters are outcompeted by the mussel B. exustus. The monopolization of space by C. virginica at protected sites contrasts with studies north of Cape Hatteras where the abundance of predators produces a more diverse and heterogeneous community. Predation was unimportant in Beaufort because predators were absent at the exposed areas and the oyster drill Urosalpinx cinerea was restricted to the subtidal zone at the protected site. This absence of predators indicates a higher level of environmental stress near Beaufort compared to areas farther north.     

Distribution and variability in abundance ofSchistomysis spiritus (Crustacea: Mysidacea) in the Bristol Channel in relation to environmental variables,with comments on other mysids

Mysids are important components of the zooplankton biomass of estuaries and coastal regions. Twelve species were identified from the Bristol Channel and Severn Estuary (British Isles). The most abundant species wasSchistomysis spiritus (Norman), the majority of which occurred in the Channel within a salinity range of 27 to 34 S. The seasonal distribution and numerical abundance of this species are described over the period November 1973 to February 1975. The peak of abundance ofS. spiritus occurred in the Inner Channel in September (mean of 14 individuals m^-3, ca. 250 individuals m^-2 for this sub-region) when it represented 76% of the omnivore biomass (g C m^-3); for the 364 d from 4 November 1973, the species contributed 43% of the total integrated omnivore standing stock. This peak of biomass was the product of the development of the juveniles from the reproductive period in the spring. Correlation analyses were carried out betweenS. spiritus biomass and 10 physical and biotic variables for 2 mo, November 1973 and September 1974. Temperature and salinity, which are simple indices of seasonal change, exhibited significant correlations with the mysid's abundance in both months. These correlations do not necessarily imply causal relationships or mechanisms between the distribution and abundance of the species and these variables. Clearly, in such a complex environment as an estuarine ecosystem a single variable is unlikely to control the abundance of a species, it is more the result of the combined influence of a number of variables acting in concert.     

Migrational orientation in two species of littoral gastropods (Littorina angulifera and Nerita senegalensis)

Littorina angulifera (Lamark) and Nerita senegalensis Gmelin, both intertidal gastropods, appear to have a preference for specific levels of the shore. In some experiments conducted both in the laboratory and in the field, displaced snails exhibited a general homing behaviour when displaced upshore or downshore. Cues involved in this migrational behaviour have not clearly been identified.     

Effect of freezing on photosynthesis of intertidal macroalgae: relative tolerance of Chondrus crispus and Mastocarpus stellatus (Rhodophyta)

The effect of freezing on photosynthetic metabolism was studied in the red algae, Chondrus crispus and Mastocarpus stellatus. Plants of both species were collected from the intertidal at Chamberlain or Kresge Point, Maine, USA (43°56N, 69°54W) between February and March 1987. Photosynthetic rates were measured immediately after freezing at-20°C and following recovery periods in seawater. Photosynthesis in C. crispus declined rapidly following freezing, falling to 70% of control values within 1 h and 30% after 3 h exposure. Minimum photosynthetic rates (7 to 9% of controls) occurred following freezing exposures of 12 h or more. Full photosynthetic recovery in C. crispus after 3 h at-20°C required 48 h. Photosynthesis in C. crispus did not fully recover in plants frozen for 6 h or more. In contrast, photosynthesis in M. stellatus was relatively unaffected by freezing exposures of <12 h. Twelve hours or more at-20°C reduced photosynthesis to 55% of controls. Photosynthesis in M. stellatus fully recovered from 24 h at-20°C within 24 h. In both species the reduction of photosynthesis by freezing was associated with damage to the plasma membrane and reduced efficiency of energy transfer from phycobilisomes to chlorophyll a, but did not appear to involve ribulose-1,5-bisphosphate carboxylase oxygenase activity. The freezing tolerance of C. crispus and M. stellatus positively correlates with their respective intertidal distributions, suggesting that freezing may be involved in controlling the distributions of these species on the shore.