PHYSIOLOGICAL AND POPULATION ECOLOGY OF INTERTIDAL AND SUBTIDAL ASCOPHYLLUM NODOSUM (PHAEOPHYTA)1

Morphological, demographic and physiological characteristics of Rhode Island intertidal and subtidal populations of Ascophyllum nodosum (L.) Le Jolis were compared in order to examine factors influencing vertical distribution. The two populations had distinctive morphologies: subtidal plants were narrower (more terete) and highly branched compared with intertidal plants. The subtidal population showed signs of necrosis and breakage, which was reflected in significantly shorter mean plant size. High survivorship and low recruitment of both population resulted in relatively constant densities, averaging 91 and 50 plants per m² in the intertidal and subtidal habitats, respectively. Intertidal plants had higher mean annual growth rates (25 cm.yr^?1) than subtidal plants (2 cm.yr^?1). In general, intertidal plants had higher photosynthetic capacity and nutrient (NO₃^?) uptake rates than the subtidal population but maintained lower light-harvesting pigment and tissue nitrogen concentrations. Although Ascophyllum nodosum is capable of survival and growth in subtidal as well as intertidal areas, results of this study suggest that different selective pressures affect persistence in each habitat. The scarcity of plants in the subtidal environment may be due to the lack of a critical balance between algal production, allocation of photosynthate, and the negative effects of grazers or competitors.     

Intraspecific variation in the population dynamics and growth of the limpet,Cellana tramoserica

Summary Populations of the limpet Cellana tramoserica (Sowerby) from high, mid, and low intertidal regions, and from a subtidal zone, were studied at Cape Banks (N.S.W.), Australia. Individuals from the subtidal population had the largest mean and maximum shell size, the low and high shore populations were intermediate, while the midshore population had both the smallest mean and maximum size. The density of adults showed the reverse trend: the midshore region had the greatest adult population density while the subtidal population had the smallest density. The density of juveniles and recruits was negatively correlated with tidal height in the intertidal areas, but the density of both age-classes was smallest in the subtidal region. The rate of growth of individuals was negatively correlated to the adult density of a site; individuals from the subtidal population grew the fastest, while those from the midshore region grew the slowest. The level of adult mortality of the four populations was similar, with an annual rate of between 50–60%. Juvenile mortality did, however, differ among populations; reduced percentages of juveniles reached adult size in the mid and low shore populations than in the highshore and subtidal populations.The expected lifetime fecundity of individual females differed among the four populations. Subtidal individuals were expected to spawn 40 times the amount of gonad material during their life than were individuals from the midshore population. Individuals from the high and low populations probably spawn 10 times more than those from the midshore region. Differences in the fecundities of individuals were not likely to be the result of genetic differences because experimental transplants and manipulations of density showed that individuals from all the intertidal populations could increase their growth rate to match that of subtidal individuals. The implications of the difference in lifetime fecundity among populations are discussed.     

Micropopulation differentiation in phenol content and susceptibility to herbivory in the Chilean kelp Lessonia nigrescenss (Phaeophyta,Laminariales)

Micropopulation differences in phenol content between intertidal and subtidal individuals of the kelp Lessonia nigrescens were found. Subtidal plants showed: (1) significantly higher phenol content than intertidal individuals, in vegetative and reproductive tissues, (2) intra-plant differences, with higher content in apical frond tissues, (3) higher resistance to consumption by herbivorous fishes. The microscopic progeny of subtidal plants showed the same trend as adult plants: (1) haploid spores from subtidal plants had higher phenol content than spores from intertidal individuals, and (2) the microscopic sporophytes derived from subtidal spores and gametophytes were less consumed by herbivorous snails (Tegula tridentata) than those derived from intertidal plant propagules. No increase in phenol content was detected after mechanical injury to experimental fronds, or after transplantation to the subtidal environment.In addition to the absence of inducible responses, the different phenol content between intertidal and subtidal individuals, in adult diploid plants and also in the haploid progeny, suggests that both environments differ someway enough to fix the mentioned features on the plants of Lessonia nigrescens. It is likely that the differences in herbivory between the two distributional extremes contributed to the observed pattern.     

Variability in the reproductive effort of the limpet,Cellana tramoserica

Summary The reproductive effort of the limpet, Cellana tramoserica was investigated in high, mid, and low intertidal areas of the rocky shore as well as in a subtidal population. The reproductive effort of individuals within a population was independent of age. There was, however, a noticeable difference among populations with respect to the relative amount of gonad material spawned per year. The individuals from the subtidal population had the greatest reproductive effort, spawning the equivalent of 101% of their somatic weight per year, while the low, mid and high intertidal populations spawned the equivalent of 66%, 20%, 51%, respectively.Experimental transplants showed that individuals from all three intertidal populations increased their reproductive effort to match that of subtidal individuals. It was unlikely, therefore, that the observed patterns were the result of genetic differences among the populations. The reproductive effort of the populations was also not correlated with the rate of mortality of the adults in the four populations. The observed differences could be explained by a combination of the availability of food and intraspecific competition. The flexibility in the reproductive pattern of Cellana is hypothesised to be a result of this species having a dispersive larval stage, which may result in selection for plasticity of reproductive allocation, depending upon the environmental conditions at the site of settlement and growth to adult size.     

Microbial-Activated Sediment Traps Associated with Oncolite Formation Along a Peritidal Beach,Northern Arabian (Persian) Gulf,Kuwait

Identification of microbial communities within shoreline sediments and sediment precipitates from the Tigris-Euphrates delta (northern Kuwait) were determined by microscopic/nanoscopic studies, and by molecular analysis. Oncolites are syn-diagenetic carbonate precipitates that are surviving in a shallow subtidal to intertidal siliciclastic environment with periodically excessive hydraulic energy, extreme salinity (up to 47 per mil), and high concentrations of organic matter. X-ray diffraction techniques reveal that oncolite cortices are predominantly composed of calcite, quartz, halite and dolomite, associated with minor fractions of clay minerals. Quantitative analysis of the Corey Shape Factor reveals distinct morphological populations but with local overlap. A plot of the Equivalent Diameter vs. Corey Shape Factor provided the best indicator of the morphological relationships within the total oncolite population, indicating a hydrodynamically controlled morphological distribution defining intertidal and subtidal oncolite classes. Direct microscopic examination of the samples indicates that diatoms are the most abundant eukaryotic algae in subtidal sediments and within actively precipitating carbonate cements, especially the genus Navicula. In contrast, filamentous cyanobacteria from the genus Anabaena are most abundant in the intertidal zone sediments. The PCR-DGGE of the 16SrRNA gene of the cyanobacteria shows a higher diversity for this genus of bacteria in all sediment samples and that the cyanobacterial population in the diagenetically precipitating oncolites are closely related to the population found in the subtidal sediments. Dunaliella viridis dominates the culturable algae obtained from the four tidal zones. Our results indicate that a range of microbial populations are actively contributing to the formation of microbially-induced sedimentary structures in the extreme conditions of the southern Tigris-Euphrates delta.     

Size and Performance of Juvenile Marine Invertebrates: Potential Contrasts Between Intertidal and Subtidal Benthic Habitats

Newly settled or hatched juveniles of marine benthic invertebratesgenerally experience very high mortality. Juvenile mortalitycan profoundly affect adult populations, but little is knownabout how individual variation in juvenile quality affects performance.Several recent studies have demonstrated that differences insize, larval nutrient stores, or larval feeding history canstrongly affect the performance (measured as growth and survivorship)of juveniles. Additional research suggests that the strengthof the effect of juvenile size on performance may be mediatedby variation in environmental stress in the intertidal, a habitatcharacterized by strong fluctuations in abiotic factors. Themajor sources of juvenile snail mortality are likely to differin intertidal and subtidal habitats; abiotic stresses relatedto exposure, such as desiccation, are important in the intertidalbut far less severe in subtidal environments. Previously observedtrends in hatching or settlement size between intertidal andsubtidal species from three gastropod taxa may be due to differingselective regimes acting on initial juvenile size.     

Modelling nearshore intertidal habitat use of young-of-the-year Pacific sand lance (Ammodytes hexapterus) in Barkley Sound, British Columbia, Canada

The Pacific sand lance (Ammodytes hexapterus) is an ecologically important prey species for many vertebrate marine predators in the Pacific Northwest. In this study, we examined the use of intertidal habitat by young of the year (YOY) sand lance in Barkley Sound, British Columbia. We selected 14 of the original 29 independent environmental variables for modelling based on univariate analysis for variable selection. These were then used to model sand lance presence-absence using a classification tree approach. Based on our models we found that sand lance avoided mud and intertidal eelgrass. For sites that had very little mud and no intertidal eelgrass, sand lance preferred sediment size mean ≥1,918 μm or sites with sediment size mean ≤1,918 μm but with relatively well sorted sediment (sorting values ≤2.56 SDs, used as a heterogeneity index of the substrate grain size). Adjacent subtidal characteristics were not found to be important at this scale of study. This suggests intertidal substrate characteristics and presence-absence of intertidal eelgrass are the main influences on occurrence of YOY in the intertidal. Our results support the hypothesis that sand lance are associated with particular sediment types, however intertidal sediment types used by sand lance in our study differ from known preferences of sand lance for subtidal sediment types. This difference may be due to unique habitat constraints for intertidal versus subtidal regions. Although it is unknown if the results of this study are widely applicable, the results begin to identify intertidal habitat features that are important for Pacific sand lance.     

INTRASPECIFIC VARIATION IN STRESS‐INDUCED HYDROGEN PEROXIDE SCAVENGING BY THE ULVOID MACROALGA ULVA LACTUCA1

We investigated the presence and kinetics of the oxidative stress response in intertidal and subtidal individuals of the ulvoid macroalga Ulva lactuca L. Stress responses, as measured with both enzymatic and fluorescent‐based antioxidant assays, differed between individuals collected from a subtidal and an intertidal habitat. Subtidal individuals secreted significantly more hydrogen peroxide (H₂O₂) than intertidal individuals when subjected to osmotic stress or desiccation. The activity of reactive‐oxygen‐scavenging enzymes and the ability to scavenge exogenous H₂O₂ were lower in subtidal than in intertidal individuals, suggesting that subtidal individuals are less stress tolerant. In vitro experimentation demonstrated that millimolar concentrations of dimethylsulfoniopropionate (DMSP) and its breakdown products could efficiently scavenge H₂O₂, with DMSP being a less‐effective scavenger than dimethyl sulfide (DMS), acrylic acid, and acrylate. The addition of H₂O₂ at concentrations of 2.5 mM or greater induced the cleavage of DMSP into DMS and acrylic acid in subtidal individuals. Intertidal individuals were affected in the same manner with the addition of 5 mM H₂O₂. There were no differences in the amounts of DMSP cleavage in subtidal and intertidal algae when the algae were subjected to hyposaline conditions. Our data suggest that the oxidative‐stress‐induced cleavage of DMSP affords products with efficient H₂O₂‐scavenging abilities. In addition, U. lactuca individuals growing in intertidal habitats are better acclimatized to changing environments and thus have a higher threshold for oxidative stress than conspecifics in subtidal habitats.     

Spatial and temporal variation of photosynthesis in intertidal Mazzaella laminarioides (Bory)Fredericq (Rhodophyta, Gigartinales)

The red alga Mazzaella laminarioides is an economically important species with an extended latitudinal distribution along the Chilean coast. Its populations form mid-intertidal stands, several meters wide, and therefore are differentially exposed to environmental variables that result in temporal and spatial variability in productivity. We evaluated the effect of latitude and intertidal height on productivity by in situ measurement of photosynthetic performance. Daily and seasonal variations of O₂-evolution rate and maximal quantum yield (F _v/F _m) were determined in plants from the upper and lower intertidal zone at two localities 1500,km apart. Results suggest that plant responses were mainly affected by irradiation, temperature and desiccation. At local level, upper intertidal plants showed a reduced photosynthetic rate and quantum efficiency as compared to those displayed by plants from the lower intertidal, indicating their higher level of excitation energy acclimation. Stronger acclimation differences between upper and lower intertidal plants were observed in spring and summer. Differences in photosynthetic parameters between reproductive phases were recorded in autumn and winter, regardless of the position of the individuals in the intertidal zone. The effects of tidal elevation on seasonal patterns of photosynthesis were also influenced by latitude. Seasonal variation in photosynthetic efficiency was observed in plants from the northern population at both intertidal elevations, but only at the upper intertidal level in the southern population. This study shows that production variability in M. laminarioides results from differences in the intensity of environmental factors observed seasonally at local (intertidal) and latitudinal scales.     

Amphipods exclude filamentous algae from the Western Antarctic Peninsula benthos: experimental evidence

Hard bottom, subtidal communities along the Western Antarctic Peninsula are dominated by forests of large, chemically defended macroalgae that support a very dense assemblage of amphipods. Free-living filamentous algae are rare in the subtidal, but filamentous algal endophytes are common in many of the larger macroalgae, both likely as the result of amphipod grazing pressure. Filamentous algae are common in the intertidal, but primarily in the upper intertidal and on high-energy shores where amphipods are likely to be excluded much of the time. We tested the hypothesis that free-living, filamentous algae would be rapidly consumed if transplanted from the intertidal to the subtidal, and our results clearly supported this hypothesis. The filamentous, intertidal green alga Cladophora repens was transplanted to the benthos in 6 different macroalgal habitats. Control algae were transplanted in 3 m deeper waters nearby (usually 12 m or less laterally) but suspended 3 m off the bottom where amphipods are absent or rare. Overall consumption during approximately 6 h on the bottom ranged from 22 to 98% of the initial biomass, while significantly less biomass loss occurred in the water column.     

High morpho‐anatomical variability in Halimeda macroloba (Bryopsidales,Chlorophyta) in Thai waters

Halimeda macroloba is a common widespread and morphologically variable species in the Indo‐Pacific Ocean. A series of morphometric data (16 morphological and 46 anatomical variables) and the chloroplast‐encoded tuf A gene were examined to explore the morphological and genetic variations within the species and to better determine species boundaries in H. macroloba in Thai waters. Moreover, the environmental conditions, which may affect morphological variations in H. macroloba, especially of Haplotype I were examined. Our results showed that H. macroloba has more morpho‐anatomical variability and broader species boundaries than previously thought in several characters: holdfast type, segment shape and size, node height with differences in pore size, the presence of two additional patterns of peripheral utricle as seen in surface view, and additional layers of utricles (up to seven layers). Additionally, two new variable characteristics have been reported for the first time, i.e. shape and margin of peripheral utricles. Water depth and light correlated with thallus morphology, larger thalli and segments with a greater number of utricle layers were associated with subtidal plants, while smaller thalli and segments were found in the intertidal or high light intensity areas. The high plasticity of this species may explain why in the Indo‐Pacific Ocean this species is common and dominant.     

Is there a risk to living large? Large size correlates with reduced growth when stressed for knapweed populations

A central hypothesis in ecology is that plant life history evolution is constrained by fundamental “compromises between the conflicting selection pressures resulting from particular combinations of competition, stress, and disturbance”, with stress being defined as abiotic conditions that restrict production. Biogeographic differences among native and non-native ranges of invasive plants may provide unique opportunities for tests of this theory. We conducted a greenhouse experiment with Centaurea stoebe plants from North American and European populations. We compared the total biomass and phenotypic plasticity indices for plants from the native and non-native ranges under stressed and non-stressed conditions. The average size of Centaurea stoebe plants from 13 North American populations was greater than that of plants from 18 European populations regardless of stress treatment. However, when plants from the same populations were exposed to lower resources the differences in biomass between plants from North American and European populations were significantly less, suggesting that large plants were poorer stress tolerators. For all 31 populations the regression slope for the relationship between mean mass for populations in non-stressful conditions and mean mass in stressful conditions was less than 1.0, indicating that populations that produced large plants in good conditions also produced plants that grew disproportionately less in stressful conditions. These findings suggest that Centaurea stoebe may be evolving towards being a good “competitor” (sensu Grime, 1977) in its invasive range, but at the cost of being a good “stress tolerator”.     

Production and population ecology ofPhyllospadix iwatensis Makino. II. Comparative studies on leaf characteristics,foliage structure and biomass change in an intertidal and subtidal zone

A seagrass in Japan,Phyllospadix iwatensis Makino, is distributed in the lower intertidal zone and upper subtidal zone making a dense population on the Choshi coast, Japan. IntertidalP. iwatensis is able to receive sufficient light for photosynthesis but experienced severe exposure to the air, which decreased a large amount of aboveground biomass in April to June (i.e. the daytime exposure season). SubtidalP. iwatensis was never exposed throughout the year and the aboveground biomass increased gradually over the daytime exposure season. However, the maximum aboveground biomass and shoot density of the subtidal plant never exceeded that of the intertidal plant. The dense foliage, large aboveground biomass and high shoot density of both intertidal and subtidal plants is likely to be an adaptation to heavy water movement, but the subtidal plants always received insufficient light for photosynthesis as a result of having dense foliage, particularly in turbid water. In choppy and swell sea,P. iwatensis did not seem to be adapted to growing in the subtidal zone where there was shortage of light.     

Disturbance and organization of macroalgal assemblages in the Northwest Atlantic

Large seaweeds are often structurally dominant in subtidal and intertidal rocky shore benthic communities of the N.W. Atlantic. The mechanisms by which these algal assemblages are maintained are surprisingly different in the two habitats. In the subtidal community, kelps are dominant space competitors in the absence of strong grazing interactions. In contrast, the large perennial seaweeds of intertidal zones (fucoids and Chondrus crispus) are competitively inferior to both sessile filter feeders and ephemeral, pioneer algal species. Intertidal seaweed beds are maintained by carnivory of whelks, which reduces filter feeder populations, and by herbivorous periwinkles which reduce ephemeral algal populations. Through most of the intertidal zone, disturbance, both biological and physical, dictates which species shall compete and equilibrium conditions obtain subsequently.The roles of subtidal consumers are quite different. Sea urchins are the major algal herbivores and these voracious animals maintain an equilibrium state in which large tracts of subtidal coralline pavement are kept free of kelp forests. Urchins do not seem to play a successional facilitative role for kelps in the way that periwinkles do for fucoids in the intertidal. Control of herbivore populations is thus a key to the maintenance of subtidal foliose algal beds. It is clear that parasitic amoebas can decimate sea urchin populations so that kelp forest dominance is assured. However, the importance of carnivory in limiting urchins in the subtidal community is unclear in the absence of appropriate manipulation experiments. It is possible that carnivorous decapods and fin fish control sea urchin populations and hence foliose algal abundance, but this must remain speculative. The seaweed-dominated state of the subtidal system is an alternative equilibrium condition to the urchin/coralline alga configuration. The structure of the kelp beds is relatively uniform in responding to frequent small-scale, infrequent large-scale, or no, disturbance.     

Morphological variability of Carex spicata Huds. utricles among plant communities

Differentiation in the size and number of seeds among populations or particular individuals of a given species may depend on genetic features and environmental conditions. The objective of our study was to answer two questions: whether any differences exist in the size and shape of utricles among Carex spicata populations growing in several plant communities and whether the hypothesized differences remain constant in ‘common garden’ conditions (i.e. if the sedges grown in different plant communities are evolving distinctly separate ecotypes). We studied utricle morphological traits (projected area, width, length, W/L ratio and projected perimeter) and number of utricles per spike collected from plants grown in five different plant communities (natural sites) and from plants transferred to common garden conditions. C. spicata utricles showed significant variability of morphological traits that depended on the plant community of origin. Among five plant communities, the largest utricles were found in Agropyron repens–Urtica dioica and Convolvulo arvensis–Agropyretum repentis communities, while the smallest ones were in plants from the Lolio-Plantaginetum community. The similarity of the analyzed populations regarding sedge utricle traits corresponded to the similarity of plant communities where the populations had grown. Moreover, the differences in utricle traits collected from natural conditions remained stable in common garden conditions. This indicated that differences in traits among utricles had a physiological dependence on different biotope conditions or different interspecific interactions prevailing in the particular plant communities. The relationship between the similarities in the diaspores of the populations studied and the similarities in the ecological conditions of the communities may also indicate that the variability of utricles is important for evolution and adaptation.The results support the hypothesis that C. spicata populations in different communities are producing separate ecotypes, i.e. specific species forms with genetically fixed traits adapted to narrowly determined habitat conditions.     

Contrast Between Solitary and Clonal Lifestyles in the Sea Anemone Anthopleura elegantissima

The common intertidal sea anemone Anthopleura elegantissima(Brandt) occurs in two forms the clonal aggregating form andthe solitary form The obvious differences between the two apparentlyresult from the presence of asexual reproduction in the clonalaggregating form and its absence in the solitary form resultingdifferences in growth form suit the two anemones to differentlifestyles The clonal form is well suited to life higher inthe mid intertidal Asexual reproduction resulting in moderatelysmall individual size and close association with clonematesimproves its resistance to physical stress (drag and desiccation)and makes it a superior competitor in exposed habitats higherin the intertidal where species diversity and predator pressureare low The larger solitary animals live in more protected microhabitatslower in the intertidal and subtidal and are probably more resistantto predation and less able to withstand physical stress or intenseintraspecific competition The two forms also have differentbiogeographic ranges While the clonal aggregations are quitecommon at rocky sites at least as far north as Vancouver Islandin British Columbia populations of solitaries have not beenfound north of Point Reyes California Apparently the two formsare re productively isolated phenotype frequencies are verydifferent for clonals and solitaries living at the same locationswhile the between-sample variation within each form is relativelysmall These differences in phenotype frequencies biogeographicrange and microhabitat suggest that the clonal anemones andthe solitary anemones known as Anthopleura elegantissima areactually a sibling species pair.     

Localization and antioxidant capacity of flavonoids from intertidal and subtidal Halophila johnsonii and Halophila decipiens

In this study, flavonoid localization, content and total antioxidant capacity in leaves of subtidal Halophila decipiens were compared to intertidal and subtidal Halophila johnsonii. H. johnsonii leaves had significantly higher flavonoid content (3.5 and 3.8 nmol quercetin equivalent mm⁻² leaf for intertidal and subtidal H. johnsonii, respectively) and antioxidant capacity (101.7 and 224.2 nmol Trolox equivalent mm⁻² leaf for intertidal and subtidal H. johnsonii, respectively) than H. decipiens leaves (1.4 nmol quercetin equivalent mm⁻² leaf and 21.0 nmol Trolox equivalent mm⁻² leaf). Flavonoid content did not significantly differ between intertidal and subtidal H. johnsonii, however, antioxidant capacity was significantly higher in subtidal plants. Confocal laser scanning microscopy of fresh leaf cross sections indicated that both species contained flavonoids in the cuticle, but only H. johnsonii contained intracellular flavonoids. Intracellular flavonoids are better situated to perform antioxidant functions in planta. These results suggest that flavonoid compounds in H. johnsonii are capable of sunscreen and antioxidant functions while an antioxidant role for flavonoids within H. decipiens is not supported.     

Differential population response of allocation,phenology, and tissue chemistry in <Emphasis Type="Italic">Spartina alterniflora</Emphasis>

Phenotypic variation within species is widespread among salt marsh plants. For Spartina alterniflora, the dominant species of low intertidal wetlands across the Altantic and Gulf coasts of the US, distinct phenological and morphological differences among populations from different latitudes have been found. To determine whether S. alterniflora plants from lower latitudes and those regenerated from Delaware tissue cultures would maintain differences from that of native plants, we conducted a field study in a natural salt marsh in Delaware, US. After two growing seasons, plant height, stem density, above- and belowground biomass, elemental composition, and nutrient resorption were measured. Natural variation in porewater salinity influenced physiological traits of Na⁺/K⁺ ratio regulation and nitrogen resorption efficiency similarly across populations. While plant height exhibited plasticity where populations tended to converge to a similar height, several other traits remained distinct. Delaware plants had a greater rate of rhizome growth than Georgia and Louisiana plants, which correlated with a greater magnitude of fall senescence. If traits such as seasonal translocation are plastic and can change with the length of the growing season, climate warming may alter belowground biomass production of S. alterniflora in wetlands of the mid-Atlantic.     

Tidal activity rhythms and depth distribution of rocky shore fish in an altered intertidal environment

In this note we report on changes of activity level of littoral fish and their use of different microhabitats in an altered intertidal environment, where intertidal fish are never emersed and turbulence is confined to high tides. Despite these atypical conditions, the structure of the assemblage is basically the same found in a typical rocky-shore situation. Two intertidal fish known to possess internal tidal rhythms, Lipophrys pholis and Coryphoblennius galerita, retain those rhythms in these altered conditions, and the associated social changes in L. pholis are also retained. These observations support the studies of Gibson (1971), who showed that changes in hydrostatic pressure were of prime importance in keeping the tidal rhythm entrained. The subtidal Parablennius pilicornis, on the contrary, is more active during low tide than at high tide. The hypothesis that some subtidal species are excluded from the intertidal by a turbulence-avoiding mechanism is discussed.     

Direct effects of physical stress can be counteracted by indirect benefits: oyster growth on a tidal elevation gradient

The paradigmatic gradient for intertidal marine organisms of increasing physical stress from low to high elevation has long served as the basis for using direct effects of duration of water coverage to predict many biological patterns. Accordingly, changes in potential feeding time may predict the direction and magnitude of differences between elevations in individual growth rates of sessile marine invertebrates. Oysters (triploid Crassostrea ariakensis) experimentally introduced at intertidal (MLW+0.05 m) and subtidal (MLW–0.25 m) elevations in racks provided a test of the ability to use duration of water coverage to predict changes in growth. During early-to-mid winter, a depression of 38–47% in shell growth of intertidal oysters matched the 36% reduction in available feeding time relative to subtidal oysters. In late winter as solar heating of exposed oysters increased, growth differences of 52–55% departed only slightly from the predicted 39%. In spring, however, duration of water coverage failed to predict even the correct direction of growth change with elevation as intertidal oysters grew 34% faster despite 39% less feeding time. Intense seasonal development of shell fouling by other suspension feeders like ascidians, mussels, and barnacles on subtidal (94% incidence) but not on aerially exposed intertidal (21–38% incidence) oysters may explain why duration of water cover failed to predict spring growth differences. Less intense fouling develops on intertidal oysters due to the physiological stress of aerial exposure on settlers, especially during higher temperatures and longer solar exposures of spring. Fouling by suspension feeders is known to reduce growth of the host through localized competition for food and added energetic costs. Thus, in springtime, indirect effects of aerial exposure providing a partial refuge from biological enemies overwhelmed direct effects of reduced duration of water coverage to reverse the expected pattern of slower intertidal growth of a marine invertebrate.