Comparative population biology and reproduction of two sympatric crabs (Grapsidae) on Azores cobble beaches

Population biology of the intertidal crabs Pachygrapsus marmoratus Fabricius, 1787 and P. maurus (Lucas 1846) was investigated at cobble beaches with different levels of wave exposure. Surveys were taken monthly over 1.5 years at three intertidal levels. The growth rate of both species was similar, although P. marmoratus reached larger sizes. This species was more abundant on the mid-littoral zone of sheltered beaches, whereas P. maurus was more abundant on more exposed ones. The sex ratio was male biased in both species, particularly for the smaller and larger size classes. Both species showed an extended reproductive season, coinciding with the rising temperatures of spring and summer, with ovigerous females being found from June to September for P. marmoratus and from May to August for P. maurus. The present study indicates that both could develop adaptive strategies allowing them to coexist in the Azores littoral. The greater size of P. marmoratus seems to be an advantage on the competition for refuges with P. maurus, which results in its higher abundance and general dominance in the higher shore levels and in less exposed shores. However, P. maurus seems to be more fitted to live in the higher hydrodynamic habitats.     

A refutation of critical tidal levels as determinants of the structure of intertidal communities on British shores

Early hypotheses to account for the pattern of zonation and vertical distribution of species on rocky shores in Britain invoked the concept of critical tidal levels. These levels were at heights on the shore where the upper or lower boundaries of distribution of a number of species coincided. Critical levels were correlated with heights on the shore where there were changes in the rate of change, with height, of the annual proportion of the time spent emersed. This was calculated from predicted tide tables. One implications of this was that different assemblages of intertidal species were present between particular levels on the shore. Futhermore, this hypothesis depended on patterns of vertical distribution of several species being identically controlled by physical factors associated with the rise and fall of the tide. This conflicts with more recent hypotheses, based upon experimental evidence, which include the effects of biological interactions among species on their patterns of distribution.The methods of calculation of the annual emersion curve were approximate, with a high degree of extrapolation. The hypothesis that critical levels exist has not been quantitatively tested.New, more accurate calculations of the emersion curve, from predicted tidal heights, indicate a smooth, monotonic curve against height on the shore. This eliminates the possibility of correlation between the coincident boundaries of distribution of a group of species and a height at which the curve changes in slope.The upper and lower boundaries of species were recorded in transects on five shores in different parts of Britain. If critical level exist, the upper or lower boundaries of a set of species must depart from random dispersion up and down the shore, to be clumped or aggregated at the critical levels. A test for non-random dispersion of the boundaries indicated no significant departure from random on any of the shores sampled.There is thus no evidence that critical tidal levels exist, because there is no evidence that the upper, or lower, boundaries of vertical distribution of intertidal species are in any way aggregated. There is no possibility of correlation between changes of slope of the emersion curve at particular heights on the shore and the patterns of distribution of intertidal species on British shores. The hypothesis of critical tidal levels must be abandoned and the rôle of the effects of tidal rise and fall on the vertical distribution of intertidal species must be re-examined in any new hypotheses to account for observed patterns of zonation.     

Thermal tolerance,evaporative water loss,air-water oxygen consumption and zonation of intertidal prosobranchs: a new synthesis

Duration of emergence increases with tidal height on rocky shores therefore, emergence adaptations in intertidal species such as littorine and other prosobranch gastropods have been considered correlated with zonation patterns; temperature tolerance, desiccation resistance and aerial respiration rate all commonly assumed to increase progressively with increasing zonation level. Such direct correlations are rarely observed in nature. Maximal aerial gas exchange occurs in mid-shore, not high shore species. Temperature tolerance and desiccation resistance do not increase directly with shore height. Thus, hypotheses regarding physiological correlates of zonation require revaluation. A new hypothesis is presented that the high tide mark presents a single major physiological barrier on rocky shores. Above it, snails experience prolonged emergence and extensive desiccation; below it, predictable submergence and rehydration with each tidal cycle. Thus, desiccation stress is minimal below the high tide mark and maximal above it. Therefore, species restricted below high tide (the eulittoral zone) should display markedly different adaptive strategies to emergence than those above it (the eulittoral fringe). A review of the literature indicated that adaptations in eulittoral species are dominated by those allowing maintenance of activity and foraging in air including: evaporative cooling; low thermal tolerance; elevated aerial O₂ uptake rates; and high capacity for radiant heat absorption. Such adaptations exacerbate evaporative water loss. In contrast, species restricted to the eulittoral fringe display adaptive strategies that minimize desiccation and prolong survival of emergence including: foot withdrawal, preventing heat conduction from the substratum; aestivation in air; elevated thermal tolerance reducing necessity for evaporative cooling; position maintenance by cementation to the substratum and increased capacity for heat dissipation. In order to test of this hypothesis the upper thermal limits, tissue and substratum temperatures on emergence in direct sunlight and evaporative water loss and tissue temperatures on emergence in 40 °C were evaluated for specimens of six species of eulittoral and eulittoral fringe gastropods from a granite shore on Princess Royal Harbour near Albany, Western Australia. The results were consistant with adaptation to the proposed desiccation barrier at high tide. The eulittoral species, Austrocochlea constricta, Austrocochlea concamerata, Nerita atramentosa and Lepsiella vinosa, displayed adaptations dominated by maintenance of activity and foraging during emergence while the eulittoral fringe littorine species, Bembicium vittatum and Nodilittorina unifasciata displayed adaptations dominated by minization of activity and evaporative water loss during emergence. The evolution of adaptations allowing tolerance of prolonged desiccation have allowed littorine species to dominate high intertidal rocky shore gastropod faunas throughout the world's oceans.     

Physical and biological conditions on a steep intertidal gradient at Rottnest Island,Western Australia

Seven species of grazing molluscs, two littorinids, one nerite, three limpets and one chiton, lived on the vertical intertidal rock wall at the landward edge of the coastal limestone platforms at Rottnest Island, Western Australia. On the average, the vertical ranges of these species overlapped broadly, although a consistent zonation pattern was conspicuous from the platform surface upwards for 2 m. On these vertical intertidal shores, physical and biological conditions were predicted quantitatively from easily made measurements of vertical height on the shore; the percentage of time any shore level was immersed in seawater, the percent weight loss of plaster clods, the standing crop of algae, and the growth rate of the limpet, Notoacmea onychitis, all decreased linearly or semi-logarithmically with increasing height on the shore. The standing crop of animals was greatest, largely due to the presence of the chiton, between 40 and 70 cm from the platform surface and decreased rapidly down the shore and more gradually towards higher shore levels. We interpret this information and the positive correlation between algal production rate, and egestion rate of the animal community at various levels of the shore as evidence supporting the idea that food may be in short supply on these shores.     

Dislodgement force and shell morphology vary according to wave exposure in a tropical gastropod (Cittarium pica)

Wave exposure has strong influences on population density, morphology and behaviour of intertidal species in temperate zones, but little is known about how intertidal organisms in tropical regions respond to gradients in wave exposure. We tested whether dislodgement force and shell shape of a tropical gastropod, Cittarium pica, differs among shores that vary in wave exposure. After adjusting for body size, we found that C. pica from exposed shores required greater dislodgement force to remove them from the shore, had slightly larger opercula (the closure to the shell aperture), and were slightly squatter in shape (reduced in shell height relative to shell width) than C. pica from sheltered shores. These morphological adjustments are consistent with those observed in temperate gastropods, which are argued to represent adaptive responses to the risk of mortality associated with dislodgement.     

Influence of shell morphometry,microstructure, and thermal conductivity on thermoregulation in two tropical intertidal snails

Tropical intertidal organisms tolerate large fluctuations in temperature and high desiccation rates when exposed during low tide. In order to withstand the short‐term heat stress, intertidal organisms adopt behavioral responses to maximize their survival. Our previous research showed that tropical littorinids found at the upper and lower intertidal shores in Singapore exhibited different behavioral adaptations during low tide. Most of the upper‐shore Echinolittorina malaccana kept a flat orientation, with the aperture against the substrate and the long axis of the shell towards the sun, whereas a majority of the lower‐shore individuals of Echinolittorina vidua stood with the edge of the aperture perpendicular to the substrate on the rocky shore during low tide. This prompted analyses of the shells of these two species to determine whether the differences in the shell morphometry, microstructure, and thermal conductivity of shells of E. malaccana and E. vidua were associated with their respective behavioral responses to thermal stress. Analyses of shell morphometry and thermal conductivity showed that shells of E. malaccana were more likely to minimize heat gain, despite having a higher thermal conductivity on the outer surface, due to their light‐gray, elongated shell. By contrast, the dark‐colored, globose shells of E. vidua probably gain heat more readily through solar radiation. Scanning electron microscopy images of the shells of both littorinid species further revealed that they have cross‐lamellar structure; however, only individuals of E. vidua showed the presence of disjointed rod layers and a pigmented inner shell surface. Individuals of E. malaccana had a rough outer shell surface with holes that inter‐connect to form water‐trapping channels that probably aid cooling. Individuals of E. vidua, however, had a smooth outer surface with rows of kidney‐shaped depressions as microsculptures which probably help to stabilize shell shape. In both Echinolittorina species, behavioral responses were used to overcome thermal stress during low tide that was associated with shell morphometry and shell thermal conductivity. Such combined adaptations increase survivability of the littorinids at their respective tidal levels.     

Spatial and vertical patterns in the tidepool fish assemblage on the island of O`ahu

The microtides, wave regimes, and relative isolation of the Hawaiian archipelago may provide unique environmental and biogeographic effects that shape the structure of tidepool fishes. We sampled fishes across a narrow gradient at low tide from 6 sites on the island of O`ahu. We tested predictions of the hypotheses that environmental conditions (pool depth, volume, macroalgal cover, temperature, and salinity) would result in a vertically structured tidepool fish assemblage unique to basalt or limestone rocky shores. 343 fish were recorded from 40 pools, and 19 species from 10 families were identified. Tidepool fish diversity (H’: O`ahu = 2.4; Sites Average = 0.0–0.9) was typical for tropical islands, with members from Gobiidae (5 species), Blenniidae (4 species), Pomacentridae (3 species), Acanthuridae (2 species) and Kuhliidae (2 species) among the most common. Endemism (32%) was higher than other well studied assemblages yet similar to Hawaiian reef fishes (25%). Assemblage abundance varied among shores with basalt or limestone substrate, among sites, and vertically among high, mid, and low pools. In general, blenniids occurred at higher proportions on limestone shores and gobiids were more common on basalt shores. High pools were characterized by an abundance of a small sized (29.0 mm median standard length) blenniid Istiblennius zebra, while the blenniid Entomacrodus marmoratus and wrasses Thalassoma spp. were more common in low pools. Temperature was the best environmental predictor of assemblages and this relationship warrants further investigation. Our findings indicate that assemblages can vary across a narrow geographical range and intertidal shore.     

Patterns of organizations of intertidal and shallow subtidal vegetation in wave exposed habitats of central Chile

Wave-exposed rocky intertidal habitats of central Chile exhibit zonation of algal morphologies rather than strict patterns of species zonation. In low shore areas, there is a vertical sequence of perennial belts of calcareous crusts, kelp-like forms and expanded cushions or non-calcareous crusts. The calcareous crusts are represented by species of Mesophyllum, the kelp-like forms include Lessonia nigrescens and Durvillaea antarctica, while the cushions are represented by Gelidium chilense and G. lingulatum and the noncalcareous, expanded crusts by Codium dimorphum. Thin and thick blades, represented by Iridaea laminarioides, Ulva rigida and Porphyra columbia and filamentous forms including Ceramium rubrum, Centroceras clavulatum and Polysiphonia spp. are more patchy than the lower, perennial belts. They may, however, form distinct temporal monocultures at upper intertidal levels. Upper and lower limits of the various zones are set by interactions of several factors, the relative importance of which can change seasonally. When some of the factors restricting species distribution are experimentally removed, other interactions among factors become limiting.Within each zone, species are morphologically similar, with the abundance of species being regulated by symmetric competitive interactions. Competition is often asymmetric at the boundaries of zones except when adults of small-sized forms interact with morphologically similar juveniles of larger forms. Irrespective of their extremely different morphologies, the permanent, zone-forming algal species generally combine escape from grazers or defensive adaptations with clear competitive abilities. Nevertheless, there is a clear competitive hierarchy which is expressed in vertical displacements and zonation. The lowershore habitats could potentially be occupied by any of the different types of algae. Fast growth and large size allow the kelps to occupy this zone pushing the calcareous crust dominated-zone down into shallow subtidal areas and displacing the cushions and fleshy crusts into the low and middle intertidal regions. In turn, these last forms can displace thick and thin foliose forms and filaments to upper levels on the shore. Displaced forms may exist as patches at various levels of the shore.     

Intertidal zonation of prosobranch gastropods: Analysis of densities of four co-existing species

The densities of populations of Nerita atramentosa Reeve, Austrocochlea constricta Lamarck, Bembiciuin nanum (Lamarck), and Cellana tramoserica (Sowerby) were recorded in 0.25 m² quadrats on landward and seaward halves of a sheltered and a moderately exposed rock platform from June 1972 to June 1973. The data have been subjected to variance and multiple regression analysis. Differences between densities of each species on the two shores and in the two areas of each shore are discussed with respect to the regressions on littoral height. Densities of Nerita on the sheltered shore, Austrocochlea and Bembicium on both shores, and Cellana, on the exposed shore, increased with height. The density of Cellana on the sheltered shore decreased as height increased. The density differences in the landward and seaward areas on the two shores were attributable to the effect of height for Nerita and Austrocochlea. Height was the major variable affecting the density of these two species on the sheltered shore. Bembicium and Cellana densities on both shores, and Austrocochlea density on the exposed shore, are not primarily determined by height.On the sheltered shore, Nerita and Cellana increased in density with increasing cover of water in rock pools. On the exposed shore, Bembicium and Cellana became sparser with increasing water cover. Bembicium on both shores increased in density with increasing cover of the alga Peyssonelia, as did Cellana on the exposed shore. Cellana and Austrocochlea tended to increase in density with Nerita on the sheltered shore. Cellana tended to be sparse where Bembicium was dense on the exposed shore. The biological effects of substrata and other species on the densities of each species remain unexplained.Significant time effects were found for variations in the densities of Nerita and Cellana on the sheltered shore and of Austrocochlea and Cellana on the exposed shore. The seasonal change in density of Nerita was parabolic with a minimum in summer, and is consistent with the known period of recruitment of this species. The lack of significant time effects on density of Austrocochlea on the sheltered shore is consistent with the known continuous recruitment of this species. The declining density of Austrocochlea on the moderately exposed shore was found, by size-frequency analysis, to be due to reduced recruitment compared with surrounding shores. The density of Cellana on both shores showed seasonal changes, with a maximum in summer, which is consistent with the known period of recruitment in Cellana. The lack of seasonal changes in density of Bembicium is briefly discussed.This type of multifactor analysis identifies biological interactions affecting variation in density of these species. It also corroborates the results of other ecological investigations and determines the relative importance of a variety of physical and biological factors in the distribution of density of intertidal gastropods.     

The vertical distribution of Chthamalus montagui and Chthamalus stellatus (Crustacea,Cirripedia) in two areas of the NW Mediterranean Sea

The Mediterranean Sea is characterised by a small tidal range (0.3–1 m). Despite this, intertidal communities are well established and their upper limits often extend above mean high water level. Organisms living in the intertidal region and in the supralittoral zone rely on both tides and wave action to perform their biological functions. Lack of food, desiccation and predation are common stresses in such a harsh environment. The present study deals with the vertical distribution of two species of intertidal barnacles, Chthamalus montagui Southward and Chthamalus stellatus (Poli), which are the main constituents of the barnacle belt along Mediterranean rocky shores. Previous work, carried out in the Atlantic, showed that where the distribution ranges of the two Chthamalus species overlap, C. montagui is more common in the upper barnacle zone while C. stellatus is dominant lower down. The main aims of our study are: (1) to establish if there is a relationship between position and extension of the barnacle belt on the shore and tidal range and/or wave exposure, (2) to test the hypothesis that in the study areas C. montagui is more abundant than C. stellatus high on the shore, and that the pattern is reversed lower down. Barnacle populations were monitored in summer 1998 in the Gulf of Genoa (Ligurian Sea) and in the Gulf of Trieste (North-Adriatic Sea). The two areas differ in tidal range and hydrodynamism, the former presenting quite strong wave action and a tidal range of 30 cm, the latter having limited wave action and 1 m tidal range. Three shores were randomly selected in each gulf and two transects on each shore. Counts of barnacles in 10 ^* 10 cm quadrats were done at different shore heights along each transect. The data was subjected to analysis of variance. Results showed that a more pronounced hydrodynamic regime corresponded to a shift of the barnacle belt towards the higher shore (Gulf of Genoa), while in more sheltered areas (Gulf of Trieste), the barnacle distribution was confined to the intertidal region. The relative spatial distribution of C. montagui and C. stellatus within the barnacle belt varied locally, even between transects on the same shore, and this obscured the distribution pattern along the vertical gradient. Nevertheless, it was still possible to conclude that at mid and high shore in Genoa, C. stellatus was more abundant than C. montagui, while in Trieste the pattern was reversed.     

Causes and consequences of thermal tolerance limits in rocky intertidal porcelain crabs, genus petrolisthes

Vertical zonation of intertidal organisms, from the shallowsubtidal to the supralittoral zones, is a ubiquitous featureof temperate and tropical rocky shores. Organisms that livehigher on the shore experience larger daily and seasonal fluctuationsin microhabitat conditions, due to their greater exposure toterrestrial conditions during emersion. Comparative analysesof the adaptive linkage between physiological tolerance limitsand vertical distribution are the most powerful when the studyspecies are closely related and occur in discrete vertical zonesthroughout the intertidal range. Here, I summarize work on thephysiological tolerance limits of rocky intertidal zone porcelaincrab species of the genus Petrolisthes to emersion-related heatstress. In the eastern Pacific, Petrolisthes species live throughouttemperate and tropical regions, and are found in discrete verticalintertidal zones in each region. Whole organism thermal tolerancelimits of Petrolisthes species, and thermal limits of heartand nerve function reflect microhabitat conditions. Speciesliving higher in the intertidal zone are more eurythermal thanlow-intertidal congeners, tropical species have the highestthermal limits, and the differences in thermal tolerance betweenlow- and high-intertidal species is greatest for temperate crabs.Acclimation of thermal limits of high-intertidal species isrestricted as compared to low-intertidal species. Thus, becausethermal limits of high-intertidal species are near current habitattemperature maxima, global warming could most strongly impactintertidal species.     

Barnacle larval supply to sheltered rocky shores: a limiting factor?

In northwest Europe, sheltered rocky shores are dominated by fucoid canopy algae and barnacles are rare, although the latter are extremely abundant on exposed shores. The supply of the intertidal barnacle Semibalanus balanoides (L.) to sheltered, fucoid dominated rocky shores was investigated to determine the importance of larval supply in limiting the abundance of adults in shelter. Larval supply was measured at two spatial scales, at the scale of shore (100s of metres), by comparing larval concentrations at exposed and sheltered sites, and at a smaller spatial scale (m), by examining the role of fucoid canopies in limiting supply to the substratum. Replicate plankton trawls were carried out above the intertidal zone at high water at two sheltered sites and nearby exposed headlands. The concentration of S. balanoides cyprid larvae was significantly higher at the sheltered sites on two out of three sampling occasions with up to 14 times greater larvae on one occasion than the nearby exposed site. The effect of the macroalgal canopy on supply to the substratum was assessed in two ways: directly, by pumping water from the substratum in areas with and without Ascophyllum nodosum (L.) Le Jolis, and indirectly by measuring cyprid settlement in a canopy-manipulation experiment. Pumped plankton samples from mid tide level showed that the A. nodosum canopy did not form a barrier to larval supply and may have had a positive effect on larval concentrations at the substratum. Cyprid settlement was assessed in the mid shore A. nodosum and low shore Fucus serratus L. zones to areas with canopy algae (but protected from the sweeping effects of macroalgal fronds) and without canopy. Settlement over three consecutive 24-h periods showed a consistent pattern; settlement was consistently lower beneath the F. serratus canopy than in cleared areas, suggesting that this algal species forms a barrier, limiting supply of cyprid larvae to the substratum.     

Trends in abundance of rocky intertidal seaweeds and filter feeders across gradients of elevation,wave exposure,and ice scour in eastern Canada

On NW Atlantic rocky shores, the main basal organisms in intertidal communities are seaweeds (Ascophyllum nodosum, Fucus spp. and Chondrus crispus) and filter feeders (barnacles, Semibalanus balanoides, and mussels, Mytilus spp.). Their ecology has been extensively studied in New England (United States), but knowledge gaps exist for northern shores, which are subjected to stronger environmental stress. Therefore, we studied the above organisms on Canadian shores. We quantified the summer abundance of these seaweeds and filter feeders across full vertical (intertidal elevation) and horizontal (wave exposure and winter ice scour) environmental gradients on the Gulf of St. Lawrence and open Atlantic coasts of Nova Scotia. At the regional scale along the open Atlantic coast, seaweeds showed similar abundances in Nova Scotia than values reported for New England. However, both filter feeders were considerably less abundant in Nova Scotia. At the local scale in Nova Scotia, intense winter ice scour (which only occurs on the Gulf of St. Lawrence coast) was associated with a very low abundance of all species except barnacles. Spatial trends in Nova Scotia were similar to patterns known for certain species elsewhere, such as A. nodosum being almost restricted to sheltered habitats, regardless of elevation, and C. crispus being almost restricted to low elevations, regardless of exposure. Other trends were, however, characteristic of Nova Scotia, such as C. crispus being frequent at low elevations in exposed habitats, unlike in New England, where mussels often predominate there because of competitive advantages. In Nova Scotia, mussels were always restricted to cracks and crevices, unlike in New England, where they form extensive intertidal beds on exposed shores. The direct effects of increased environmental stress and indirect effects through altered interspecific interactions might explain the regional differences in local species distribution, which will require experimental validation. Handling editor: K. Martens     

Spatial analyses of intertidal assemblages on sheltered rocky shores

Abstract Understanding processes in complex assemblages depends on good understanding of spatial and temporal patterns of structure at various spatial scales. There has been little quantitative information about spatial patterns and natural temporal changes in intertidal assemblages on sheltered rocky shores in temperate Australia. Natural changes and responses to anthropogenic disturbances in these habitats cannot be accurately measured and assessed without quantitative data on patterns of natural variability in space and through time. This paper describes some suitable quantitative methods for examining spatial and temporal patterns of diversity and abundances of highshore, midshore and lowshore intertidal assemblages and the important component species for a number of shores in a bay that has not been severely altered by human disturbance. Despite a diverse flora and fauna on these shores, the midshore and lowshore assemblages on sheltered shores were characterized by a few species which were also the most important in discriminating among assemblages on a shore and, for each assemblage, among different shores. The same set of species was also important for measuring small-scale patchiness within each assemblage (i.e. between replicate sites on a shore). Therefore, these data provide a rationale for selecting species that are useful for measuring differences and changes in abundance among places and times at different scales and, hence, can be used in the more complex sampling designs necessary to detect environmental impacts. There was considerable spatial variability in all assemblages and all species (or taxa) examined at scales of metres, tens of metres and kilometres. There were no clear seasonal trends for most measures, with as much or more variability at intervals of 3 months as from year to year. Most interactions between spatial and temporal measures were at the smallest scale, with different sites on the same shore generally showing different changes from time to time. The cause(s) of this apparently idiosyncratic variability1 were not examined, but some potential causes are discussed. These data are appropriate for testing hypotheses about the applicability of these findings to other relatively undisturbed sheltered shores, about effects of different anthropogenic disturbances on sheltered intertidal assemblages and to test hypotheses about differences in intertidal assemblages on sheltered versus wave-exposed shores.     

An examination of species dispersion patterns along the intertidal gradient on Macquarie Island (sub-Antarctica) using a restricted occupancy model

It has been widely held that intertidal zonation boundaries and the tidal emersion levels are causally related: the critical tidal level hypothesis. In order to evaluate this hypothesis, the dispersion patterns of species boundaries on the intertidal rocky shores of Macquarie Island (sub-Antarctica) were examined using a restricted occupancy model to test the null hypothesis that the observed species were randomly dispersed along the tidal emersion gradient. Subsequently this investigation demonstrated that the intertidal species boundaries on Macquarie Island shores were randomly dispersed with respect to the tidal emersion gradient. Hence no prima facie evidence was found to support the critical tidal level hypothesis. This suggests that tidal emersion is not a significant factor structuring intertidal communities on Macquarie Island.     

Biotic zonation on rocky shores of Heard Island

Summary The zonation of rocky shore biota on Heard Island is described for the first time, related to a universal zonation scheme and compared with that recorded for other sub-Antarctic localities. The Kelp Zone of holdfasts of the giant kelp, Durvillea antarctica (Chamisso) Hariot (one of the characteristic features of these regions) is confirmed as a sublittoral fringe. The occurrence of a Bare Zone within the littoral zone on sub-Antarctic shores is discussed. The possible roles of submersion, spray, freezing and predation by gulls in influencing the extent and composition of zones are discussed.     

Dispersal mode and assessments of recovery on the shores of Gruinard,the ‘anthrax island’

Formaldehyde run-off was an unintended impact of the anthrax decontamination procedure on the island of Gruinard. The death of intertidal organisms was observed where formaldehyde reached the shore during 1986. The extent to which shores on Gruinard have recovered was assessed with survey work in 2000. Recovery estimates were based on the hypothesis that the process of recolonization is partly dependent on species’ dispersal capability. Underdevelopment of the assemblage of species lacking planktonic dispersal stages (direct developers) is therefore evidence that the process of recolonization is ongoing, rather than complete. A novel multivariate test showed that, when comparing quadrats from Gruinard and nearby mainland shores, assemblages of direct developing molluscs were significantly more distinct than assemblages of molluscs with planktonic dispersal stages. The average densities of species with direct development were generally lower on Gruinard than on mainland shores. While some species with direct development have similar densities on Gruinard and on the surrounding shores, the recovery of the overall assemblage was still incomplete after 14 years. In contrast, the harvested species, Littorina littorea, appeared to benefit from the absence of humans visiting Gruinard’s shores. Populations of L. littorea on Gruinard contained significantly higher proportions of large individuals. Depending on the dispersal capabilities of different species, Gruinard is either still in recovery or acts as a reserve.     

KINETICS OF NITRATE,AMMONIUM, AND UREA UPTAKE BY FOUR INTERTIDAL SEAWEEDS FROM NEW ZEALAND1

The competitive ability for N uptake by four intertidal seaweeds, Stictosiphonia arbuscula (Harvey) King et Puttock, Apophlaea lyallii Hook. f. et Harvey, Scytothamnus australis Hook. f. et Harvey, and Xiphophora gladiata (Labillardière) Montagne ex Harvey, from New Zealand is described by the uptake kinetics for NO₃^?, NH₄⁺, and urea. This is the first study to report uptake kinetics for N uptake by a range of southern hemisphere intertidal seaweeds in relation to season and zonation. Species growing at the highest shore positions had higher NO₃^? and urea uptake at both high and low concentrations and had unsaturable NH₄⁺ uptake in both summer and winter. Although there was evidence of some feedback inhibition of V_max for NO₃^? uptake by Stictosiphonia arbuscula growing at the lower vertical limits of its range, rates were high compared with species growing lower on the shore. Our results highlight the superior competitive ability for N uptake of certain high intertidal seaweeds, and consistent with our previous findings we can conclude that intertidal seaweeds in southeast New Zealand are adapted to maximizing N acquisition in a potentially N‐limiting environment.     

Nest site choice by the intertidal spider Desis formidabilis (Araneae: Desidae) and nest utilisation by its hymenopteran egg parasitoid

1. Echthrodesis lamorali Masner, 1968 is the only known parasitoid of the eggs of the intertidal rocky shore spider Desis formidabilis O.P. Cambridge 1890 and is endemic to a small area of South Africa. 2. The abundance of spider nests and parasitoid presence were assessed in relation to their in‐ and between‐shore location at multiple sites within the distribution of E. lamorali along the Cape Peninsula (Western Cape, South Africa). 3. Desis formidabilis nests were more abundant in the mid‐shore zone than higher up or lower down the shore. Spider population sizes also differed between collection sites, with higher numbers recorded on the cooler western coast of the peninsula. 4. Evidence of parasitoid activity was recorded in 43.31% of the 127 nests and 13.85% of the 592 egg sacs they contained. 5. Where parasitoids gained entry to a spider egg sac, oviposition took place into all of the eggs present. 6. Incidence of wasp activity was positively correlated with spider nest concentration, not with height up the shore, suggesting that both the host and parasitoid are tolerant of salt‐water inundation. 7. These results should assist managers of the Table Mountain National Park, in which the full distribution of E. lamorali falls, to better understand this component of rocky shore community dynamics.     

ASYMMETRICAL DEVELOPMENTAL PLASTICITY IN AN INTERTIDAL SNAIL

Field and laboratory experiments were used to investigate the variation and phenotypic plasticity in the adhesive abilities of the intertidal snail Nucella lapillus between high- and low-wave-energy environments. Whelks from an exposed coast produced a larger pedal surface area and were more resistant to dislodgement than were similar-sized individuals from a protected shore. Tenacity (g/cm²) was similar between individuals from exposed and protected shores, indicating that variation in resisting dislodgement was solely a function of pedal surface area. Whelks from exposed and protected shores did not differ in pedal surface area as they emerged from egg capsules or when reared in the laboratory under uniform conditions, suggesting that variation between populations does not represent genetic differentiation. Individuals from high-and low-wave-energy environments reared intertidally produced a larger pedal surface area than did those reared in the laboratory. The extent to which pedal surface area increased corresponded to the intensity of wave action. These findings suggest that pedal surface area is a highly plastic character modulated or induced by the water turbulence accompanying breaking waves. A reciprocal-transplant experiment confirmed this notion but revealed an asymmetry in the plasticity. Snails from the protected site transplanted to an exposed shore formed a much larger pedal surface area than did controls reared on the protected shore. In contrast, whelks from a wave-swept shore transplanted to a protected shore differed little from their controls reared on the exposed shore. The asymmetric response parallels a possible asymmetry in the risks of acclimating to a temporally unpredictable environmental cue, such as wave action.