Effects of interactions between algae and grazing gastropods on the structure of a low-shore intertidal algal community

Summary At low levels on shores in New South Wales, foliose algae are abundant and often occupy all substrata; microalgal grazing gastropods are rare or absent. At higher levels, foliose algae are sparse or absent and grazing gastropods are abundant. Hypotheses for the causes of the lower vertical limits of distribution of these grazers include the effects of increased predation or the deleterious physiological effects of increased period of submergence at lower levels on the shore. Alternatively, the presence of the algae, because they occupy space and deprive the grazers of substratum for feeding, may prevent the downward movement, or survival of the grazers at low levels. Under the first two of these hypotheses, algae are able to colonize and grow in low-shore areas as an indirect result of factors which remove grazers. Under the third hypothesis, the algae are directly responsible for the lack of grazers.Experimental clearings of the low-shore algae and introductions of the mid-shore limpets Cellana tramoserica and Siphonaria denticulata were used to test these hypotheses. C. tramoserica grazes microalgae and removes them from the substratum, preventing colonization. S. denticulata, in contrust, crops the algae, leaving a visible cover of algae on the substratum, which can grow rapidly. Because of its method of feeding, S. denticulata had no measurable impact on the rates of colonization, nor on the dry weights of algae, compared with those of ungrazed areas. C. tramoserica could keep cleared areas tree from foliose algae, but only when the limpets were mainfained in great density (10 per 900 cm²). They were less effective where wave-action was greater.Neither species of limpets could survive when placed onto beds of mature algae, because they had no substratum on which to cling and were swept away by the waves. C. tramoserica did not invade clearings below their lower limit of distribution where they had to move over a bed of foliose algae. Few C. tramoserica moved directly downshore into cleared areas. When placed on bare rock within low-shore beds of algae of different ages, S. denticulata remained amongst the algae and maintained their tissue-weights. Few C. tramoserica remained in areas with well-developed algae, compared with areas having sparse algal growth. Those Cellana which remained amongst well-developed algae lost weight, whereas limpets in areas with less algal growth mammtained their weights. In experimental cages in low-shore beds of algae, where the limpets were inaccessible to potential predators, C. tramoserica lost weight and died. On cleared areas they survived for many weeks, but lost weight and died as algae grew and covered the substratum. In the absence of predation, the micro-algal grazer C. tramoserica could not survive in lowshore areas because algae grew too fast and occupied the substratum, making it inaccessible for the limpets to graze; the algae, once grown beyond small sporelings, are not a suitable food-source for C. tramoserica, and the loss of weight and death of these limpets is attributable to starvation.The lower limit of distribution of C. tramoserica is not due to the direct effects of physical factors associated with prolonged submersion, nor to the impact of predators, but is apparently determined by the presence of rapidly growing, extensive beds of foliose algae at low levels on the shore. The cause of the limit of distribution of S. denticulata is not yet known and predation may prove to be important. Removal of S. denticulata from low-shore algal beds would not, however, affect the domination of substrata by algae. Grazing by S. denticulata at very great density had no effect on algal cover nor weight. In the intertidal community studied, the persistence of a low-shore algal zone, bounded above by abundant grazers is not influenced by the activities of predators, but is a direct result of interactions between the grazers and the algae.     

Spatial distribution of algae and invertebrates in the rocky intertidal zone of the Strait of Magellan: are patterns general?

Spatial variability in the distribution of macroalgae and invertebrates was examined at a number of sites in the Strait of Magellan. Two main predictions of models of zonation were tested in this study: (1) consistency in patterns of vertical distribution among sites one to tens of kilometers apart, and (2) homogeneity between areas at the same level on the shore, tens to hundreds of meters apart. Two types of habitat were considered: continuous rocky shores and blocks of rock in gravel beaches. In the former habitat, percent cover estimates of sessile organisms (algae and mussels) were obtained from three replicated plots (0.5 × 0.5 m) sampled non-destructively in each of three random areas (stretches of shore about 20 m long) at each of three levels on the shore: high, middle and low intertidal zone. This procedure was repeated at six different sites along the Chilean coast of the Strait of Magellan. Blocks of rock were sampled at two sites about 1 km apart. Estimates of the abundance of sessile (barnacles and mussels) and mobile (limpets) organisms were obtained for six blocks at each of two levels on the shore (high and low intertidal zone) at each site. At each level on the shore, three blocks were sampled on the top and three on the vertical sides. A single plot was sampled on each block. This design allowed a test of the null hypothesis of no interactive effects between position on the blocks and level on the shore. Both the predictions were falsified: (1) there were large between-site differences in the vertical structure of assemblages and (2) variability between areas at the same level on the shore was large for some of the most common algae. In contrast, the null hypothesis of no interactive effects of position on the blocks and level on the shore was retained. The results of this study show that vertical position on the shore alone is not a good predictor of the structure of assemblages of benthic organisms in the rocky intertidal of the Strait of Magellan. Received: 9 December 1996 / Accepted: 2 May 1997     

Grazing by two species of limpets on artificial reefs in the northwest Mediterranean

The extensive presence of artificial reefs in marine coastal habitats demands a better understanding of the extent to which these structures can be considered surrogates of natural rocky shores for populations of plants and animals. The primary aim of this study was to test the hypothesis that removing limpets from the midlittoral of artificial breakwaters in the northwest Mediterranean led to changes in assemblages similar to those observed on rocky shores in the same area. Orthogonal combinations of the presence/absence of two species of limpets, P. aspera and P rustica, were produced using manual removals from June 1997 to February 1998. To test the hypothesis that the effects of limpets were variable at spatial scales comparable to those investigated on rocky shores, we repeated the experiment at two locations tens of kilometres apart, and on two reefs within each location a few kilometres apart. The results revealed strong and relatively consistent negative effects of limpets on filamentous algae, whereas interactions with other members of assemblages were complex and variable. Several taxa (Cyanophyta, encrusting and articulated coralline algae, Ralfsia and Rissoella) were abundant at one location but nearly absent at the other. This large-scale variability in patterns of distribution generated inconsistencies in the effects of limpets between locations. Within locations, several effects of P. aspera and P. rustica were observed, ranging from independent effects on some organisms, to additive or interactive effects on others. Apparently, the removal of filamentous algae by limpets resulted in positive indirect effects on Ralfsia and Rissoella. Collectively, these effects were comparable to those described for rocky shores in the northwest Mediterranean. The processes accounting for large-scale variation in grazing, however, appeared different between the natural and the artificial habitat.     

Morphological and behavioral evidence for adaptive diversification of sympatric Hawaiian limpets (Cellana spp.)

The endemic Hawaiian limpets (Cellana exarata, Cellana sandwicensis, and Cellana talcosa), reside at different elevations on wave-exposed rocky shores and comprise a monophyletic lineage that diversified within Hawai'i. Here, I report phenotypic differences in shell, soft tissue, and behavioral characters among these limpets and discuss their potential utility in exploiting their respective niches. The high-shore limpet, C. exarata, is characterized by a tall round shell, short mantle tentacles, and long evasion distance when confronted by a predatory gastropod. The mid-shore limpet, C. sandwicensis, is characterized by a shorter oblong shell, long mantle tentacles, and a short evasion distance when confronted by a predatory snail. The low-shore, shallow-subtidal limpet, C. talcosa, is characterized by a flat shell that is thin in juveniles and disproportionately massive in large adults (relative to the other two species), and mantle tentacles of varying lengths (some individuals exhibit short tentacles, some long). These species-specific suites of characters are likely to confer specific fitness advantages on the high shore (C. exarata) where thermal and desiccation stress is severe, on the mid shore (C. sandwicensis) where hydrodynamic forces are severe, and on the low-shallow subtidal shore (C. talcosa) where pelagic predators have free access to the limpets. These data add to the growing body of evidence for adaptive diversification and speciation in the Hawaiian Cellana, and in marine species in general.     

The effects of grazing on the distribution and composition of low-shore algal communities on the central coast of Portugal and on the southern coast of Britain

A red algal turf is often found just below the barnacle/limpet zone of many European shores, especially on steep shores of moderate exposure. The hypothesis that grazing by limpets determines the upper limit of distribution of this red algal turf was tested on moderately exposed shores in Portugal and Britain. We also aimed to assess whether the grazing effect is modified at various spatial scales. Grazers were excluded by fences, with half-fenced and unfenced controls. Exclusion plots were rapidly colonised by green ephemeral algae in the months immediately after the beginning of the experiment (summer); these algae were later replaced by perennial algae. The percentage cover of turf-forming macroalgae showed a significant increase at both locations. The upper limit of distribution extended more than 50 cm on most of the shores studied. In contrast, control and half-fenced plots remained devoid of algae. After 2 years, ungrazed plots were mainly colonised by a red algal turf (e.g. Caulacanthus ustulatus, Gelidium spp., Laurencia pinnatifida) in Portugal, while canopy cover (Fucus serratus and Himanthalia elongata) dominated in Britain in marked contrast to the grazed plots. Physical factors acting at both local and geographical scales may explain these differences. However, although physical factors probably have an important influence on the identity, size and abundance of sublittoral fringe macroalgae, grazers play a major role in directly setting their upper limits. The effect of grazing by limpets was not consistent for all of the morphological algal groups and spatial scales considered in the present study. The effect of grazing on the cover of turf algae varied between Portugal and Britain (location scale), while effects on ephemeral and canopy algal cover varied at the shore scale within location.     

Responses of the limpet, Cellana grata (Gould 1859), to hypo-osmotic stress during simulated tropical, monsoon rains

Although heat stress is often cited as the dominant physical stress on tropical shores, intertidal organisms in regions with monsoonal climates are also regularly exposed to prolonged periods of heavy rainfall. Such events are predicted to have adverse physiological effects on individuals and may result in mortality. In a series of laboratory experiments, the impact of simulated monsoonal rains was investigated on the patellid limpet, Cellana grata. Sub-lethal responses in terms of body water content, body fluid osmolality and heart rate were measured in two different size cohorts maintained on horizontal and vertical substrata. Limpets were unable to achieve any effective behavioural isolation, and exposure to either simulated rainfall or diluted seawater resulted in both large and small C. grata gaining water with subsequent dilution of mantle water and haemolymph osmolalities. With increased duration of rainfall, dilution of body fluids increased with little difference between individuals on horizontal and vertical surfaces. Body fluids generally showed proportional dilution during prolonged rain, but in some individuals there was evidence for regulation of the haemolymph relative to the mantle fluid. Overall, smaller limpets were more susceptible to prolonged rainfall than large animals in terms of swelling of soft tissues and detachment and also had higher heart rates than large limpets. Both cohorts reduced heart rates with prolonged rainfall, suggesting a degree of metabolic depression, especially on horizontal surfaces. In small limpets, no difference in heart rate was found with substratum orientation, whereas large limpets had elevated heart rates on vertical as compared to horizontal substrata, when exposed to either simulated rainfall or washed with dilute seawater. This may reflect the increased energetic costs required to maintain a relatively larger body on a vertical surface under stressful conditions. Monsoonal rainfall during emersion, and subsequent dilution of seawater, therefore, have sub-lethal physiological and possible lethal effects on intertidal limpets. This influence has been largely overlooked, but coupled with the possible synergistic effects of thermal stress, monsoon rains are likely to play an important role in community dynamics on tropical shores.     

The influence of canopy algae on vertical patterns of distribution of low-shore assemblages on rocky coasts in the northwest Mediterranean

Canopy-forming algae have been shown to play a fundamental role in the maintenance of understorey assemblages on rocky shores. In the Mediterranean, low-shore habitats are often monopolised by canopy algae of the genus Cystoseira and, in particular, by the species Cystoseira amentacea var. stricta. Alternatively, low-shore habitats are dominated by turf-forming algae and/or mussels. Previous studies showed that loss of Cystoseira, by natural or anthropogenic disturbances, resulted in the increase of turfing algae. Here, we propose that turf-forming algae may act as colonisers from nearby habitats, specifically from lower portions of the shore. The following hypotheses were tested to investigate this proposition: (1) in presence of Cystoseira, the assemblage living under its canopy will be distinct, in terms of composition and structure, from that found lower on the shore, (2) if the canopy of Cystoseira was removed, the differences between the two assemblages would decrease or disappear, (3) the effects of the canopy would be more important than other causes of variability at the spatial scale investigated (hundreds to thousands of metres), and (4) that effects of removal of the canopy would be consistent through time. These hypotheses were tested by means of a canopy removal experiment, involving several spatial and temporal repetitions of the manipulation.The assemblage underneath the canopy of Cystoseira was distinct from that found immediately lower on the shore, but when the canopy was removed there were no differences between the two. The effects of the canopy were consistent at the spatial and temporal scales investigated. Several species living under the canopy disappeared and were replaced by turf-forming species, resulting in a homogenisation of the two habitats and in a drastic loss of habitat diversity. This study shows that Cystoseira can be considered a habitat-forming species, responsible for the maintenance of two distinct low-shore assemblages. In this light, its importance should be taken into account by policies aiming to preserve biodiversity on rocky shores.     

LINKING BENTHIC ALGAL BIOMASS TO STREAM SUBSTRATUM TOPOGRAPHY1

The physical properties of substrata significantly influence benthic algal development. We explored the relationships among substratum surface texture and orientation with epilithic microphytobenthic biomass accumulation at the whole‐substratum and micrometer scales. Unglazed clay tiles set at three orientations (horizontal, vertical, and 45°), and six substrata of varying surface roughness were deployed in a prairie stream for 3 weeks. Substrata were analyzed for loosely attached, adnate, and total benthic algal biomass as chl a, and confocal laser scanning microscopy was used to measure substrata microtopography (i.e., roughness, microscale slope angles, and three‐dimensional surface area). At the whole‐substratum level, vertical substrata collected significantly (P < 0.05) less algal biomass, averaging 34% and 36% less than horizontal and 45° substrata, respectively. Benthic algal biomass was also significantly less on smoother surfaces; glass averaged 29% less biomass than stream rocks. At the microscale level, benthic algal biomass was the greatest at intermediate values, peaking at a mean roughness of approximately 17 μm, a mean microscale slope of 50°, and a projected/areal surface area ratio of 2:1. The proportion of adnate algae increased with surface roughness (26% and 67% for glass and brick, respectively), suggesting that substratum type changes the efficiency of algal removal by brushing. Individual substrata and microsubstrata characteristics can have a strong effect on benthic algae development and potentially affect reach scale algal variability as mediated by geomorphology.     

A continental scale evaluation of the role of limpet grazing on rocky shores

It is critical for our knowledge of biodiversity and ecosystem processes to understand how individual species contribute to ecosystem processes and how these contributions vary in space and time. We used a manipulative field experiment in five locations over 17° of latitude [from southern Portugal to the Isle of Man (British Isles)] to determine the relative response of rocky intertidal algal assemblages released from control by the grazing of limpets. Response ratios showed that when limpets were removed there was a trend of effects from north to south. In the north, grazing had a strong effect on algal assemblages, but removing grazers reduced spatial variability in assemblages. In the south, the effect of limpet grazing was far weaker and removal of grazers had a much reduced impact on spatial variability. Here we show a clear trophic control of an ecosystem in that grazing by limpets not only determines macroalgal abundance overall but also modifies ecosystem stability via variability in cover of algae.     

Effects of Chiton granosus (Frembly, 1827) and other molluscan grazers on algal succession in wave exposed mid-intertidal rocky shores of central Chile

Molluscan grazers can have important effects on the abundance, colonization rates, and successional pathways of algal assemblages and the entire intertidal community. In general, early successional algae are more readily consumed than corticated algae and kelps, which usually get established later in the community succession. To generalize, however, the effect of different grazers on algal assemblages must be examined on different coasts and under different scenarios. This information could help us understand the mechanisms of ecosystem processes and situations in which general models do not apply. Along the coast of Chile, humans harvest large keyhole limpets, which seem to be the only invertebrate grazers capable of controlling the dominant corticated alga Mazzaella laminarioides, a canopy-forming species that can cover extensive areas of the mid intertidal zone. In this scenario, where large limpets are harvested, the overall effects of the diverse molluscan assemblage of limpets, chitons and snails on algal succession and on corticated algae in particular are not clear. We conducted a 26-month-long experiment to evaluate the effects of molluscan grazers on mid-intertidal algal succession and to isolate the effects of Chiton granosus, the most conspicuous member of the assemblage at these tidal elevations. At sites heavily impacted by humans the molluscan grazer assemblage had strong negative effects on colonization and abundance of green algae such as ulvoids and Blidingia minima. In doing so, the grazer assemblage had a strong negative indirect effect on the establishments of chironomid fly larvae, which were only observed on green algal mats and rarely on bare rock. No significant effects were detected on epilithic microalgae, and effects on sessile invertebrates were highly variable over space and time. C. granosus also had significant negative effects on green algae but did not account for the total grazing pressure exerted by the guild. Limited foraging excursions (ca. 35 cm) from refuges and moderate site (crevice) fidelity in this species may contribute to the patchiness in green algal distribution observed in the field. Nearly 13 months after rock surface were experimentally cleared, M. laminarioides appeared in all experimental plots, but increased over three times faster in enclosures containing C. granosus than in exclosures plots or controls, suggesting that moderate levels of herbivory could actually facilitate the establishment of this alga in the succession and that the green algal cover found in the absence of grazers may delay its establishment.     

Disturbance and predation in an assemblage of herbivorous Diptera and algae on rocky shores

Summary Speculation about the effects of disturbance in marine benthic communities is often based on competition theory. Disturbances are thought to provision numerically depleted or competitively inferior species with resources associated with open substrate. However, disturbances that remove entire assemblages of sessile species also alter trophic structure, and thereby, influence the outcome of predator/prey relations. Aspects of community structure may be determined by patterns of disturbance and predation.The influence of disturbance and predation on the distribution and seasonality of blooms of ephemeral algae and associated Diptera was investigated with field experiments at several rocky beaches in central California. Blooms of ephemeral algae developed on high intertidal rock faces that were subject to severe seasonal disturbances caused by shifting sediment. These were subsequently colonized by the herbivorous larvae of several Diptera species for predictable periods each year. Other areas, without blooms, were not so disturbed.Experiments were done to determine if seasonal blooms were caused by seasonal disturbances that remove predators which otherwise might prevent the establishment of the Diptera/algae assemblage. The predators were crabs and limpets which eat both algae and larvae while foraging. Blooms of algae and larvae did not develop when limpets were transplanted to disturbed areas in periods between disturbances. Adjacent control areas did support blooms. Transplanted limpets did not survive periods of burial. When both limpets and crabs were excluded from treatment plots in undisturbed areas, blooms developed where they would not otherwise have occurred; controls remaied unchanged. Crabs and limpets differed in their effects on this assemblage. Crabs recruited quickly to the site of a bloom, but did not crop algal cover as closely, nor decrease larval density as much as the slowly recruiting limpets.The results suggest that disturbances favor blooms of some species by reducing predation. Severe localized disturbances increased the variability of the upper shore community by creating a patchwork of differing predator/prey abundances.     

Substratum‐dependent responses of ciliate assemblages to temperature: a natural experiment in Icelandic streams

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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.     

Experimental manipulation of shell color and morphology of the limpets Lottia asmi (Middendorff) and Lottia digitalis (Rathke) (Mollusca: Patellogastropoda)

We experimentally show that substratum can determine an individual patellogastropod's color and morphology. Lottia asmi (Middendorff), previously considered a high-domed, black stenotopic species on trochid gastropods, is similar to other northeastern Pacific limpet species with low profile and tessellated color patterns, when it is translocated to rock substrata. Lottia digitalis (Rathke), a highly variable species with at least two ecological variants that have been considered to be genetically determined, changes between these two forms when translocated from stalked barnacles to rocks. Color changes because the ingested pigments (algal, periostracal, etc.) and other materials (e.g., calcium carbonate) vary between substrata; morphological changes occur as responses to the topographical complexity and constraints of substrata. The geographical distribution of ecological variants of polytypic limpet species, and the ability to achieve cryptic coloration, may be determined by the geographic distributions of algae on which the limpets feed rather than by intrinsic characteristics of the limpets.     

Feast or famine for intertidal grazing molluscs: a mis-match between seasonal variations in grazing intensity and the abundance of microbial resources

Distinct seasonal variations in the abundance of photosynthetic microbiota and limpet grazing intensity were recorded at Port St Mary, Isle of Man between January 1994 and June 1996. Microbial abundance was negatively correlated with insolation stress, while grazing intensity was positively correlated with sea and air temperature. These patterns result in a mis-match between the supply of and the demand for microbial resources with maximal grazing intensity during the summer and autumn, but maximal microbial standing stock during the winter and early spring. The importance of top-down control of microbial assemblages by grazing was demonstrated by experimental exclusion of limpets during autumn 1993. This resulted in a four-fold increase in the abundance of cyanobacteria within 6 days, followed by a more gradual proliferation of ephemeral algae during the next 4 weeks. The abundance of diatoms remained relatively constant and was not influenced by the removal of grazers at this time of year. The influence of microbial resource availability on the growth and mortality of limpets was examined using experimental enclosures of differing densities of either Patella vulgata or P. depressa. After 6 months, there were significant relationships between grazer density and both mortality and growth with increased mortality and reduced growth for P. vulgata at increased densities, and reduced growth for P. depressa at increased densities. Hence, the availability of microbial resources may also influence the biomass of grazers on rocky shores from the bottom upwards. A conceptual model is presented which describes seasonal and annual variations in microbial resources and grazing intensity and their potential consequences for other shore dwellers.     

Population regulation in the intertidal limpet Patelloida alticostata (Angas, 1865)

Summary Subpopulations of the limpet, Patelloida alticostata, converged slowly toward a density of about 75 per linear meter of vertical rocky shore two years after experimental alteration of densities of adult animals. The changes in number of limpets in subpopulations occurred because large limpets suffered considerable mortality, while recruits experienced almost none. Neither migration, cannibalism, selection of settlement site by the limpets, nor predation by a whelk, acted in a density-dependent manner.Growth rates were density-dependent, and juveniles and recruits reached a much larger size in the absence than in the presence of adult limpets. Increasing the density of adult limpets did not increase the extent of grazing areas and subpopulations of different size compositions utilized similar grazing areas. Food may have been in short supply.These limpets have a great deal of population inertia, accommodating sporadic good recruitment by compensations in growth rates.     

Distribution and abundance of the acmaeid limpet,Patelloida latistrigata,and its interaction with barnacles

Summary At Cape Banks, New South Wales, adults of the small intertidal limpet, Patelloida latistrigata occur exclusively in the barnacle zone, and are primarily associated with the barnacle, Tesseropora rosea. Limpet density increases with barnacle density. Juvenile limpets can be found throughout the barnacle zone, and on patches of bare rock that may be temporarily available at lower levels on the shore. The failure of juveniles to survive and grow in places other than among Tesseropora is due to a combination of factors. These include desication at high levels on the shore, smothering by rapidly growing algae low on the shore, and the grazing activities of the larger limpet, Cellana tramoserica. These latter two factors also reduce the survival of experimentally transplanted adult Patelloida: algae by covering the substratum and smothering the limpets, and Cellana by outcompeting them for food. The density of Cellana is greater on patches of bare rock than among barnacles, and these large limpets may be unable to move and feed effectively over the irregular surface created by Tesseropora. Patelloida, however, is small enough to feed over and among these barnacles, and hence has a refuge from competition with Cellana. Barnacles may also provide shelter from the effects of desiccation and strong wave action, and thus increase the survival of juvenile Patelloida. By being associated with barnacles, however, Patelloida becomes vulnerable to intermittent predation by the whelk, Morula marginalba. This effect may be serious enough to eliminate small local populations of limpets, either by direct predation or by removing the refuge-providing barnacles. The association with barnacles may also limit the maximum size to which Patelloida can grow. Patelloida is not always found with Tesseropora, and adults in different localities can be found in association with other sessile organisms. It may be argued that small species of limpets require a spatial refuge from physical and/or biological pressures. To examine this hypothesis, the relationship between Patelloida and Tesseropora is compared to other published accounts of limpets with specialised modes of life.     

The effect of substratum type, orientation and depth on the development of bacterial deep-sea biofilm communities grown on artificial substrata deployed in the Eastern Mediterranean

An increasing number of deep-sea studies have highlighted the importance of deep-sea biofouling, especially in relation to the protection of deep-sea instruments. In this study, the microbial communities developed on different substrata (titanium, aluminum, limestone, shale and neutrino telescope glass) exposed for 155 days at different depths (1500?m, 2500?m, 3500?m and 4500 m) and positions (vertical and horizontal) in the Eastern Mediterranean Deep Sea were compared. Replicated biofilm samples were analyzed using a Terminal Restriction Fragment Length Polymorphisms (T-RFLP) method. The restriction enzymes CfoI and RsaI produced similar total numbers (94, 93) of different T-RFLP peaks (T-RFs) along the vertical transect. In contrast, the mean total T-RF number between each sample according to substratum type and depth was higher in more samples when CfoI was used. The total species richness (S) of the bacterial communities differed significantly between the substrata, and depended on the orientation of each substratum within one depth and throughout the water column (ANOVA). T-RFLP analyses using the Jaccard similarity index showed that within one depth layer, the composition of microbial communities on different substrata was different and highly altered among communities developed on the same substratum but exposed to fouling at different depths. Based on Multidimensional Scaling Analyses (MDS), the study suggests that depth plays an important role in the composition of deep-sea biofouling communities, while substratum type and orientation of substrata throughout the water column are less important. To the authors' knowledge, this is the first study of biofilm development in deep waters, in relation to the effects of substratum type, orientation and depth.     

Changes in habitat complexity negatively affect diverse gastropod assemblages in coralline algal turf

The physical structure of a habitat generally has a strong influence on the diversity and abundance of associated organisms. I investigated the role of coralline algal turf structure in determining spatial variation of gastropod assemblages at different tidal heights of a rocky shore near Sydney, Australia. The structural characteristics of algal turf tested were frond density (or structural complexity) and frond length (the vertical scale over which structural complexity was measured). This definition of structural complexity assumes that complexity of the habitat increases with increasing frond density. While frond length was unrelated to gastropod community structure, I found significant correlations between density of fronds and multivariate and univariate measures of gastropod assemblages, indicating the importance of structural complexity. In contrast to previous studies, here there were negative relationships between the density of fronds and the richness and abundance of gastropods. Artificial habitat mimics were used to manipulate the density of fronds to test the hypothesis that increasing algal structural complexity decreases the richness and abundance of gastropods. As predicted, there were significantly more species of gastropods in loosely packed than in tightly packed turf at both low- and mid-shore levels. Despite large differences between gastropod assemblages at different tidal heights, the direction and magnitude of these negative effects were similar at low- and mid-shore levels and, therefore, relatively independent of local environmental conditions. These novel results extend our previous understanding of the ecological effects of habitat structure because they demonstrate possible limitations of commonly used definitions of structural complexity, as well as distinct upper thresholds in the relationship between structural complexity and faunal species richness.     

Aerial and underwater metabolism of <Emphasis Type="Italic">Patella vulgata</Emphasis> L.: comparison of three intertidal levels

Intertidal molluscs are known to possess specific respiratory organs that permit aerial breathing during emersion. Patella vulgata is a widely distributed intertidal species found from low-water spring tide to high-water neap tidal level. In order to determine metabolic adaptations to habitat, carbon fluxes associated with respiration and calcification of P. vulgata living at high-shore, middle-shore and low-shore levels were compared. Seasonal aerial respiration was measured using an infrared gas analyser; seasonal underwater respiration and calcification were calculated from dissolved inorganic carbon and total alkalinity. P. vulgata showed net CaCO₃ deposition at all seasons, although the high-shore level limpet annual calcification rate was relatively low due to longer air exposure. Both aerial and underwater respiration rates were highly correlated with seasonal temperature variations and followed the vertical shore gradient, with stronger fluxes for low-shore tidal level limpets and lower fluxes for high-shore level limpets that must limit energy expenditure. P. vulgata appears to be well adapted to aerial exposure, with average hourly respiration fluxes stronger in air than in water. This study demonstrates that P. vulgata calcification and respiration are reduced in upper shore levels and are important factors determining the upper distribution limit of the species.