The ecological impacts of marine debris: unraveling the demonstrated evidence from what is perceived.

Anthropogenic debris contaminates marine habitats globally, leading to several perceived ecological impacts. Here, we critically and systematically review the literature regarding impacts of debris from several scientific fields to understand the weight of evidence regarding the ecological impacts of marine debris. We quantified perceived and demonstrated impacts across several levels of biological organization that make up the ecosystem and found 366 perceived threats of debris across all levels. Two hundred and ninety-six of these perceived threats were tested, 83% of which were demonstrated. The majority (82%) of demonstrated impacts were due to plastic, relative to other materials (e.g., metals, glass) and largely (89%) at suborganismal levels (e.g., molecular, cellular, tissue). The remaining impacts, demonstrated at higher levels of organization (i.e., death to individual organisms, changes in assemblages), were largely due to plastic marine debris (> 1 mm; e.g., rope, straws, and fragments). Thus, we show evidence of ecological impacts from marine debris, but conclude that the quantity and quality of research requires improvement to allow the risk of ecological impacts of marine debris to be determined with precision. Still, our systematic review suggests that sufficient evidence exists for decision makers to begin to mitigate problematic plastic debris now, to avoid risk of irreversible harm.

Linking effects of anthropogenic debris to ecological impacts.

Accelerated contamination of habitats with debris has caused increased effort to determine ecological impacts. Strikingly, most work on organisms focuses on sublethal responses to plastic debris. This is controversial because (i) researchers have ignored medical insights about the mechanisms that link effects of debris across lower levels of biological organization to disease and mortality, and (ii) debris is considered non-hazardous by policy-makers, possibly because individuals can be injured or removed from populations and assemblages without ecological impacts. We reviewed the mechanisms that link effects of debris across lower levels of biological organization to assemblages and populations. Using plastic, we show microplastics reduce the 'health', feeding, growth and survival of ecosystem engineers. Larger debris alters assemblages because fishing-gear and tyres kill animals and damage habitat-forming plants, and because floating bottles facilitate recruitment and survival of novel taxa. Where ecological linkages are not known, we show how to establish hypothetical links by synthesizing studies to assess the likelihood of impacts. We also consider how population models examine ecological linkages and guide management of ecological impacts. We show that by focusing on linkages to ecological impacts rather than the presence of debris and its sublethal impacts, we could reduce threats posed by debris.

Effects of zinc on microalgal biofilms in intertidal and subtidal habitats.

Microalgal biofilms are sensitive to environmental conditions. Impacts of contaminants on assemblages of marine biofilm are often investigated in laboratories or in mesocosms. Such experiments are rarely representative of the effects of contaminants on biofilms under natural conditions. Studies in field situations, with enough power to detect impacts, are necessary to develop a better understanding of the effects of contaminants on ecological processes. Metals are a common contaminant of marine systems and can cause disturbances to assemblages. Using a new technique to experimentally deliver contaminants to microalgal assemblages, hypotheses were tested regarding the effects of zinc on microalgal biofilms growing on settlement panels in subtidal and intertidal habitats. PAM fluorometry was used to assess the amount and physiological state of biofilms on panels. Control panels deployed for 1 month in each habitat had significantly greater amounts of biofilm than those exposed to zinc. After deployment for 3 months, the results varied with location. The observed effects on the biofilm did not, however, cause significant changes in the macro-invertebrate assemblages that developed on the panels.

Urban structures provide new insights into interactions among grazers and habitat.

A major challenge in ecology is to understand and predict consequences of environmental changes to biological assemblages. Urbanization and associated alteration and destruction of habitat cause profound changes in local biodiversity and the ecology of and interactions among organisms. This study tested hypotheses about interactions among intertidal species on urbanized shores to examine predictions from their known ecology. On natural shores, grazing limpets, Cellana tramoserica, outcompete the smaller limpets, Patelloida latistrigata. The latter shelter among barnacles. On seawalls in Sydney Harbour (Australia), P. latistrigata were found among and on oysters. C. tramoserica were found directly on walls where there were no oysters. To explain these observations, several models were proposed and tested by manipulative experiments in the field: (1) oysters provide habitat for P. latistrigata; (2) negative interactions with C. tramoserica cause P. latistrigata to move on and among oysters; and (3) in areas with oysters, space available for grazing is insufficient for C. tramoserica to survive well. The results showed that C. tramoserica had a negative impact on the survival of P. latistrigata, although not causing the latter to move onto oysters. The oysters directly and indirectly affected P. latistrigata. First, oysters provided habitat and increased survival of P. latistrigata. Second, oysters influenced the interaction between C. tramoserica and P. latistrigata. This "interaction modification" diminished the negative impact of C. tramoserica. The material used to construct a wall did not directly influence the distribution of the limpets. Maintaining oysters on seawalls is, however, important because of their direct and indirect effects on P. latistrigata. It was possible to predict some processes on urbanized shores from known ecology. Other processes could not be predicted, making it necessary to do experiments to understand how built structures influence biological assemblages. This research contributes to understanding how to conserve biodiversity in urban areas.

Independent effects of patch size and structural complexity on diversity of benthic macroinvertebrates.

Despite a long history of work on relationships between area and number of species, the details of mechanisms causing patterns have eluded ecologists. The general principle that the number of species increases with the area sampled is often attributed to a sampling artifact due to larger areas containing greater numbers of individuals. We manipulated the patch size and surface area of experimental mimics of macro-algae to test several models that can explain the relationship between abundance and species richness of assemblages colonizing different habitats. Our results show that patch size and structural complexity have independent effects on assemblages of macroinvertebrates. Regardless of their structural complexity, larger habitats were colonized by more species. Patch size did not have a significant effect on numbers of individuals, so the increased number of species in larger habitats was not simply a result of random placement associated with sampling increased number of individuals. Similarly, random placement alone could not explain differences in numbers of species among habitats with different structural complexity, contrary to suggestions that the relationship between number of species and surface area might also be a sampling artifact due to more complex habitats having larger areas and therefore sampling more individuals. Future progress would benefit from manipulating properties of habitat in conjunction with experimental manipulations of area.

Do modified habitats have direct or indirect effects on epifauna?

Replacing natural habitats with artificial structures such as pier-pilings, jetties, and seawalls has important consequences to abundances of biota. It is, however, not often known whether these are direct (the novel habitat alters abundances of some species) or indirect (the novel habitat directly alters some aspect of the behavior or ecology of some species, which, in turn, alter abundances of other species). Marine animals in some modified habitats in Sydney Harbour provide experimental opportunities to test hypotheses to distinguish between direct and indirect processes. Covers of bryozoans and hydroids were greater on kelp growing on pilings than on kelp growing on natural reefs. The epifauna may be affected directly by the pilings or indirectly, i.e., the structure affects characteristics of the kelp which, in turn, influence covers of epifauna. Thus, differences in covers of epifauna on kelp can be due to: (1) factors associated with the primary habitats (pilings vs. reefs), (2) differences between characteristics of the kelp found in each habitat, or (3) an interaction between these factors (habitat and/or type of kelp). Kelp were experimentally transplanted between pilings and reefs, demonstrating that properties of the habitat directly affected covers of epifauna, which were not influenced by the type of kelp that grows on pilings or rocky reefs. Manipulative experiments to unconfound multiple components of habitats influencing disturbances to biota are needed to understand human impacts on natural systems.

Evaluating accuracy and precision of species-area relationships for multiple estimators and different marine assemblages.

This paper evaluates the accuracy and precision of six nonparametric estimators and six regression models for predicting the number of species in an area using random subsamples of that area. Such extrapolations are important for increased understanding of species-area relationships in different assemblages of species to inform ecological theories, and because they have direct implications in many conservation issues. Often, such extrapolations are evaluated by considering how well they fit existing data, rather than the actual number of species in the larger area. Also, many studies have focused on larger species in terrestrial habitats, where numbers of species are quite well known. Assemblages of invertebrates and smaller plants, especially in marine habitats, are less well known, and their distributions are dominated by small-scale variability. We examined species-area relations for two assemblages on boulders from six locations, for rocky shore assemblages in three locations, and for gastropods colonizing artificial habitat. Changing the focus of subsampling relative to the extent of the study and changing the size of the subsamples was also evaluated for subsets of data. Most estimators fitted the number of species poorly, with the second-order Jackknife consistently the best of the nonparametric measures, and the Negative Exponential the best of the regression models. Increasing the number of replicates improved most models, but some only slightly, and others considerably. Changing focus had little effect, probably because marine assemblages such as these vary at small scales as much as at many larger scales. These extensive analyses provide clear evidence about which estimators should not be used for measuring species-area relationships in assemblages such as these, and which will consistently over- or underestimate the number of species. The findings are applicable to many assemblages dominated by small organisms with strong stochastic variation.

Evidence necessary for taxa to be reliable indicators of environmental conditions or impacts.

Using taxa as indicators of environmental impacts is widespread. Indicators are chosen because they are considered to be easy to measure, sensitive to stresses and respond to stresses in predictable ways. Here, we review these criteria by addressing the nature of the relationships between some characteristic of taxa and the environmental variables they are supposed to indicate. It is crucial that variables measured as indicators be strongly and consistently correlated (through space and time) with levels of the environmental variables. Appropriate experiments must be done to establish that an observed correlation is causal, or the correlation cannot be considered sufficient to identify a useful indicator. Finally, it is necessary to establish that the taxa directly respond to changes in the environmental variables they are supposed to indicate. Appropriate methodologies to establish these criteria are considered and we evaluate studies in which these criteria have or have not been met.

Complexity and idiosyncrasy in the responses of algae to disturbance in mono- and multi-species assemblages.

There is considerable debate about whether stability (e.g. inertia) of an assemblage, or of individuals in an assemblage, is positively associated with the number of species or whether there are idiosyncratic effects of particular species. We assessed the general model that the loss of an individual alga, caused by trampling, is greater in monospecific than in multi-species stands but that the responses of algae are idiosyncratic, depending on the morphology of the species. The experiment was done on conspicuous and dominant algae with different morphology on temperate Australian rocky shores: the fucalean algae Hormosira banksii and Sargassum sp. and the coralline alga Corallina officinalis. We assessed the relative and interactive effects of the extent of trampling (number of paths) and the localised intensity of trampling (number of travels per path) on the three algae. The number of paths trampled (the extent of disturbance) had more impact on each alga than the number of times paths were travelled (the intensity of disturbance). As predicted, H. banksii was most susceptible to trampling at each level than were the coarser algae Sargassum sp. and C. officinalis. There was a consistent trend for each alga to be more inert to trampling when in the presence of the other two species than when in monospecific stands, but this was only statistically significant (P < 0.05) for the softer alga H. banksii. The responses of H. banksii and Sargassum sp. to disturbance seemed, in many cases, to be due to the presence of C. officinalis rather than to "diversity" per se. The relationship between the number of species and stability is complex in intertidal habitats, depending on the species and the combinations of species with which it grows.

Interactions of components of habitats alter composition and variability of assemblages.

1. The nature and resources supplied by different components of habitats influence species, creating variability from place to place within a habitat. 2. Experiments were done to investigate the effects of altering components of habitats on the variability of assemblages of numerous species of intertidal gastropods. 3. Artificial habitats with three levels of structure, combining different types of turf (i.e. different densities and height of fronds) were sampled 8 weeks after deployment in the intertidal. They were rapidly colonized by up to 66 species of gastropods. 4. Independently of the types of turf combined to form different habitats, there were differences in assemblages where there was more than one type of component present. Multivariate dissimilarities among units making up each habitat were also greater where there were more than one type of unit, but there was no such difference in the variance of numbers of species per unit. 5. Altering the relative abundances of different types of components made little change to the assemblages, nor their multivariate variability among units of habitat and the variance in number of species per unit in each habitat. 6. Differences in assemblages due to the different structure of habitat are complex to interpret and simple characterizations of structure of habitat are inadequate. Comparing different habitats requires appropriate experimentation to ensure that variability within habitats is also investigated.

Quantification of radular marks as a method for estimating grazing of intertidal gastropods on rocky shores.

Wax discs have been used previously on intertidal rocky shores to record the grazing activity of gastropods. This study has evaluated this methodology for recording grazing of four common intertidal microalgal grazers on intertidal shores in New South Wales, Australia. In the laboratory, the four species examined-the patellid limpet, Cellana tramoserica (Sowerby), the trochid, Austrocochlea porcata (A. Adams), the neritid, Nerita atramentosa Reeve and the littorinid, Bembicium nanum (Lamarck)-made distinctive marks in the wax. These allowed identification of each species or combinations of species grazing over the different discs. Field experiments showed that the intensity of grazing, as indicated by the mean number of scratches per disc, was positively related to the number of gastropods in the surrounding area during low tide for C. tramoserica. The number of scratches per disc in any area was correlated with the percentage of discs scratched. The relationship for C. tramoserica was found at two scales-in sites (approximately 3x3 m) and also in plots (50x50 cm) within sites. Therefore, densities that were measured when these limpets were inactive during low tide provided good estimates of grazing activity during high tide. This is largely because these limpets do not move far between where they rest and where they feed. The amount of microalgal food in the vicinity was not correlated with density, nor with grazing intensity. No relationship between density and grazing intensity was found for N. atramentosa, although experiments were only done in the field at one spatial scale (in sites, 3x3 m). Results obtained in the laboratory and in the field show that wax discs are useful to distinguish grazing by different species of gastropods on Australian rocky shores and allow tests of hypotheses about grazing activity at different spatial scales.

A method for analysing spatial scales of variation in composition of assemblages.

In several areas of research on ecological assemblages, it is useful to be able to analyse patterns of spatial variation at various scales. Multivariate analyses of dissimilarity or similarity in assemblages of species are limited by problems of non-independence caused by repeated use of the sample-units. Where rank-order procedures are used, no comparative quantitative measurements of dissimilarity at different scales are produced. An alternative method is described that uses the sample's average assemblage (or centroid). These estimates are themselves averaged to give centroids for larger spatial scales. Dissimilarities from the centroids at each scale are then calculated using independent replicates for each scale from those in each sample. The dissimilarity measures can then be examined by analysis of variance to detect spatial scales of differences for each sample at every level of a hierarchy of scales. The method is illustrated using data from mangrove forests and rocky shores, involving up to 97 taxonomic groups (species, other taxa). Differences among assemblages at the scales of sites (tens of meters apart) or locations at shores (hundreds of meters apart) were identified. Consequences of different numbers of replicates are discussed, with some potential problems (and their solutions) in application.

Effects of gastropod grazers on recruitment and succession of an estuarine assemblage: a multivariate and univariate approach.

Grazers have been shown to affect assemblages of species in many habitats. Here we studied the effects of the gastropod grazers, Austrocochlea porcata and Bembicium auratum, on intertidal estuarine assemblages in a sheltered bay in New South Wales, Australia. We examined the effects of gastropods on individual species and on the assemblage as a whole. The multivariate response was compared with data on succession in these assemblages to estimate potential effects of grazers on succession. The experiment was repeated several times to determine the generality of grazer effects in the light of possible variation in the timing or intensity of recruitment. There were different responses of individual species to the presence of grazers. Grazers reduced the abundance of ephemeral algal species, bryozoans, copepods, insect larvae and Balanus spp. barnacles. They had a positive effect on oysters and spirorbids and no effect on the barnacles Elminius covertus and Hexaminius spp. These effects were consistent through time. Multivariate analyses confirmed that grazers caused significant changes to whole assemblages and that these effects were far-reaching and not only caused by changes to algal species. The removal of grazers appeared to neither speed up nor slow down succession, but rather caused a completely different assemblage to develop. Apparent important mechanisms affecting the composition of animal species when grazers were removed included accumulation of sediments and detritus and pre-emption of space by algae.

Sampling to differentiate between pulse and press perturbations.

There is great inconsistency in the use of the terms 'pulse' and 'press' when describing types of perturbations. This is due primarily to a failure to distinguish between the cause and the effect of the perturbation in question. The cause and effect may be either short- or long-term and clearly one may be short-term and the other long-term. Distinction between these two types of disturbance is crucial for management to prevent further impact. Thus, it is important to describe separately these two aspects of a perturbation. Here, we define a protocol for sampling perturbations which enables the cause and effect to be distinguished between short- or long-term. Existing (i.e., already established) assemblages and newly-established assemblages are sampled and compared among disturbed and control locations. Existing assemblages may have been affected by past (pulse) disturbances and/or ongoing (press) disturbances, whereas the establishment of new assemblages can only be influenced by ongoing disturbances. We describe the procedures for assessing impacts of estuarine marinas as an illustration of the issues to be considered in any habitat. Settlement plates and defaunated sediment are suggested for sampling the establishment of new assemblages in aquatic environments.

Scales of spatial patterns of distribution of intertidal invertebrates.

Few comparative studies of spatial patterns at different scales have examined several species in the same habitat or the same species over a range of habitats. Therefore, variability in patterns among species or among habitats has seldom been documented. This study quantifies spatial patterns of a suite of intertidal snails and a species of barnacle using a range of statistical techniques. Variability in densities was quantified from the scale of adjacent quadrats (over a distance of centimeters) to tens of kilometers. Significant differences in abundances occurred primarily at two spatial scales. Small-scale differences were found at the scales of centimeters or 1-2 m and, for many species on many shores, these accounted for most of the variability in abundances from place to place. These are likely to be determined by behavioural responses to small-scale patches of microhabitat. Large-scale differences in abundance were also found in most species at the scale of hundreds of meters alongshore. These are likely to be due to variation in recruitment (and/or mortality) because of limited dispersal by adults of these species. There was little or no additional variation among shores, separated by tens of kilometers, than was shown among patches of shore separated by hundreds of meters. Identification of the scale(s) at which significant differences in abundance are found focus attention on the processes (and the scales at which these processes operate) that influence patterns of distribution and abundance. Some of the advantages and disadvantages of various procedures are discussed.

Supply-side ecology and benthic marine assemblages.

Many marine invertebrates have a planktonic stage of their life history during which widespread dispersal and much mortality occur. The numbers surviving to recruit into habitats occupied by adults are therefore very variable in time and space. Models for the structure and dynamics of benthic assemblages tend to focus on processes causing death - often assuming consistent arrivals of recruits. Supply-side ecology is a newly fashionable term to describe recent interest in the long-realized consequences of variations in recruitment. Such variations have important influences on theory and empirical research in these assemblages.

Vertical and seasonal patterns in competition for microalgae between intertidal gastropods.

Grazing by the snail Nerita atramentosa and the limpet Cellana tramoserica caused similarly great reductions in abundance of microalgae - measured by direct counts and by estimation of chlorophyll analyses. A smaller snail, Bembicium nanum, caused smaller reductions of microalgal resources, compared with ungrazed areas. These results were consistent with the competitive abilities of these three species. Chlorophyll concentrations in samples of grazed rocks were reliable estimates of the nature and abundance of food available to the grazers.Inter- and intra-specific competition amongst Nerita and Cellana were investigated at various densities in experimental cages. To examine the effects of different availability of food resources, the experiments were repeated at three heights on the shore (abundance of food decreases with height) and during autumn/winter and spring/summer periods of the year (less food is available during summer).Density-dependent mortality of Cellana was caused by the presence of other limpets, or of Nerita. Mortality was greatest at higher levels and during the spring/summer and was significantly, negatively correlated with mean chlorophyll concentration in the experimental cages. The only exception was that all limpets suddenly died in cages at the highest level during the summer period, which cannot be explained solely by competitive interactions. Nerita showed no density-dependent mortality during the short periods of these experiments. Tissue-weights of Nerita declined with increased density, but the effect of Cellana was not as great as the intraspecific effect of Nerita. Snails retained weight better at lower than at higher levels, and during the autumn/winter which is consistent with the availability of food. Tissue weights of both Nerita and Cellana were positively correlated with chlorophyll concentrations inside the cages in both seasons investigated.These experimental results demonstrate that intensity of competition will vary from place to place and time to time according to the densities and mixtures of the grazers, and according to the availability of microalgal food.

The apparent diet of predators and biases due to different handling times of their prey.

In order to estimate the true diet of predators, the prey of a number of predators is recorded at one time. Such sampling underestimates the true diet during a period of time. Where handling times of different types of prey are very different, these estimates will be biased, because prey that take a relatively long time to be eaten will be overestimated. We examined a rocky intertidal predatorprey system and demonstrated the existence of such bias. A number of hypothetical correlates of the bias were also investigated. As anticipated, variations in handling times were a major factor, but neither taxonomic affinity nor absolute size of the prey could predict the degree of underestimation in the true diet for any given type of prey. A previously described correction for this type of bias was tested, but found to be unsatisfactory. We suggest that it was too insensitive to variability in handling times.A simple computer model incorporating differences among prey in their handling times was also unable to predict the bias, but did indicate that non-random selection of prey was occurring.We concluded that where such biases are likely to occur, information on the handling times of different prey and/or accurate estimates of the true diets of the predators are essential for the predatory interaction to be interpreted properly. These results were discussed in relation to published accounts of diets of predators in rocky intertidal habitats. Many studies have not presented data on handling times of prey in the field, and the magnitude and importance of potential biases in these studies are therefore difficult to assess.

Experimental analyses of the structure and dynamics of mid-shore rocky intertidal communities in New South Wales.

At mid-shore levels on rocky shores in New South Wales, grazing gastropods are the dominant species in sheltered areas. Where wave-exposure is great, barnacles occupy most of the space. At intermediate levels of waveexposure, there are mixtures of grazing gastropods and barnacles, and the patterns of occupancy of space, and structure of the community change from time to time. The major species found in these areas are the coronuloid barnacle Tesseropora rosea, the patellid limpet Cellana tramoserica, the smaller acmaeid limpet, Patelloida latistrigata, which is mostly confined amongst barnacles, and the predatory whelk Morula marginalba. The roles of each of these species in determining the structure and persistence of intertidal communities were investigated by experimental manipulations of the densities of each of these organisms. In most experiments, a range of densities of limpets and barnacles was used, rather than the simple removal of all of one species.Recruitment of Cellana was negatively associated with increasing density of adult limpets, and with increasing density of barnacles. Growth and survival of juvenile Cellana were decreased by increasing densities of barnacles, probably because barnacles occupied space, preventing limpets from grazing. Morula had no effect on recruitment or survival of juvenile Cellana. Recruitment of juvenile Patelloida was not affected by different densities of barnacles, but survival to adult sizes was poor in areas where whelks are active. In areas where whelks were removed, Patelloida showed increased survivorship with increased cover of barnacles, probably because Patelloida amongst barnacles found refuge from the superior competitive effects of Cellana.The settlement and subsequent survival of Tesseropora were affected in complex ways by the activities of Cellana. At great densities, Cellana can have deleterious effects on newly-settled barnacles, probably by crushing them whilst grazing. In some areas, and at some densities, however, limpets can have beneficial effects on the recruitment and/or survival of Tesseropora. The limpets graze the juvenile stages of growth of foliose macroalgae, preventing them from growing up to pre-empt the rock-surface (thus preventing settlement of barnacles) or to smother alreadysettled barnacles. The effect of limpets on the recruitment and survival of barnacles in any area is a function of the densities of limpets and barnacles, and the height on the shore and local weather (the latter factors influence the rates of growth of algae).Increased cover of the rock-surface by adult barnacles caused reductions in the densities of Cellana. Limpets migrated away from areas of great cover of barnacles, and, if confined in such areas, starved and lost weight. The dispersion, as well as the density of the barnacles was important in determining the effects of barnacle cover. Where barnacles occupied half the space, but were scattered, leaving only small patches of bare rock, they had the same deleterious effects on Cellana as in areas where they were spread evenly to occupy most of the rock-surface. Thus, barnacles could invade areas dominated by limpets provided they recruited in sufficient numbers. They did not have to saturate an area to displace the Cellana.In these communities, all of the species can be considered to have important roles in the establishment and maintenance of community structure. We conclude that interpretations of the roles of individual species must be based on direct, experimental investigation. In this system, there was no indication that many of the species were functionally insignificant.The present experiments also revealed that the interactions among even a few species are very varied and complex; proper investigation requires considerable replication and repeated experimentation in different places and years.Finally, although the present studies allow reliable interpretations of observed patterns of occupancy and dominance on natural shores, the experiments did not provide a predictive framework to anticipate the future events in any area. This is because of great variability in the timing and intensity of recruitment of planktonic propagules of all the components in the system, and in the density and activity of predatory whelks in different areas. These results suggest that tightly co-evolved community relationships are not likely to be important, even if they appear to exist, in communities where most of the species have widely dispersed pelagic offspring and interact in diverse and complex ways at different densities.

Analysis of inter- and intra-specific competition amongst intertidal limpets with different methods of feeding.

The prosobranch limpet Cellana tramoserica grazes on microalgae, including the spores of macroalgae, and coexists with the pulmonate limpets Siphonaria denticulata and S. virgulata at mid-tidal levels on sandstone shores in New South Wales. These siphonarians graze on macroalgae, leaving the basal parts of the thalli intact. Where Cellana graze, they are capable of removing all algae; where Siphonaria graze, they leave at least a thin film of alga on the rocks, which is available to Cellana. S. denticulata normally show invariant homing behaviour, whereas S. virgulata tend to move around at random when Cellana are present, but apparently home when Cellana are absent. Both siphonarians have been observed to show opportunistic behavioural responses by moving towards patches of macroalgal foods when they become available.Experimental ecclosures of limpets at different densities and in different combinations revealed that Cellana tramoserica suffered increased mortality and reduced growth due to intraspecific competition when at increased densities. There was no effect on Cellana of increased densities of either species of Siphonaria. Nor was there any interspecific interaction between the siphonarians. Both species of Siphonaria showed some reduction of growth at increased intraspecific density. More importantly, both showed increased mortality when enclosed with low densities of Cellana. Larger densities of Cellana had no effect; the numbers of Cellana could not be maintained because of the reductions caused by intraspecific competition. Even after 27 weeks in enclosures with Cellana, the numbers of Siphonaria never declined to zero in any experimental enclosure. Thus, Cellana has a competitive effect on the survival of siphonarian limpets, but is unable to exclude them from an area of the shore. Siphonaria spp., in contrast, have no effect on Cellana.The nature of the competitive interactions between these types of limpets is explaied in terms of their methods of feeding; Cellana can exploit the food-resource before it reaches a suitable size for Siphonaria. The coexistence of Siphonaria spp. with Cellana is discussed with respect to the behaviour of the pulmonates. Intraspecific competition leading to reduced densitities of Cellana, however, will ensure that Cellana cannot exploit all the food resources, and some will be available to Siphonaria. The consequences of inter- and intra-specific competition among grazing gastropods are discussed with reference to the structure of intertidal communities, and it appears that competition for food is fundamentally different from competitive interactions for space in the organization of such communities.