Ecological interactions in the provision of habitat by urban development: whelks and engineering by oysters on artificial seawalls

Abstract Increases in human population cause increased urbanization of most habitats, including the shoreline. This has many consequences for coastal environments, in particular the trend for artificial structures, such as seawalls, to replace natural habitats. Seawalls and natural shores support many of the common intertidal species, but others important on rocky shores are absent from or rare on many seawalls. The whelk Morula marginalba Blainville is an abundant and important predator on rocky shores of south‐eastern Australia, but is infrequently recorded on artificial substrata. In Sydney Harbour, where the Sydney rock oyster (Saccostrea glomerata Gould) was locally abundant, densities of M. marginalba on some seawalls appeared to be similar to those on rocky shores and to be larger than where there were few oysters. We sampled densities and sizes of whelks in four habitats, predicting and corroborating that: (i) on seawalls with many oysters, there would be more whelks than on seawalls with few oysters; (ii) where there are many oysters, densities of whelks would be similar on seawalls and rocky shores; and (iii) whelks would be larger where oysters were abundant. Growth and survival of whelks were measured to test hypotheses from the observed differences in size and density. Survival was greater in habitats with many oysters, which could explain differences in density, but size‐specific differences in survival could not explain differences in size among habitats. On seawalls but not on rocky shores, slower growth could explain the smaller size of whelks where there were few oysters. Seawalls provide important habitat for M. marginalba, but only via their indirect effects, mediated by oysters. These interactions cannot be predicted from those on natural rocky shores. Predicting how developed areas provide suitable habitat, either in management of conservation or in assessments of potential impacts clearly depends on understanding the roles of biogenic habitats.     

Intertidal seawalls as habitats for molluscs

Coastal urbanization changes intertidal shorelines, by alterationand destruction of natural habitat and addition of new builthabitat. In Sydney Harbour, New South Wales, Australia, up to50% of the shoreline is composed of constructed habitat, particularlyseawalls. Research has shown that many components of intertidalassemblages live on seawalls, but patterns of abundance anddiversity are very variable. Seawalls differ physically fromnatural shores in a number of ways that are likely to influencedistribution and abundances of intertidal molluscs, which arevery important determinants of structure of intertidal assemblages. This study examined diversity, abundances and frequencies ofoccurrence of intertidal molluscs on seawalls and on naturalhorizontal and vertical shores in numerous locations in SydneyHarbour and in natural boulder-fields and artificial boulder-fieldscreated from rubble of seawalls. On seawalls, assemblages variedbetween tidal heights and among locations, but when data werecombined across locations, there were some general patterns.Sessile bivalves (oysters and mussels) and many limpets werefound in similar numbers on both habitats, or patterns variedinconsistently. Many coiled snails, in contrast, including whelksand grazing gastropods, plus opisthobranchs, which were eithercommon or relatively sparse on horizontal shores, were not foundon seawalls and found in intermediate frequencies on verticalshores. Similarly, common species of molluscs were found innatural and artificial boulder-fields in similar numbers, orpatterns were not consistent, although rarer species were notfound in these boulder-fields. Because molluscs are such importantcontributors to intertidal dynamics, these results have importantimplications for management of intertidal biodiversity in urbanizedand altered environments. (Received 18 July 2005; accepted 9 November 2005)     

Exposure of seawalls to waves within an urban estuary: effects on intertidal assemblages

Abstract Despite being largely protected from sea swell by headlands, shores within Sydney Harbour, New South Wales, Australia are subjected to considerable wave action from wind waves and the numerous recreational and commercial boats and ships using the waterways. The aim of this study was to test whether assemblages on artificial shores, that is, seawalls, either exposed to or sheltered from waves would show similar patterns to those published for natural shores exposed to or sheltered from sea swell. Specifically, the hypotheses were tested that sheltered seawalls would have greater cover of algae, whereas exposed seawalls would have greater cover of sessile invertebrates and greater abundance of mobile invertebrates. It was found that encrusting algae had greater cover on sheltered seawalls, whereas cover of turfing algae was greater on exposed walls. Sessile filter‐feeders generally had greater cover on exposed seawalls, although a dominant space occupier, the oyster Saccostrea glomerata, showed either no difference, or greater cover on sheltered seawalls among different locations. The pattern for the dominant grazer on seawalls, the pulmonate limpet Siphonaria denticulata showed the predicted pattern low on the shore, but the opposite pattern at mid‐tidal levels. Despite some inconsistencies the results were similar to those predicted from studies on natural shores, showing that assemblages on urban structures respond to wave action in a way that is predictable from studies on natural shores. Experiments were carried out where assemblages were transplanted to sites with differing exposure to waves to determine whether differences in recruitment or post‐recruitment mortality were responsible for patterns of difference. We found that some sessile invertebrates did not survive when transplanted to sheltered seawalls, suggesting that difference in survival of organisms was responsible for observed patterns. This was not, however, found when the experiment was repeated at other locations. It is likely that processes affecting the assemblages are temporally and spatially very variable, or that more than one process interacts to cause observed patterns. Identifying such complexity requires rigorously structured sampling designs and appropriate manipulative experiments.     

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.     

Fish assemblages associated with urban structures and natural reefs in Sydney,Australia

Abstract Fish ecology in urban estuaries is poorly understood. As coastal landscapes are transformed, recognizing the impact that urban structures, such as marinas, seawalls and wharfs, have on local fish populations is becoming increasingly important. The extent to which fish are able to maintain natural ecological assemblages can be measured, to a certain extent, by how closely they mimic natural habitats. In Sydney Harbour, assemblages of fish associated with artificial structures were compared with those associated with natural rocky reefs. Sampling was carried out in five locations, each with a marina, swimming enclosure and natural reef. In each location, different habitats supported different assemblages, but differences between habitats were not consistent among locations. Subsequent sampling compared artificial and natural sites in three different areas in each of three different estuaries. Results indicated that differences in fish assemblages between artificial and natural sites were greater than differences between sites within each habitat, but there were no patterns among different positions in an estuary or from estuary to estuary. This study provides initial evidence that, although artificial habitats generally support the same species as found on natural reefs, assemblages usually differed between natural and artificial habitats. In addition, without knowing if these habitats do, in fact, sustain viable populations of fish, it would be premature to label artificial structures as effective habitat for fish.     

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.     

Mussel beds on different types of structures support different macroinvertebrate assemblages

Abstract Artificial structures, such as seawalls, pilings and pontoons, are common features of urban estuaries. They replace natural structures or add to the amount of hard substratum in an area and provide habitats for many fish and invertebrates. Previous work has concentrated on fish or on the invertebrates that occupy the primary substratum of artificial structures. Mussels often grow on different types of structures (pontoons, pilings, seawalls and natural reefs) and provide a secondary substratum for other organisms to inhabit. Counting and identifying organisms associated with mussel beds is traditionally done to species level, which is very time‐consuming. To save time, organisms in this study were identified to coarse levels of taxonomic resolution (a mix of taxa, such as class, order, family and genus), which showed similar patterns to those when particularly speciose and abundant groups were identified to species. This study tests hypotheses that the distribution and abundance of mobile and sessile organisms that inhabit mussel beds will differ among natural and various types of artificial structures. When the associated assemblages of mussel beds from different types of structures and from different locations were examined, assemblages varied according to the type of structure they inhabited and its location. Assemblages associated with mussels on pontoons differed consistently from those on other types of structures. Patterns in the assemblages were also consistent through time. These data show that the types and amounts of artificial structures added to an environment can affect the types, distribution and abundances of organisms living in biogenic habitats.     

Effects of structural diversity and identity of patches of habitat on diversity of benthic assemblages

A fundamental problem in ecology, regardless of habitat or system, is understanding the relationship between habitats and assemblage of organisms. It is commonly accepted that differences in composition and surrounding landscape of habitats affect the diversity of assemblages, although there is not much empirical evidence because of difficulties of manipulating structure in many habitats. These relationships were examined experimentally, using habitats of artificial turfs that are colonized by diverse assemblages of gastropods. Each habitat was made of nine sub‐habitats, which were sampled individually to allow tests of hypotheses about the effect of type of habitat and the influence of other adjacent sub‐habitats on the colonizing assemblage. Turf habitats were deployed for 8 weeks on a rocky shore after which they were collected and the colonizing assemblages of gastropods sampled. Independently of the types of turfs combined to form different habitats, there were more species where there was more than one type of component in a habitat (i.e. structural diversity). The type of habitat (i.e. structural identity) itself had little or no influence on the colonizing assemblage. The number of species colonizing short‐sparse and short‐dense turfs was influenced by which type of habitat was adjacent. Thus, when units of one type (e.g. short‐sparse turf) were added to a patch of habitat of long‐sparse turfs, the number of species in short‐sparse turfs was greater than in patches of the same type. This also increased total number of taxa in the whole patch of habitat. These results show how diversity of gastropods colonizing heterogeneous patches of habitat is influenced not only by the number of types of sub‐habitats, but also by interactions with surrounding sub‐habitats. These findings reiterate the importance of investigating the role of structure of habitats and of their surrounding landscapes across different systems, irrespective of their size or associated assemblages of organisms.     

Variation in rocky shore assemblages in the northwestern Mediterranean: contrasts between islands and the mainland

Insular assemblages of species are often considered unique because they are exposed to unpredictable patterns of colonization/extinction that depend on distance from other sources of colonists and on size of islands. An alternative explanation is that islands provide fundamentally different habitats of those of the mainland, regardless of any possible effect of size and isolation. These alternatives were examined by comparing assemblages of rocky shores on islands of the Tuscany Archipelago with those of the mainland in the same geographical region. Sandy beaches created a pattern of discrete areas of rock along the mainland with spatial discontinuities and extents comparable to those of the insular environment. Possible effects of isolation and size were therefore controlled in this study, so that one would expect no difference between islands and the mainland if only size and isolation matter. In contrast, differences are expected if historical events or other processes have distinct influences on assemblages in these environments. These hypotheses were tested by comparing assemblages of midshore and lowshore habitats of two islands with those of two similarly distributed locations on the mainland over a period of 2 years, using a hierarchical sampling design. Multivariate and univariate analyses revealed various patterns in the data. There were differences between islands and the mainland in structure of assemblages, in mean abundance of common taxa and in the magnitude of spatial and temporal variance in abundance in both habitats. Collectively, these findings support the model that islands in the Tuscany Archipelago have distinct assemblages from the mainland, thereby contributing to the regional diversity and complexity of assemblages of rocky shores over and above any possible effect of size or isolation.     

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.     

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.     

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.     

Does colonization contribute to spatial patterns of common invertebrates in coralline algal turf?

Abstract The potential of colonization to contribute to the spatial patterns of six common invertebrates in coralline algal turf was investigated on a rocky shore near Sydney, Australia. The species, which included two amphipods (Elasmopus warra, Hyale spp.), a small bivalve (Lasaea australis), a fly larva (Limonia marina), and two microgastropods (Amphithalamus incidata and Eatoniella atropurpurea), had a range of dispersal modes (larval dispersal, crawling, swimming, rafting, and passive transport). Field sampling between May 1997 and November 1999 demonstrated that the amphipods were more abundant in low‐shore areas, the fly larvae and bivalves were more abundant in mid‐shore areas, and the abundances of gastropods did not vary with tidal height. Furthermore, abundances of all species varied among patches separated by tens of metres at one time or another. To test whether rates of colonization could contribute to established patterns of abundance, habitat mimics were deployed for 2‐week periods. The supply of new individuals matched long‐term patterns of abundance at different tidal heights for E. warra and L. marina. Colonization rates also differed among patches separated by tens of metres for three of the six species. Overall, there was little evidence to suggest that common species in coralline turf are limited by colonization on local scales, regardless of their major mode of dispersal. However, the potential for colonization to determine patterns of abundance varied from species to species.     

Floating pontoons create novel habitats for subtidal epibiota

Urban structures in the form of pontoons and pilings represent major coastal habitats for marine organisms and understanding the factors causing abundances of organisms to differ between these and natural habitat has been neglected in the study of coastal ecology. It has been proposed that composition of substrata explain differences previously described between subtidal assemblages of epibiota on rocky reef (sandstone) and pontoons (concrete) in Sydney Harbour, Australia. This study tested the hypothesis that differences in the composition of substratum (sandstone vs. concrete) independent of type of habitat (rocky reef vs. pontoon) affects the development of epibiotic assemblages. This was tested by experimentally providing substratum of the two types in both habitats. Epibiotic assemblages were unaffected by the composition of substratum but strongly affected by the type of habitat; demonstrating that pontoons constitute novel habitats for epibiota. This result highlights a need for determining how current ecological understanding of subtidal epibiota, which is heavily based on studies of urban structures (pilings and pontoons), relates to natural reef. Future tests of hypotheses about the nature of these differences will not only contribute to better ecological understanding of epibiota and their use of urban structures as habitats, but also to better predictions of future changes to the ecology of coastal habitats.     

Physical effects of habitat‐forming species override latitudinal trends in temperature

Latitudinal and elevational temperature gradients (LTG and ETG) play central roles in biogeographical theory, underpinning predictions of large‐scale patterns in organismal thermal stress, species' ranges and distributional responses to climate change. Yet an enormous fraction of Earth's taxa live exclusively in habitats where foundation species modify temperatures. We examine little‐explored implications of this widespread trend using a classic model system for understanding heat stresses – rocky intertidal shores. Through integrated field measurements and laboratory trials, we demonstrate that thermal buffering by centimetre‐thick mussel and seaweed beds eliminates differences in stress‐inducing high temperatures and associated mortality risk that would otherwise arise over 14° of latitude and ~ 1 m of shore elevation. These results reveal the extent to which physical effects of habitat‐formers can overwhelm broad‐scale thermal trends, suggesting a need to re‐evaluate climate change predictions for many species. Notably, inhabitant populations may exhibit deceptive resilience to warming until refuge‐forming taxa become imperiled.     

Sympatric speciation as a by-product of ecological adaptation in the Galician Littorina saxatilis hybrid zone

On exposed rocky shores in northwestern Spain there is a strikingpolymorphism of Littorina saxatilis that has been claimed asan example of a putative sympatric ecological speciation process.Two ecotypes of this species have evolved that are adapted todifferent shore levels and habitats, although they meet andhybridize on the mid shore where their two habitats overlap.As a by-product of adaptation these ecotypes have evolved anincomplete premating reproductive isolation where they meeton the mid shore. Although they are not yet true species, andthe final outcome of the process cannot be predicted, the ecologicalmechanisms responsible for this polymorphism could cause sympatricspeciation in similar situations. Here, I review all data insupport of these claims and discuss the interest of such a modelsystem in microevolutionary studies. (Received 13 February 2006; accepted 25 October 2006)     

Studying exotics in their native range: Can introduced fouling amphipods expand beyond artificial habitats?

Knowledge of the habitat use patterns of introduced species in their native or naturalized range can provide unique insights into processes of secondary dispersal and colonization of natural habitats. Caprellid amphipods are small mobile marine epibionts with limited natural dispersal. The global distribution of some caprellid species is mostly the result of anthropogenic transport; however, their subsequent spread beyond artificial habitats is poorly understood. A biogeographic approach, mainly focused on the native-range ecology of introduced and common fouling caprellid amphipods of southern Europe, was used to understand the implications of habitat use patterns for predicting their spread in the introduced regions. Specifically, abundance and composition of caprellid populations were compared among different primary habitats including artificial (floating pontoons), sheltered and wave-exposed rocky shores along the southern and southeastern coasts of Brazil. The findings indicated that artificial habitats act as reservoirs for globally distributed species in both their native and introduced ranges, while endemic species are more scarcely represented. Environmental conditions provided by primary habitats appear important in structuring caprellid assemblages on secondary substrata (basibiont species). Most wide-ranging caprellids were negatively correlated with the level of wave exposure, being more abundant in sheltered (artificial or natural) than in exposed habitats. In this context, Caprella scaura and Paracaprella pusilla, the two introduced caprellids recorded in the Mediterranean, where they are virtually restricted to artificial habitats, may become established in sheltered and even highly polluted natural habitats but hardly colonize wave-exposed rocky shores.     

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     

Patterns of distribution and abundance of chitons of the genus Ischnochiton in intertidal boulder fields

Abstract Chitons of the genus Ischnochiton are found predominantly on the undersurfaces of boulders, compared with other intertidal or subtidal habitats. They therefore appear to be habitat‐specialists at this scale. This, combined with the fact that boulder fields are relatively sparse compared with other intertidal habitats, makes these animals vulnerable to natural and anthropogenic disturbances. In addition, many species of Ischnochiton are relatively rare and appear to have very patchy abundances, making them likely to have very specific requirements for habitat. We need to understand the habitat requirements in order to manage, conserve and restore disturbed habitats. The present study was carried out at three intertidal boulder fields separated along approximately 200 km of the coast of New South Wales, Australia, centred around Sydney. The boulder fields were representative of those found in this region. The boulders were made of different materials: shale in the north, sandstone in Sydney and quartzitic sandstone in the south. Some boulders in each boulder field were covered by up to 0.4 m of water during low tide. The study showed that the seven species examined were overdispersed among boulders in each of these three intertidal boulder fields. Most boulders did not have associated chitons, but there were very large abundances on a very small number of boulders. Chitons were also overdispersed among boulders that they occupied. These patterns were consistent among shores and among species, even though patterns of abundance were extremely different among different species. These species appear therefore to show specific requirements for habitat at a small spatial scale, using only a small proportion of potential patches of habitat (i.e. boulders) in any place. Extremely patchy patterns of dispersion can be caused by variation in patterns of recruitment, mortality or behavioural responses to habitat or other species. Before performing experiments to investigate such processes, it is useful to test hypotheses of association with habitat using mensurative experiments to identify environmental correlates that might explain the observed patterns. In the present study, sizes of boulders and the associated sessile and mobile assemblages were proposed as mechanisms that could affect dispersion of chitons among boulders. None of these factors, however, showed strong associations with abundances of chitons. The lack of support of these models rules out some features of habitat to which species of Ischnochiton might respond, thereby precluding manipulative experiments involving these features, which are unlikely to be involved in the very patchy patterns of dispersion of species of Ischnochiton.     

Effect of diet and temperature upon muscle metabolic capacities and biochemical composition of gonad and muscle in Argopecten purpuratus Lamarck 1819

It has been difficult to understand the effects of habitat structure on assemblages because the different elements of habitat structure are often confounded. For example, few studies consider that the effects of structural components of a habitat (rocks, trees, pits, pneumatophores) may be separate from the complexity (e.g. surface area {SA}) they create. From prior observations and experiments, I developed three hypotheses about the effects of habitat structure on gastropods on rocky intertidal shores in Botany Bay, Australia. (1) The complexity of habitats positively affects the density and richness of gastropods. (2) The fractal dimension (D) represents elements of complexity that affect the density and richness of gastropods better than other indices of complexity. (3) The effects of specific structural components on the density and richness of gastropods are independent of their complexity. To test these hypotheses, treatments composed of pits and pneumatophores were used to independently manipulate complexity and structural components in experiments repeated at five different times on two shores. There was support for hypotheses (1) and (3) at most times and places but not for hypothesis (2). Richness, total density, and the densities of two of the three most common gastropods were greater in treatments with greater complexity. D was not definitively better than other indices of complexity, but D and SA were recommended for further consideration. When complexity was held constant, species richness and the density of most gastropods, except Austrocochlea porcata, was greater in treatments with pits than with pneumatophores. A common mechanistic explanation for the effects of habitat complexity on rocky intertidal gastropods relies on a specific characteristic of pits; they pool water and reduce desiccation stress. This assumption may be appropriate for many gastropods, but it was inappropriate for A. porcata. Habitat complexity affected its density, but this was not because of a characteristic specific to pits. The complexity and structural components of habitats have separate effects on assemblages, and it confuses the study of habitat structure to combine them.