A fossil Halobates from the Mediterranean and the origin of sea skaters (Hemiptera, Gerridae)

Five species of sea skaters, genus Halobates Eschscholtz, are the only insects to have successfully colonized the open ocean. In addition, 38 species are found in sheltered coastal waters throughout tropical Indo-Pacific. The taxonomy of the genus is relatively well known, and the phylogeneuc relationships between extant species have recently been analysed (using cladistic methods). In the present paper, we describe the first fossil species of sea skaters, Halobates ruffoi sp. no v. from the Eocene deposit 'Pesciara di Bolca', in the province of Verona, northeastern Italy (geological age about 45 Myr). The significance of this fossil in setting the time scale for the reconstructed phylogeny and anagenesis of adaptive features of sea skaters, and in understanding the evolution and historical zoogeography of these marine insects is discussed.     

Distribution and environmental synchronization of the marine insect, Halobates robustus, in the Galapagos islands.

The following three aspects of the biology of the marine insect Halobates robustus were studied, during a two week observation period, at several sites in the Galapagos Islands: distribution, aggregation behaviour and rhythmicity of locomotory activity. H. robustus occurred in highest numbers on the water surface at shores fringed with mangroves. The aggregations of H. robustus varied according to their location and density. Copulating adults formed dense, floating aggregations, which tended to be close to rocks or mangroves. Late instar nymphs were less aggregated and, in lagoons (where there was some shelter from direct tidal forces), were furthest from the shore. In two types of habitat (mangrove-fringed, sandy shores and rocky shores) the aggregations of H. robustus showed a pronounced ability to maintain a floating station in relation to the surrounding environment, irrespective of tidal movements (in one case at 34 m from the nearest fixed objects). Evidence of the ability of the aggregations to maintain station on the water surface was also obtained by comparing the movements of H. robustus with those of floating polystyrene particles, which move passively with wind and tide. Laboratory observations and experiments indicated no clear periodicity in locomotory movements throughout a 24 h period. However, the frequency of encounters between individuals showed two daily peaks, post-dawn and pre-dusk, with fewer encounters during the day and only occasional encounters during the night. By shifting the light-dark cycle it was demonstrated that the daily bimodal rhythm of encounters is triggered by dawn and, since it is not maintained in constant light or dark, an 'hour-glass' mechanism is suggested. The contributions of single adults, of copulating pairs and of immature stages to the overall pattern of activity were also determined. Immature stages did not affect the overall rate of encounters significantly and the interactions between single and copulating pairs of adults appear to have been responsible for the bimodal pattern.     

Marine insects: genital morphology, phylogeny and evolution of sea skaters, genus Halobates (Hemiptera: Gerridae)

Five species of sea skaters, genus Halobates Eschscholtz, are the only insects to have successfully colonized the open ocean. In addition, 36 species are found in sheltered coastal waters throughout tropical Indo-Pacific. The taxonomy of the genus is relatively well known, but reliable hypotheses about phylogenetic relationships are required if the biogeography and evolution of sea skaters is to be discussed in a meaningful way. This work presents the results of a study of new characters from the genital segments, especially those of the male phallus and the female gynatrial complex, and a reinterpretation for several other characters. In total 64 characters were scored for 26 species of Halobates , two species ofAsclepios and one species of Metrocoris. With Asclepios and Metrocoris species as outgroups, the character state sets were analysed cladistically using the computer program Hennig86. After critical evaluations of both characters and clades, a phylogeny for Halobates is presented and its taxonomic implications are discussed. A number of monophyletic species groups are delimited. One genus-level synonymy and three species-level synonymies are suggested. The evolution of Halobates is discussed in the light of the reconstructed phylogeny and present knowledge of the ecology and behaviour of sea skaters. A hypothesis of ecological evolution in halobatine water striders is proposed and tested.     

WATER MOTION AND MORPHOLOGY IN CHONDRUS CRISPUS (RHODOPHYTA)1

Morphological variability of intertidal Chondrus crispus Stackh. fronds along a small open rocky coast was related to wave exposure and emersion. Cluster analysis revealed two well-defined morphologies: filiform and planiform, named the N morphotype and B morphotype, respectively. We propose a rapid method of classifying fronds based on the morphology of the cross section at half the height on the thallus. The N morphotype is characterized by fewer dichotomies per unit length, a circular cross section with a large inner cortex, and narrow fronds. It is abundant at low intertidal and exposed sites. The B morphotype is characterized by more dichotomies, smaller sizes, a subelliptical or flattened cross section, and broad fronds. It is abundant at high intertidal sites in sheltered areas. Regression analysis revealed a major effect of water movements on frond morphology with respect to tidal level, which was more evident at high intertidal levels. No relationships were observed between morphology and life history phases.     

Chemical composition and behavioral responses of the marine insect Halobates hawaiiensis (Heteroptera: Gerridae)

Halobates is the only insect genus with representatives in the open ocean. How adults find one another at sea has long been an intriguing issue. Since chemical communications have been demonstrated in a related marine veliid Trochopus, and laboratory bioassays indicated behavioral differences between males and females when insect extracts were presented, we carried out similar studies on Halobates. Analyses of surface lipid constituents of female and male Halobates hawaiiensis revealed marked differences. Palmitic and oleic acid, major constituents in the male extracts, were absent in the female extract, whereas nonacosenol, dominating the female extracts, was not detected in the male extracts. Analyses of nymphal extracts indicated an intermediate chemical profile. Surface waxes of all insect stages investigated showed nonacosanol and isononacosanol to be main components. "Headspace" analyses of airborne chemicals showed high levels of 4-hydroxy-4-methyl-2-pentanone and benzaldehyde from the male, whereas benzyl alcohol was the main component in the female mixture.     

The habitat function of mangroves for terrestrial and marine fauna: A review

Mangroves are defined by the presence of trees that mainly occur in the intertidal zone, between land and sea, in the (sub) tropics. The intertidal zone is characterised by highly variable environmental factors, such as temperature, sedimentation and tidal currents. The aerial roots of mangroves partly stabilise this environment and provide a substratum on which many species of plants and animals live. Above the water, the mangrove trees and canopy provide important habitat for a wide range of species. These include birds, insects, mammals and reptiles. Below the water, the mangrove roots are overgrown by epibionts such as tunicates, sponges, algae, and bivalves. The soft substratum in the mangroves forms habitat for various infaunal and epifaunal species, while the space between roots provides shelter and food for motile fauna such as prawns, crabs and fishes. Mangrove litter is transformed into detritus, which partly supports the mangrove food web. Plankton, epiphytic algae and microphytobenthos also form an important basis for the mangrove food web. Due to the high abundance of food and shelter, and low predation pressure, mangroves form an ideal habitat for a variety of animal species, during part or all of their life cycles. As such, mangroves may function as nursery habitats for (commercially important) crab, prawn and fish species, and support offshore fish populations and fisheries. Evidence for linkages between mangroves and offshore habitats by animal migrations is still scarce, but highly needed for management and conservation purposes. Here, we firstly reviewed the habitat function of mangroves by common taxa of terrestrial and marine animals. Secondly, we reviewed the literature with regard to the degree of interlinkage between mangroves and adjacent habitats, a research area which has received increasing attention in the last decade. Finally, we reviewed current insights into the degree to which mangrove litter fuels the mangrove food web, since this has been the subject of long-standing debate.     

Effect of an integrated mangrove-aquaculture system on aquacultural health

Mangroves are unique intertidal halophyte formations growing in sheltered tropical and subtropical coastal areas. Due to the increasing population and economic development, mangroves have faced degrada-tion and loss, which has been mainly caused by land conversion into aquaculture ponds in Asia. In the past several decades, the rapid growth of aquaculture has induced water pollution. Using mangroves for effluent treatment from coastal aquaculture ponds could be a suitable approach for wastewater treatment and healthy aquaculture development. An Integrated Mangrove-Aquaculture System (IMAS) was established to test whether the idea of a mangrove in situ treatment for aquaculture wastewater is feasible. The monocultures of three mangroves, Sonneratia caseolaris, Kandelia obo-vata, and Aegiceras corniculatum were established with area proportions of 45%, 30%, or 15%, respectively. One control pond without mangroves was also set up. The results indicated that the mangroves had different toler-abilities to long-term inundation. The aquaculture ponds had different fishery yields, considering the mangrove species and area proportions. The water quality of most of the experimental ponds was better than the control pond, except for the planted Sonneratia. It is concluded that mangroves can reduce the concentration of dissolved inorganic nitrogen and phosphate, buffer the pH value and increase the concertration of dissolved oxygen in aqua-culture water bodies effectively. It is suggested to use 15% of the Aegiceras corniculatum area to conduct in situ purification of aquaculture wastewater and to enhance aquaculture production.     

Biology and distribution of the littoral rove beetle Psamathobledius punctatissimus (Le Conte) (Coleoptera: Staphylinidae)

The intertidal macrofauna on a small sheltered marine beach at Pawley's Island, South Carolina, was dominated by insects, primarily Psamathobledius punctatissimus, a small beetle which attained densities of up to 2 260 adults m⁻². Both adult and larval beetles remained buried in the sand when covered by the tide, then emerged to form mole-like surface trails and feed upon diatoms when exposed. Eggs and young larvae were maintained within special maternal burrows, while larger larvae and non-breeding adults occupied their own individual burrows. Both adults and larvae became comatose when in direct contact with water, but recovered after as long as six hours submergence. In the field, burrows retained air during tidal coverage, allowing the beetles to avoid direct contact with sea water. Seaward extension of the population is probably limited by time available for feeding, while longshore distribution is restricted by sand texture and diatom concentration, or by sediment mobility in more exposed locations. Although very locally distributed, P. punctatissimus has achieved considerable success in a habitat poorly exploited by competitors of either marine or terrestrial origin. Contribution No. 434 of the Belle W. Baruch Institute for Marine Biology and Coastal Research Contribution No. 434 of the Belle W. Baruch Institute for Marine Biology and Coastal Research     

Differential predation on mangrove propagules in open and closed canopy forest habitats

Amounts of seed predation by grapsid crabs (Brachyura: Grapsidae) on two species of mangroves (Aegiceras corniculatum and Avicennia marina) were compared among different habitats in an Australian mangrove forest. For Avicennia, comparisons were between canopy gaps and the adjacent forest understory for six, mid intertidal, gaps of different sizes. For Aegiceras the comparisons were among canopy gaps in the high intertidal; open, accreting mud/sand banks where mangroves were colonizing in the low intertidal; and in the forest understory in both the high and low intertidal zones. These were repeated in the high salinity (35\%) downstream portion and the low salinity (0–5\%) upstream portion of a tidal river.Predation on Avicennia was significantly higher in the understory than in adjacent canopy gaps. Within a canopy opening, predation was greatest in the smallest gaps and lowest in the largest gaps. Predation on Aegiceras was greater in the high intertidal compared to the low intertidal, but no differences were found between river mouth and upstream locations. In the high intertidal zone of the forest, there were no differences in predation between canopy gap or forest understory sites for Aegiceras. In the low intertidal zone, however, significant differences in amount of predation were found between habitats. More Aegiceras propagules were consumed in the understory than on adjacent accreting sandbanks.Frequency of tidal inundation, which in turn affects the amount of time available to forage, is hypothesized to account for differences in predation between low and high intertidal forests and between small and large canopy gaps. Our results also suggest that shade intolerance in these two species may actually reflect an escape from predators, successful when the seeds are dispersed into open areas such as canopy gaps or mud banks.     

Effect of an integrated mangrove-aquaculture system on aquacultural health

Mangroves are unique intertidal halophyte formations growing in sheltered tropical and subtropical coastal areas. Due to the increasing population and economic development, mangroves have faced degradation and loss, which has been mainly caused by land conversion into aquaculture ponds in Asia. In the past several decades, the rapid growth of aquaculture has induced water pollution. Using mangroves for effluent treatment from coastal aquaculture ponds could be a suitable approach for wastewater treatment and healthy aquaculture development. An Integrated Mangrove-Aquaculture System (IMAS) was established to test whether the idea of a mangrove in situ treatment for aquaculture wastewater is feasible. The monocultures of three mangroves, Sonneratia caseolaris, Kandelia obovata, and Aegiceras corniculatum were established with area proportions of 45%, 30%, or 15%, respectively. One control pond without mangroves was also set up. The results indicated that the mangroves had different tolerabilities to long-term inundation. The aquaculture ponds had different fishery yields, considering the mangrove species and area proportions. The water quality of most of the experimental ponds was better than the control pond, except for the planted Sonneratia. It is concluded that mangroves can reduce the concentration of dissolved inorganic nitrogen and phosphate, buffer the pH value and increase the concertration of dissolved oxygen in aquaculture water bodies effectively. It is suggested to use 15% of the Aegiceras corniculatum area to conduct in situ purification of aquaculture wastewater and to enhance aquaculture production.     

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

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

黑肩绿盲蝽耐饥饿能力的研究

【目的】为早期释放黑肩绿盲蝽Cyrtorhinus lividipennis提供理论依据,研究了黑肩绿盲蝽不同虫态耐受饥饿及低温的能力。【方法】室内条件下,比较了稻苗+水、无苗有水和无苗无水三种食物条件下黑肩绿盲蝽耐饥力大小。【结果】适温(26℃)下,黑肩绿盲蝽3龄若虫、5龄若虫和成虫在稻苗+水的存活时间显著高于无苗有水,无苗有水时的存活时间又显著高于无苗无水,表明稻苗对于黑肩绿盲蝽有营养作用,取食稻苗有助于提高黑肩绿盲蝽的耐饥力。黑肩绿盲蝽成虫耐饥力最强,3龄若虫和5龄若虫接近,有苗有水时,3龄若虫、5龄若虫和成虫的平均存活时间分别为2.10、2.22和4.25 d。在低温(15℃)下,水稻苗对黑肩绿盲蝽成虫的存活不利,有苗有水时雌雄成虫的存活时间均低于无苗有水处理,其中雌成虫两处理间的差异显著(P=0.001)。【结论】黑肩绿盲蝽对饥饿和低温有较强的耐受性,以成虫最强,水稻生长前期田间释放时以成虫为宜。     

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.     

The effects of group size on predator avoidance in a marine insect

The ocean skater, Halobates robustus (Hemiptera: Gerridae), is a flightless insect which forms loosely aggregated fluotillas of various sizes close to lava and mangrove edges in the Galapagos. Field observations showed a clear effect of group size on the distance at which an approaching predator model induced avoidance behaviour (rapid withdrawal and/or dispersal). It is suggested that in large, highly aggregated groups, the insects are maximally aroused as a result of frequent bodily encounters with adjacent members of the flotilla. An approaching object thus induces avoidance behaviour with minimal visual stimulation at maximal distance. With small groups and single individuals, which are less aroused, a closer approach of the object is required to produce visual stimulation sufficient to activate the escape response.     

A review of mangrove rehabilitation in the Philippines: successes,failures and future prospects

From half a million hectares at the turn of the century, Philippine mangroves have declined to only 120,000 ha while fish/shrimp culture ponds have increased to 232,000 ha. Mangrove replanting programs have thus been popular, from community initiatives (1930s–1950s) to government-sponsored projects (1970s) to large-scale international development assistance programs (1980s to present). Planting costs escalated from less than US$100 to over $500/ha, with half of the latter amount allocated to administration, supervision and project management. Despite heavy funds for massive rehabilitation of mangrove forests over the last two decades, the long-term survival rates of mangroves are generally low at 10–20%. Poor survival can be mainly traced to two factors: inappropriate species and site selection. The favored but unsuitable Rhizophora are planted in sandy substrates of exposed coastlines instead of the natural colonizers Avicennia and Sonneratia. More significantly, planting sites are generally in the lower intertidal to subtidal zones where mangroves do not thrive rather than the optimal middle to upper intertidal levels, for a simple reason. Such ideal sites have long been converted to brackishwater fishponds whereas the former are open access areas with no ownership problems. The issue of pond ownership may be complex and difficult, but such should not outweigh ecological requirements: mangroves should be planted where fishponds are, not on seagrass beds and tidal flats where they never existed. This paper reviews eight mangrove initiatives in the Philippines and evaluates the biophysical and institutional factors behind success or failure. The authors recommend specific protocols (among them pushing for a 4:1 mangrove to pond ratio recommended for a healthy ecosystem) and wider policy directions to make mangrove rehabilitation in the country more effective.     

The introduced clam Ensis americanus in the Wadden Sea: field experiment on impact of bird predation and tidal level on survival and growth

In the Danish Wadden Sea the intertidal distribution of the introduced bivalve Ensis americanus (syn. E. directus) is restricted to a narrow zone around the mean low water level. To test the possible impact of birds and submersion time on dynamics and distribution of the clams, adult specimens of E. americanus collected near the low water line were transplanted to two intertidal sites and established in open and net-covered experimental plots for 9 weeks (autumn 2001). The lowest survival of clams was registered at the low-shore-site (LSS) in plots open to bird predators, suggesting that birds such as Common Eider (Somateria mollissima) or Oystercatcher (Haematopus ostralegus) may control the abundance of E. americanus at the lower tidal levels. For clams showing increment in shell length during the study period, the shell growth rates were highest at the LSS and lowest in the open plots at the high-shore-site (HSS). Differences in immersion time and thus food supply may explain this pattern. Body mass index (BMI) of the clams showed basically the same pattern as the survivorship: lowest BMI in open plots at the LSS and highest in the covered plots at this site. Clams from the HSS were intermediate in their BMI. Disturbance by birds in the open plots at the LSS may explain the low BMI. In conclusion birds may be an important factor controlling abundance of E. americanus in the lower intertidal zone.     

Community composition of the rocky intertidal at Helgoland (German Bight,North Sea)

At the rocky island of Helgoland (North Sea), the distribution and abundances of intertidal communities were assessed and the effects of wave exposure and tidal height on the spatial distribution patterns of the communities were evaluated. Macroalgae and invertebrates were sampled quantitatively along line transects in three intertidal locations, a semi-exposed, an exposed and a sheltered one. The semi-exposed location was characterised by (1) Ulva spp. at the high intertidal (Ulva-community), (2) mussels and periwinkles at the mid intertidal (Mytilus-community) and (3) Corallina officinalis and mainly the large brown alga Fucus serratus at the low intertidal (Fucus-community). The exposed location encompassed the mid and low intertidal; at both zones the Fucus-community occurred. The sheltered location was characterised by (1) barnacles (Balanus-community) and (2) bryozoans, hydrozoans and mainly the large brown alga Ascophyllum nodosum (Ascophyllum-community). At the semi-exposed, but not at the exposed location the communities changed with the intertidal position. A relationship between wave exposure and the occurrence of specific communities was shown for the sheltered location; in contrast, communities of the semi-exposed and the exposed location appear to be little influenced by wave exposure directly. The community concept and the potential causes of distribution patterns of the defined communities are discussed and suggestions for a future monitoring are given. Variations in the communities at different spatial scales speak in favour of a multiple scale sampling design to monitor changes in the intertidal communities at Helgoland. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Competition in Bembicium auratum (Gastropoda) and its effect on microalgal standing stock in mangrove muds

Summary Field experiments were undertaken on the intertidal gastropod Bembicium auratum, which is one of the dominant organisms in sheltered bays and mangroves in New South Wales. Animals were caged at various densities ranging from natural levels to about four times normal density. Increased density resulted in increased mortality (particularly of juveniles), reduced body weight, and (less conclusively) a decline in growth rate. Chlorophyll levels of the mangrove mud within the cages were monitored as an index of food availability, and increased rapidly in the control cages in the absence of Bembicium, remained steady at normal Bembicium densities, and declined at higher densities. Parallel experiments were conducted on the effects of substratum on Bembicium. One series of animals was allowed access to a hard substratum and the other only to mud. Body weights and survivorship were proportionally lower in animals deprived of a hard substratum. Juveniles were particularly susceptible to a shortage of food and more dependent on a hard substratum than adults. The population structure also suggests a high juvenile mortality but high adult survival and longevity. Bembicium is very abundant low on the shore but numbers decline and body weights increase higher up. Chlorophyll levels and amounts of hard substratum also decline up the shore. Possibly juveniles settle selectively (or survive better) low on the shore where oysters provide a dense hard substratum. Adults are less dependent on the hard substratum and may disperse up the shore, explaining the patterns of abundance and size.As Bembicium limits its own food supply and seemingly is limited by it, predation is unlikely to play an important role in restricting population density.     

Large scale patterns and trophic structure of southern African rocky shores: the roles of geographic variation and wave exposure

In this study we revise the biogeographic delimitation, and large-scale patterns of community structure of the intertidal rocky shores of southern Africa. We use binary (presence/absence) and per-species biomass data collected at fifteen localities and thirty-seven different rocky sites, encompassing the shores of southern Namibia, South Africa and southern Mozambique. Multivariate analyses revealed that the shores of southern Africa (south of 25°) can be divided into three main biogeographic provinces: the west coast or Namaqua province, the south coast or Agulhas province and the east coast or Natal province. The biomass structure of the intertidal rocky shores communities of southern Africa varied at a large scale, corresponding to biogeographic differences, while local-scale variation accorded with the intensity of local wave action. The average biomass of west coast communities was on average significantly greater than that of the south and east provinces. At a local scale, the community biomass on exposed shores was an order of magnitude greater than on sheltered shores, within all biogeographic provinces. Semi-exposed shores exhibited intermediate average biomass. The trophic structure of these communities varied significantly with wave action: autotrophs, filter-feeders and invertebrate predators were more prevalent on wave exposed than sheltered shores, whereas grazers were more abundant on sheltered and semi-exposed shores. Exposed shores were consistently dominated by far fewer species than semi-exposed and sheltered shores, independently of biogeographic differences. Within all biogeographic provinces semi-exposed and sheltered shores were more diverse than exposed shores. West coast intertidal communities therefore had high levels of biomass, but were consistently species-poor. Several working hypotheses that could explain these large and small-scale patterns are presented.