Tide-model analysis of the zonation of intertidal prosobranchs. I. Four species of littorina (L.)

Experiments were carried out in a tide model to determine the patterns of zonation of four species of periwinkles under laboratory conditions. Both upper shore littorinids, Littorina neritoides (L.) and L. saxatilis (Olivi), showed the same zonation in relation to the tidal cycle as observed on the shore at Plymouth. In the model, the animals adopted a zone above the level of high water and then remained inactive. L. littoralis (L.) showed the same distribution unless fucoid algae were provided, in which case the periwinkles clustered on them. L. littorea (L.) showed the ability to adopt a pattern of zonation in the model, which was at, and just below, high tide level, but higher than that normally observed on the shore.     

The Response of Experimental Rocky Shore Communities to Nutrient Additions

The aim of this study was to determine whether the experimental nutrient enrichment of littoral rocky shore communities would be followed by a predicted accumulation of fast-growing opportunistic algae and a subsequent loss of perennial benthic vegetation. Inorganic nitrogen (N) and potassium (P) was added to eight concrete mesocosms inhabited by established littoral communities dominated by fucoids. The response to nutrient enrichment was followed for almost 2 1/2 years. Fast-growing opportunistic algae (periphyton and ephemeral green algae) grew significantly faster in response to nutrient enrichment, but the growth of red filamentous algae and large perennial brown algae was unaffected. However, these changes were not followed by comparable changes in the biomass and composition of the macroalgae. The biomass of opportunistic algae was stimulated only marginally by the nutrient enrichment, and perennial brown algae (fucoids) remained dominant in the mesocosm regardless of nutrient treatment level. Established rocky shore communities thus seem able to resist the effects of heavy nutrient loading. We found that the combined effects of the heavy competition for space and light imposed by canopy-forming algae, preferential grazing on opportunistic algae by herbivores, and physical disturbance, succeeded by a marked export of detached opportunistic algae, prevented the fast-growing algae from becoming dominant. However, recruitment studies showed that the opportunistic algae would become dominant when free space was available under conditions of high nutrient loading and low grazing pressure. These results show that established communities of perennial algae and associated fauna in rocky shore environments can prevent or delay the accumulation of bloom-forming opportunistic algae and that the replacement of long-lived macroalgae by opportunistic species at high nutrient loading may be a slow process. Nutrient enrichment may not, in itself, be enough to stimulate structural changes in rocky shore communities.     

The zoogeography of algae-associated peracarids along the Pacific coast of Chile

Abstract Aim To describe the zoogeography of the algae‐associated peracarid crustaceans from exposed rocky shores along the SE‐Pacific. Location Chile, 18° S to 42° S. Methods A standardized sampling programme was used at all sites. Samples of macroalgae were taken at twenty sites distributed along the entire study area. Quantitative samples (n = 6 replicates of 8 cm^?2 surface area each) of calcareous and non‐calcareous red algae were taken in the low intertidal, preserved immediately in 4%‐formalin and washed over a 0.2‐mm mesh before sorting. All peracarid individuals were sorted, identified to the species level and then categorized in separate functional groups according to their feeding habits. Graphical representations of species replacement within each functional group along the latitudinal gradient are provided. A classification analysis employing the unweighted paired group method using arithmetic average (UPGMA) was conducted in order to reveal the main zoogeographical zones. Results A total of forty epifaunal peracarid species was found. A gradual replacement of species within different functional groups (grazing and suspension‐feeding species) was observed in the central region (c. 26° S?37° S). In this central region, species with northern and those with southern distribution overlapped, while other species were only found here, resulting in high species richness. The number of species/site/algal species in the northern (north of c. 25.5° S) and southern region (south of c. 38.5° S) was considerably lower than in the central region. The distribution of most grazing peracarids showed a more continuous pattern than that of suspension‐feeding amphipods. The distribution of the remaining species (predators, scavengers, deposit‐feeders, unknowns) was scattered along the examined sites. The cluster analysis for the epifaunal peracarid assemblage confirmed the separation of a northern and southern zone connected by a central (transitional) zone between c. 26° S and c. 37° S. Similar zonation patterns have been found by most other studies on the zoogeography of the Chilean coast, although little agreement exists about the exact limits of this transitional zone. It is discussed that the distribution limits of algae‐associated peracarids (and other macroinvertebrates) – particularly in the transitional zone – may show interannual variations as a result of varying oceanographic conditions. The large affinity of the algae‐associated peracarid fauna from the central and southern Chilean coast to those of other regions indicates that dispersal may be facilitated by rafting with floating algae transported in the Antarctic Circumpolar Current. Main conclusions The zoogeographical analysis of algae‐associated peracarids confirms the existence of a northern and a southern zone connected by an extensive transitional zone. General biology, habitat use and the abundant presence of dispersal vectors such as floating macroalgae may affect the zoogeography of species living in transitional zones with strong interannual variations in current regimes. In these areas, species associated with substrata of high dispersal potential may show different distribution patterns than species inhabiting other substrata.     

Indirect effects shape macroalgal epifaunal communities

We tested the response of algal epifauna to the direct effects of predation and the indirect consequences of habitat change due to grazing and nutrient supply through upwelling using an abundant intertidal rhodophyte, Gelidium pristoides. We ran a mid‐shore field experiment at four sites (two upwelling sites interspersed with two non‐upwelling sites) along 450 km of the south coast of South Africa. The experiment was started in June 2014 and ran until June 2015. Four treatments (predator exclusion, grazer exclusion, control, and procedural control) set out in a block design (n = 5) were monitored monthly for algal cover for the first 6 months and every 2 months for the last 6 months. Epifaunal abundance, species composition, algal cover, and algal architectural complexity (measured using fractal geometry) were assessed after 12 months. Predation had no significant effect on epifaunal abundances, while upwelling interacted with treatment. Grazing reduced the architectural complexity of algae, with increased fractal dimensions in the absence of grazers, and also reduced algal cover at all sites, though the latter effect was only significant for upwelling sites. Epifaunal community composition was not significantly affected by the presence of herbivores or predators but differed among sites independently of upwelling; sites were more similar to nearby sites than those farther away. In contrast, total epifaunal abundance was significantly affected by grazing, when normalized to algal cover. Grazing reduced the cover of algae; thus, epifaunal abundances were not affected by the direct top‐down effects of predation but did respond to the indirect effects of grazing on habitat availability and quality. Our results indicate that epifaunal communities can be strongly influenced by the indirect consequences of biotic interactions.     

The importance of shore height and host identity for amphipod assemblages

The spatial distribution of organisms associated with marine intertidal macroalgae may be a direct result of their tolerance to air exposure or an indirect consequence of the distribution of their host. We compared amphipod assemblages from five intertidal macroalgae to investigate their relationship with algal identity. To test the effect of height regardless of algal characteristics, we transplanted coralline algal turfs to three different levels within the intertidal zone and compared amphipod assemblages after 1 and 14 days. Interstitial volume was positively correlated to the abundance of amphipods, suggesting that this attribute may correspond better to the potential space for their occupation when compared to algal biomass, thallus volume or the ratio between thallus and interstitial volume. Algal level determined the structure of the amphipod assemblages. Upper-level (Acanthophora spicifera and Caulerpa racemosa) and intermediate-level (coralline) algae host similar amphipod assemblages dominated by Apohyale media, but different from lower-level algae (Padina gymnospora and Sargassum cymosum), which were dominated by Hyale niger. Ten of the 15 amphipod species reported from natural communities were found in the transplanted plots. Distinct pools of amphipod species colonized coralline transplants at upper and lower levels after 1 day. However, regardless of the position on the shore, transplanted coralline turfs supported similar assemblages after 14 days, indicating that algal identity is also important for species assemblages. Our results suggest that both height on the shore and host identity combine to determine the vertical structure of amphipod assemblages in the rocky intertidal.     

Scale-dependant control of motile epifaunal community structure along a coral reef fishing gradient

Large-scale fishing is mostly conducted using towed gears that reduce the biomass and diversity of benthic invertebrates. However, it is impossible to differentiate between the physical disturbance effect of towed gears from the effect of fish predator removal upon benthic invertebrate communities. Here we explore the impact of fish removal alone on the community structure of small motile coral reef invertebrates (epifauna) along a subsistence fishing intensity gradient in the Lau group, Fiji. We deployed settlement plates at three areas in each of six fishing grounds and examined the density and class richness of the motile epifaunal communities and the associated algal communities in relation to the structure of fish and benthic communities. Motile epifaunal density was unrelated to fishing intensity. However, at smaller inter-area scale (0.5-10 km) motile epifaunal density was negatively related to plate algal biomass, whereas at the larger inter-fishing-ground scale (4-180 km) motile epifaunal density was positively related to the rugosity (substrate complexity) of the surrounding benthos. The class richness and diversity (Margalef's d) of motile epifaunal communities were negatively related to fishing intensity, but unrelated to grazing intensity, rugosity or algal biomass at either scale. Benthic community structure varied significantly with fishing intensity; hard-coral cover was lower and turf-algal cover was higher at high fishing pressure. The variation in benthic community structure was associated with variation in fish community structure, which in turn varied with fishing intensity. Motile epifaunal community structure upon plates was linked to the structure of the surrounding benthic community, but was not directly linked to the plate algal community. We suggest the decline in richness of the motile epifauna community along the fishing gradient is attributable to either to exploiter-mediated coexistence or the reduction in ‘habitat quality’ of the surrounding benthos. At the large spatial scale substrate complexity is the key determinant of motile epifaunal density, suggesting predation by fishes plays an important structuring role at this scale. Assuming that rugosity is inversely related to predation risk then this study represents the first evidence for spatial-dependence on the top-down (predation) vs. bottom-up (algal biomass) control of community structure. We argue fisheries exploitation, in the absence of a physical disturbance can negatively influence motile epifaunal community structure at large spatial scales.     

Effects of height on the shore and complexity of habitat on abundances of amphipods on rocky shores in New South Wales, Australia

Patterns of abundance of epifaunal crustaceans were compared between two common brown algae in intertidal rock pools on a shore near Sydney, New South Wales, Australia. Amphipods were significantly more abundant on Sargassum globulariaefolium (which was most common low on the shore) than on Hormosira banksii (more common at mid-shore levels). Experiments tested the hypotheses that height on the shore and structural complexity of the algae were the main factors influencing patterns of abundance of amphipods. Reciprocal transplants using defaunated plants and similar experiments using artificial plants allowed simultaneous tests of these hypotheses. The abundance of a common intertidal amphipod, Sunampithoe graxon was significantly smaller on Sargassum transferred from low to mid-tidal level of the shore than on any other treatments, indicating that height on the shore affects numbers of this species. There were no Sunampithoe on Hormosira even when transplanted to low shore areas, indicating that other factors, apart from the height on the shore, are also important for this amphipod. Another common amphipod, Hyale maroubrae was generally found in greatest abundances on Hormosira transplanted from mid- to low shore and on Sargassum low on the shore, again indicating the importance of height on the shore. There was no difference in abundance of amphipods on ‘complex’ and ‘simple’ artificial plants indicating that structural complexity (as defined in this study) did not influence these amphipods.     

The habitat and feeding preferences of Littorina obtusata (L.) and L. mariae sacchi et rastelli

The vertical zonation of the flat periwinkles Littorina obtusata (L.) and L. mariae Sacchi et Rastelli is different, but overlaps. At a sheltered location on the Isle of Cumbrae, Scotland, L. obtusata, although most abundant on Fucus vesiculosus L. and Ascophyllum nodosum (L.) Le Jol. on the midshore, ranged over the entire shore from the Pelvetia zone down to the upper limits of Laminaria. Littorina mariae exhibited a much more restricted distribution being associated predominantly with Fucus serratus L. on the lower shore. In laboratory trials with adult plants, flat periwinkles found fucoid algae both more attractive and more palatable than other seaweeds. Littorina obtusata also preferred the reproductive receptacles of fucoid algae to their vegetative tissue. Tiny germlings of a diversity of seaweeds were readily consumed by L. obtusata. In choice experiments germling Ulva lactuca L. was preferred to adult U. lactuca, whereas adult Fucus serratus was preferred to germling F. serratus. Contrasting foraging behaviours between different species of flat periwinkle are discussed in relation to differences in their radular dentition.     

Nutrient Addition to Experimental Rocky Shore Communities Revisited: Delayed Responses, Rapid Recovery

Coastal eutrophication may alter the dominance patterns of marine macroalgae, with potential consequences for the associated fauna and the entire ecosystem. Benthic macroalgae and animals in control and nutrient-enriched mesocosms were monitored to investigate eutrophication-induced changes in rocky shore communities. During a 3-year project, nutrient addition had only minor effects on the community structure, such as increased cover and biomass of green Ulva spp. and increased abundance of certain animal species at high nutrient levels. This study is a 4-year extension of a previously reported project, with 2 extra years of effect studies (altogether 5 years) and a subsequent 2 years for recovery. During the 4th year of nutrient enrichment, the cover of Fucus vesiculosus and Fucus serratus started to decline. In the 5th year, these canopy species crashed and there was an evident take-over by green algae at high nutrient addition levels. The previously observed abundance stimulation for fauna disappeared later in the time series, probably due to the loss of the macroalgal canopy. After less than 2 years on regular seawater, the algal and animal communities had returned to within the range of normal variability. The results indicate that established rocky shore communities of perennial algae with associated fauna are able to persist for several years, even at very high nutrient levels, but that community shifts may suddenly occur if eutrophication continues. They also indicate that rocky shore communities have the ability to return rapidly to natural undisturbed conditions after the termination of nutrient enhancement.     

Relative influence of habitat complexity and proximity to patch edges on seagrass epifaunal communities

Habitat structure at many scales influences faunal communities. Although habitat structure at different scales often covaries, studies rarely examine the relative effects of structure at multiple scales on faunal density and diversity. In shallow‐water seagrass systems, epifaunal density at local scales generally increases with increased habitat structural complexity (e.g. shoot density per unit area). In turn, structural complexity often varies with other aspects of habitat structure at patch scales, such as proximity to patch edges, which itself modifies ecological processes that structure epifaunal communities. We conducted surveys and a manipulative experiment in the eelgrass Zostera marina beds of San Diego Bay, California, USA, to determine (1) whether eelgrass structural complexity, epifaunal density and diversity, and fish (predator) density and diversity vary with proximity to patch edges, and (2) the relative influences of structural complexity, proximity to patch edges and predator presence on epifaunal distribution. Seagrass structural complexity generally increased from patch edges to patch interiors at all sites and in all sampling periods. However, patterns of epifaunal density, diversity, and biomass varied among sites and sampling periods, with density and biomass increasing from patch edges to interiors at some sites and decreasing at others. In the manipulative experiment, we allowed epifauna to colonize sparse or dense artificial seagrass habitat at both the edge and interior of a seagrass patch, and enclosed a subset of experimental units in predator exclusion cages. Overall, proximity to patch edges had a larger influence on epifaunal density and community structure than did structural complexity or predation, with the exception of some common taxa which responded more strongly to either complexity or predator exclusion. Our results emphasize the importance of addressing and evaluating habitat structure at multiple scales to better understand the distribution and interactions of organisms in a particular environment.     

Maintenance of zonation patterns in two species of flat periwinkle,Littorina obtusata and L. mariae

The zonation patterns of Littorina obtusata (L.) and Littorina mariae Sacchi et Rastelli were shown to be quite distinct on a sheltered rocky shore. L. obtusata was found at all the heights sampled; it reached peak numbers at mid shore on the alga Ascophyllum nodosum L. (Le Jol). There was no difference in the tidal height occupied by adults or juveniles; or in the mean size of L. obtusata along the vertical gradient of the shore. In contrast L. mariae occurred exclusively low on the shore, on Fucus serratus L. Translocation of the two species within their respective levels resulted in random movement after 4 days, although initial movements after 1 and 2 days were sometimes directional. Animals transplanted to the normal level of the other species showed directional movement towards their home zone; this was most pronounced after 4 days. There was no difference in the distance moved by the two species, although the distance moved did vary with tidal height, both species moving further at mid shore than low shore. Distances moved by littorinids at replicate areas in the low shore were similar but those at mid shore did vary. There was an interaction between the species and the different tidal heights which revealed that transplanted species moved further than translocated species at the same tidal level. However, this was only significant in the case of L. mariae. It is suggested that the close relationship between the winkles and their host algae may direct the homing behaviour of displaced individuals.     

Direct observations on the sublittoral marine algae of Argyll,Scotland

Summary The distribution with depth of sublittoral marine algae has been investigated at 11 sites on the west coast of Scotland. More than half of the species found were confined to the sublittoral zone. The greatest variety of species was found in shallow water at sites sheltered from excessive turbulence. With increasing depth the number of species found steadily decreased. There was no evidence of a specifically distinct algal flora confined to deeper water. No algae were found deeper than 36 m below ELWS at any site, but the lower limit of algal growth was reduced to 9 m in a turbid water loch and to 3 m at a site where the herbivorous echinoderm Ophiocomina nigra was abundant. Fairly distinct algal communities were found on different substrates. The major communities recognised underwater were algae attached to stable substrata, algae attached to unstable substrata and epiphytic on other algae, especially on the stipes of Laminaria hyperborea. On stable rock in areas subject to water movement L. hyperborea forest was the dominant vegetation whereas on unstable substrata and in sheltered localities L. saccharina was dominant.     

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     

Ecology and dynamics of two intertidal algal communities on the littoral of the island of São Miguel (Azores)

The intertidal benthic algal communities of two sites located on opposite coasts of São Miguel Island (Azores), were studied over a 2-year period (September 1993–September 1995). At both sites (São Roque on the south coast and São Vicente on the north), the littoral region was surveyed from the upper intertidal down to the sublittoral fringe. The survey revealed five distinct zones, with a variable degree of overlapping. The two upper zones were characterized by animals (littorinids and barnacles, respectively). Lower down, algal communities formed three distinct zones: an upper Fucus spiralis/Gelidium microdon association, a more extensive turf zone, and a belt featuring erect or frondose algae. Upper in the eulittoral, the turf was mainly monospecific, and dominated by Caulacanthus ustulatus. The lower eulittoral turf was dominated by articulated coralline algae, the associated species differing between the two sites studied. Mainly erect algae (Pterocladiella, Asparagopsis, etc.) occurred furthest down the shore and extended into the nearby shallow sublittoral. Two intertidal communities were studied at each locality: the upper eulittoral (Caulacanthus turf in São Roque and the Fucus spiralis/Gelidium microdon association at São Vicente), and the lower eulittoral (the coralline turf). The lower littoral communities had a higher algal diversity. A general pattern was observed in the seasonal variation of biomass: the lower levels exhibited the higher values in late summer/early autumn, the period in which the upper levels had the lowest standing crops. Physical factors are proposed to account for this. No significant inter-annual variations could be detected, indicating relatively stable communities, at least on a short-term basis.     

Sessile invertebrate coelobite communities from reefs of Bermuda: Species composition and distribution

Sessile invertebrate coelobite communities attached to the walls and roof of cavities in coralgal reefs on the annular rim of the Bermuda platform near North Rock (4 sites) and in the algal cup reef tract on the south shore (3 sites) have been studied by belt photo-transects and direct sampling. Irradiance measurements reveal a light gradient which appears to exert a strong influence on the composition, relative coverage and zonation of the attached biotic communities. Two main communities are recognised from cluster analysis and relative areal coverage data. Near the cavity entrances is a community dominated by crustose coralline algae, with subsidiary ascidians, demosponges, bryozoans and Foraminifera. Species richness is high and there is total biotic coverage of walls and roof. This community grades laterally into an exclusively animal community characterised by encrusting sponges and Foraminifera, with subsidiary bryozoans and unidentified branching organisms. Coverage varies from 100% to 30%, the substrate often exhibiting high micro-relief from the branching growth forms of the Foraminifera. Species richness is high at North Rock sites, less so on the south shore. The distribution of coelobite species is compared with that described from previous studies in Bermuda, Grand Cayman and Madagascar.     

Littoral macrofauna (secondary) responses to experimental nutrient addition to rocky shore mesocosms and a coastal lagoon

Macrofauna secondary responses to controlled eutrophication within two MARICULT/MAST-III projects, EULIT and COMWEB, are compared. EULIT utilises a nutrient gradient established in eight hard bottom mesocosms, whereas the data from COMWEB originate from a whole-ecosystem study – the case of experimental nutrient addition to Hopavågen lagoon. In both systems, nutrient addition started in May 1998, after initial studies of the background macrofauna communities, thus allowing application of Before-After-Control-Impact-Pairs techniques (BACIP). The main objectives have been to investigate the macrofauna responses to eutrophication in littoral rocky shore ecosystems and to evaluate if similar responses could occur in mesocosm and `field' systems, despite their inherent differences. Apart from a distinct increase in numbers of Littorina littorea L. and some indications of increased abundance of the genus Jaera, no significant changes, caused by 2.5 years of nutrient addition, could be detected in the mesocosm fauna. It is interesting, however, that these two possibly stimulated animal groups have two things in common: (1) both belong to the rather few mobile taxa, which are not flushed out of the system through the mesocosm outlets, (2) both feed on microalgae and green algae in the upper littoral zone, i.e., algal groups that have shown the clearest response to the nutrient addition. In Hopavågen, 1.5 years of nutrient addition only caused modest plant and animal responses in the rocky shore ecosystem, although there was a markedly increased settlement of Mytilus edulis L. at one lagoon site. This increase was probably due to elevated levels of digestible particles (increased phytoplankton production) caused by the nutrient addition. There were also some indications of increased production of filamentous algae at this same lagoon site. All in all, both studies demonstrate only minor responses to increased nutrient levels within the littoral community (both plants and animals). One explanation to this may be that longer time-scales are needed in order for clear-cut changes to occur. Several observations also indicate that wave exposure (significant in both systems) may largely modulate the impact of increased nutrient load on the structure of littoral communities. Another explanation is the high degree of stability among littoral macroalgae-dominated communities with internal biological regulation factors (like grazing) possibly being able to counteract effects.     

Temporal comparison of the composition and zonation of rocky intertidal organisms at Cocos Island National Park, Pacific, Costa Rica

Several biological and physical factors change the rocky shore communities. The desiccation time and the tolerance of the intertidal species produce the vertical zonation. In many studies around the world, a temporal change in this zonation is presented.In Costa Rica, only studies that include temporal trends were carried out in Punta Mala and Montezuma, Pacific coast in 80's. The rocky intertidal of the Cocos Island National Park, Costa Rica were surveyed photographically. The Chatham bay was sampled in three expeditions (January 2007, October 2007 and April 2008). Photos corresponding to 25x25cm quadrats were taken with the goal to determine diversity and composition differences in rocky shore organisms between sampling dates. The Wafer bay was sampled in January and October 2007. The intertidal of Chatham consists of basaltic rock, while Wafer has basaltic and ignimbrite boulders. The main difference between sites were the higher algae cover (erect-frondose forms) and number of organism bands at Chatham bay. Temporal change was not found in the total cover of sessile fauna and autotrophs. The barnacle Tetraclita stalactifera, that occurs above the algal fringe (lower intertidal), was the invertebrate with the highest coverage. The mobile fauna biodiversity presented no significant trend between sampled months. However, the identity of species, their cover and their abundance showed a moderate temporal change. In October 2007, when the sea surface temperature was 23 degrees C the infralittoral zone had an increase in green algae cover. The red algae (crust and erect-frondose forms) were dominant in January and April. The pulmonate limpet, Siphonaria gigas and a bacterial biofilm at mid littoral showed a negative association. The snails of the high littoral and the supralittoral zone showed a temporal change in their abundance, but with contrasting patterns between sites. The temporal variation in the assemblages increased from the supralittoral to the infralittoral possibly due to changes in the water temperature and climatic conditions, that could influence the intertidal zone during the high and low tide, respectively.     

Zooplankton and meiobenthos diversity at the intertidal sandy shores of Jizan and Farasan coastal areas of Red Sea

Distinctive studies were conducted for the identification of meiobenthos and zooplankton at Farasan Islands and Jizan sandy shores. The present work compares the meiobenthos and zooplankton communities at Alhsas sandy shore at Farasan Islands and As-Suways sandy shore at Jizan. Population density, species richness and Shannon-Weiner diversity index were determined for meiobenthos and zooplankton inhabiting both the studied sites. Water criteria; surface water temperature, pH and conductivity were determined for each investigated site. Eleven zooplankton species were defined at Alhsas sandy shore Farasan, nine species were identified at the littoral zone at As-Suways sandy shore, Jizan. Ten meiobenthos species were defined at Alhsas site, Farasan. Only eight meiobenthos species were defined at the intertidal zone of As-Suways site, Jizan. The results were discussed to highlight the effect of water criteria on the spatial distribution of zooplankton and meiobenthos at the investigated sites.     

Spatial Structure of the Insect Community of a Small Dimictic Lake in the Laurentians (Québec)

The distribution of merolimnic benthic insects was studied in Lake Cromwell, a small (9 ha) and shallow (max. depth 9 m) dimictic lake with a summer hypolimnetic warming on the Laurentian highlands of Québec. Communities were described from 22 sets of emergence data. Two hundred and twelve species were recognized of which 153 were Chironomidae (Diptera). Through clustering and ordination techniques, five communities were recognized: one in the flooded shrub zone along the shore, two littoral, one sublittoral and one profundal. Differences between the communities were more quantitative than qualitative. The majority of species inhabited the shore and littoral stations, and only the hardiest reached the deepest zone. There were few species characteristic of the profundal, none of them abundant. Discriminating, dominant, and characteristic species were determined for each community, as were the ubiquitous species. There is a general decrease in species richness, species diversity and population density, and a change in community structure along the depth gradient. A discontinuity at 1–2 m is related to a sharp increase in Chaoboridae in the deeper waters.     

The effects of tidal height,wave-exposure,seasonality and rock-pools on grazing and the distribution of intertidal macroalgae in New South Wales

The effects of grazers (mostly gastropods), height on the shore, wave-exposure, season of the year and the presence of shallow rock-pools on the abundance of intertidal macroalgae were examined on shores at Cape Banks (Botany Bay). At the beginning of each of the four seasons, experimental plots with and without grazers were cleared at four heights on three shores, of increasing exposure to waves, The colonization and growth of algae in these plots were monitored (by measurement of per cent cover and dry wt) for approximately the next 3 months in each season.In grazed plots, foliose algae only grew at the lowest levels on the shores. They were more abundant where wave-action was greater, and during the cooler periods of the year, when growth of the plants was enhanced. Higher on the shore, there was a positive correlation between algal cover in grazed plots and the amount of rainfall during the previous 10 days. In all seasons, there was much greater colonization of foliose algae where grazers were excluded. There was greater algal growth at lower levels on the shore, and where wave-action was stronger. Less algae grew in sheltered areas during warmer times of the year. The major seasonal difference found was the more rapid growth and occupancy of the rock by algae during the cooler seasons of the year.Experimental rock-pools were colonized more rapidly at lower levels on the shore, and during the winter. There was no difference between pools and control (non-pool) areas during winter. During summer, however, there was a greater per cent cover and biomass of algae in pools from which grazers had been excluded than in similar control areas.The results can be interpreted as being due to the greater survival and more rapid growth of algae under conditions of increased moisture, decreased emersion and decreased temperatures and light regimes during low tide (i.e. when physical stresses were reduced). These physical factors were, however, less important to the distribution of the algae than were the effects of grazers. Although few algae were present in any experimental plot, the number of species of algae per plot was considerably reduced in grazed areas.The results confirm that the patterns of vertical distribution and abundance of algae on rock-platforms in New South Wales are primarily the result of the activities of grazers. All algae in the present study were capable of living higher on the shore than they were normally found. Much of the variation along a gradient of wave-exposure, from season to season, and small-scale variation from place to place at any time can be explained by the complex interactions between the activities of grazers, and the effects of variations in the physical environment that influence the recruitment, survival and growth of the algae.