Vertical and seasonal patterns in competition for microalgae between intertidal gastropods

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

Herbivorous caddisflies,macroalgae, and epilithic microalgae: dynamic interactions in a stream grazing system

Summary 1. During the low-flow period (April–October) in sunlit pools of Big Sulphur Creek (northern coastal California), the attached algal community predictably changes from an assemblage dominated by lush, upright Cladophora glomerata filaments in spring and early summer to one dominated by epilithic diatoms and blue-green algae (together=microalgae) in late summer through early autumn. Previous studies in this stream indicated that grazing by the caddisflies Helicopsyche borealis and Gumaga nigricula maintain low algal biomass during the latter part of this period. We used a combination of in situ exclusion/enclosure experiments to examine (1) the separate and combined effects of these grazers on Cladophora and microalgal assemblages, and (2) food preferences, growth, and microdistribution patterns of grazers when offered these different algal foods. 2. Grazers exerted strong but divergent effects on algal assemblages. Selective grazing on Cladophora by G. nigricula greatly accelerated the transition from upright Cladophora to epilithic microalgae, whereas selective grazing on microalgae by H. borealis dramatically reduced biomass of these forms. Grazers were largely ineffective at reducing the non-preferred algal food source (i.e. Cladophora by H. borealis, microalgae by G. nigricula). In the case of each grazer, growth was highest on the preferred algal food. Together, the activity of these grazers produced a low-biomass assemblage dominated by microalgal cells. 3. Removal of the Cladophora overstory by G. nigricula resulted in a three-fold increase in the abundance of epilithic microalgae, the preferred food of H. borealis. Elimination of Cladophora by G. nigricula can increase food availability for H. borealis and, in so doing, can indirectly facilitate the growth of this grazer during food-limited conditions. However, microdistribution of G. nigricula shifts from high overlap with H. borealis in spring and early summer when Cladophora is abundant to low overlap in late summer after Cladophora has been eliminated. This may indicate intense competition between these species for limited epilithic algae, and a concomitant movement by G. nigricula to areas in the stream where food resources are more available.     

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.     

Compound‐specific isotope analysis of benthic foraminifer amino acids suggests microhabitat variability in rocky‐shore environments

The abundance and biomass of benthic foraminifera are high in intertidal rocky‐shore habitats. However, the availability of food to support their high biomass has been poorly studied in these habitats compared to those at seafloor covered by sediments. Previous field and laboratory observations have suggested that there is diversity in the food preferences and modes of life among rocky‐shore benthic foraminifera. In this study, we used the stable nitrogen isotopic composition of amino acids to estimate the trophic position, trophic niche, and feeding strategy of individual foraminifera species. We also characterized the configuration and structure of the endobiotic microalgae in foraminifera using transmission electron microscopy, and we identified the origin of endobionts based on nucleotide sequences. Our results demonstrated a large variation in the trophic positions of different foraminifera from the same habitat, a reflection of endobiotic features and the different modes of life and food preferences of the foraminifera. Foraminifera did not rely solely on exogenous food sources. Some species effectively used organic matter derived from endobionts in the cell cytoplasm. The high biomass and species density of benthic foraminifera found in intertidal rocky‐shore habitats are thus probably maintained by the use of multiple nitrogen resources and by microhabitat segregation among species as a consequence.     

Grazing and UV radiation effects on an Antarctic intertidal microalgal assemblage: a long-term field study

A 15 week field experiment (austral summer Nov–Mar) was carried out in an intertidal hard bottom platform in Antarctica (King George Island). To test whether grazing and ultraviolet radiation (UVR) influenced the succession of a benthic microalgal assemblage, a two-factorial design was used (1) ambient radiation, >280 nm; (2) ambient minus UV-B, >320 nm; (3) ambient minus UVR, >400 nm versus grazer–no grazer). On four sampling occasions microalgae were identified, counted and carbon contents were calculated. The assemblage was dominated by the diatom genera Navicula and Cocconeis. Biomass was generally low in all treatments but was significantly reduced by grazing throughout the experiment. No significant UV effects were found. Grazer absence particularly favoured diatoms of the genus Cocconeis. We conclude that the Antarctic microalgal assemblage was unaffected by present day UVR whereas grazers acted as important drivers on the intertidal microalgal community structure.     

Food supply, grazing activity and growth rate in the limpet Patella vulgata L.: a comparison between exposed and sheltered shores

The limpet Patella vulgata L. is an important microphagous grazer on intertidal rocky shores of north-west Europe, occurring across the wave exposure gradient. Groups of P. vulgata were selected at mid-tide level of two exposed shores and two sheltered, fucoid dominated shores on the Isle of Man, British Isles, and manipulated to equivalent densities and population structure. The level of grazing activity and growth rate were determined over a 1-year period. At the same time, the abundance of epilithic microalgae, measured as the concentration of chlorophyll a, was determined as an estimate of food supply. Microagal abundance showed a seasonal pattern in both exposed and sheltered conditions, with higher levels in winter compared to summer. In both seasons, the microalgal resource was more abundant on the sheltered shore studied. The level of grazing activity in P. vulgata showed a seasonal pattern on the exposed but not the sheltered shores. Averaged over the year, grazing activity on the exposed shores was over double that on sheltered shores. Thus, in sheltered conditions, food supply for limpets was high and grazing activity low; in exposed conditions, food supply was low and grazing activity high. The growth rate of P. vulgata, measured as increase in shell length, showed no significant difference between exposed and sheltered shores. Growth rate was also determined in P. vulgata at natural densities. Although the overall density declined with decreasing exposure to wave action, the density per unit area of grazeable substance was higher in shelter. In these populations, the mean growth rate was over twice as high on exposed compared to sheltered shores.     

Intertidal assemblages on seawalls and vertical rocky shores in Sydney Harbour,Australia

Abstract Understanding the ecological role of artificial structures, such as seawalls, in shallow coastal waters is necessary in order to plan sound strategies of conservation and management of natural habitats. In Sydney Harbour (NSW, Australia), about 50% of the foreshore is made of retaining seawalls This study evaluates the changes caused to natural assemblages of organisms by these structures, by comparing intertidal assemblages between seawalls and vertical rocky shores. The following hypotheses were tested: that assemblages on seawalls would differ from those on rocky shores at mid‐, but not at low‐shore levels; where assemblages differ between habitats, there would be differences in cover/abundances of widespread species; patterns would be consistent among locations and through time; the variability of assemblages at the scales of 10s of cm and metres would differ between seawalls and rocky shores at mid‐ and low‐shore levels. To test these hypotheses, assemblages on seawalls and rocky shores were sampled at three locations, at roughly 4‐monthly intervals, over a period of about 18 months. Results indicated that mid‐shore assemblages on seawalls were different from those on rocky shores, but this was not the case at low‐shore levels. Few taxa were unique to either habitat. Cover of common species of algae and sessile animals and abundances of mobile grazers were variable with few consistent patterns. Variability at the scales sampled differed between habitats and heights on the shore. Seawalls and rocky shores, in general, supported a similar suite of species, but patterns of abundance and variation differed among locations and from height to height in each habitat. The implications of these findings for the future management of seawalls are briefly considered.     

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     

Microalgal mediation of ripple mobility

The interaction between physical and biological factors responsible for the cessation of ripple migration on a sandy intertidal flat was examined during a microalgal bloom period in late winter/early spring, as part of a wider study into the biostabilisation of intertidal sediments. Ripple positions and ripple geometry were monitored, and surface sediment was sampled, at weekly intervals over a 5-week period. Ripples remained in the same position for at least 4 weeks, during which time there was a progressive reduction in bedform height (smoothing) and deposition of some 1.5 cm sediment, mainly in the ripple troughs (surface levelling). The mean chlorophyll a (chl a) sediment content was 6.0 microg gDW(-1) (DW: dry weight) (0-1 mm depth fraction), with a maximum value of 7.4 microg gDW(-1) half way through the bloom. Mean colloidal-S carbohydrate (S: saline extraction) content was 131 microg GE gDW(-1) (GE: glucose equivalent) (0-1 mm), with a maximum of 261 microg GE gDW(-1 )towards the end of the bloom. Important accessory pigments were peridinin (indicative of dinophytes) and fucoxanthin (diatoms). Stepwise multiple regression showed that peridinin was the best predictor of chl a. For the first time, in situ evidence for the mediation of (wave) ripple migration by microalgae is provided. Results indicate that diatoms, and quite possibly dinophytes, can have a significant effect on intertidal flat ripple mobility on a temporal scale of weeks. In addition, microalgal effects appear capable of effecting a reduction in bed roughness on a spatial scale of up to 10(-2 )m, with a subsequent reduction in bottom stress and bed erodability. It is suggested that a unique combination of environmental conditions, in conjunction with the microalgal bloom(s), promoted the initial cessation of ripple movement, and that stationary-phase, diatom-derived extracellular polymeric substances (EPS) (and possibly dinophyte-derived EPS) may have prolonged the condition. It is reasonable to suppose that ripple stabilisation by similar processes may have contributed to ripple mark preservation in the geological record. A conceptual model of sandy intertidal flat processes is presented, illustrating two conditions: (i) a low EPS/microalgae sediment content with low ripple stabilisation and preservation potential; and (ii) a high EPS/microalgae content with higher preservation potential.     

Grazing dynamics in intertidal rockpools: Connectivity of microhabitats

Differences between rockpool and emergent rock communities are often attributed to their contrasting physical conditions. However, differences in grazing pressure between rockpools and open rock could also exert an important structuring role. Greater densities and/or the lack of tidal constraints on foraging may allow grazing intensity to be greater in rockpools. Here, wax discs were deployed to compare grazing intensity between rockpool and emergent rock habitats at each of three tidal heights on a moderately exposed shore in SW England. Grazing intensity was then examined in relation to herbivore density. Grazing intensity in pools was twice that on emergent rock, despite a lower density of herbivores in the rockpools. Of these herbivores, patellid limpets are the dominant grazers on rocky shores throughout the NE Atlantic and are recognised to have a major role in structuring intertidal communities. Thus, subsequent experiments focussed on the influence of limpets in determining the differences in consumer pressure between rockpools and emergent rock. Three alternative explanations were considered: (1) the effect of continuous immersion on grazing intensity in rockpools; (2) differences in limpet species abundance between the two habitats; (3) movement of limpets from emergent rock into pools to feed. The level of grazing pressure exerted by Patella ulyssiponensis (Gmelin), the predominant species living constantly immersed in rockpools, was similar to that of P. vulgata (Linnaeus) which is predominantly found on emergent rock. P. vulgata were observed moving from emergent rock into rockpools during high tide. Manipulative experiments confirmed that these foraging excursions resulted in a 2-fold increase in grazing intensity in the pools. Grazing activity of P. vulgata in rockpools was not consistent between sites and may be influenced by differences in wave exposure and/or the abundance of microbial resources. Elevated consumer pressure in rockpools may be an important factor influencing algal assemblages and probably explains the predominance of grazer resistant-species in these pools.     

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.     

Neutral processes and species sorting in benthic microalgal community assembly: effects of tidal resuspension

Benthic microalgae (BMA) provide vital food resources for heterotrophs and stabilize sediments with their extracellular secretions. A central goal in ecology is to understand how processes such as species interactions and dispersal, contribute to observed patterns of species abundance and distribution. Our objectives were to assess the effects of sediment resuspension on microalgal community structure. We tested whether taxa‐abundance distributions could be predicted using neutral community models (NCMs) and also specific hypotheses about passive migration: (i) As migration decreases in sediment patches, BMA α‐diversity will decrease, and (ii) As migration decreases, BMA community dissimilarity (β‐diversity) will increase. Co‐occurrence indices (checkerboard score and variance ratio) were also computed to test for deterministic factors, such as competition and niche differentiation, in shaping communities. Two intertidal sites (mudflat and sand bar) differing in resuspension regime were sampled throughout the tidal cycle. Fluorometry and denaturing gradient gel electrophoresis were utilized to investigate diatom community structure. Observed taxa‐abundances fit those predicted from NCMs reasonably well (R² of 0.68–0.93), although comparisons of observed local communities to artificial randomly assembled communities rejected the null hypothesis that diatom communities were assembled solely by stochastic processes. No co‐occurrence tests indicated a significant role for competitive exclusion or niche partitioning in microalgal community assembly. In general, predictions about relationships between migration and species diversity were supported for local community dynamics. BMA at low tide (lowest migration) exhibited reduced α‐diversity as compared to periods of immersion at both mudflat and sand bar sites. β‐diversity was higher during low tide emersion on the mudflat, but did not differ temporally at the sand bar site. In between‐site metacommunity comparisons, low‐ and high‐resuspension sites exhibited distinct community compositions while the low‐energy mudflats contained higher microalgal biomass and greater α‐diversity. To our knowledge this is the first study to test the relevance of neutral processes in structuring marine microalgal communities. Our results demonstrate a prominent role for stochastic factors in structuring local BMA community assembly, although unidentified nonrandom processes also appear to play some role. High passive migration, in particular, appears to help maintain species diversity and structure communities in both sand and muddy habitats.     

The effects of grazing by gastropods and physical factors on the upper limits of distribution of intertidal macroalgae

Summary The cover of foliose algae is sparse to non-existent above a low-level algal zone on many shores in N.S.W., except in rock-pools. Above this algal zone, encrusting algae, mostly Hildenbrandia prototypus, occupy most of the primary substratum on sheltered shores. Experimental manipulations at midtidal levels were used to test hypotheses about the effects of grazing by molluses and of physical factors during low tide on this pattern of algal community structure.Fences and cages were used to exclude grazers: molluscs grazed under roofs and in open areas. Cages and roofs provided shade, and decreased the harshness of the environment during low tide: fences and open areas had the normal environmental regime.In the absence of grazers, rapid colonization of Ulva and slower colonization by other foliose algae occurred in all experimental areas. The rate of colonization by Ulva sporelings was initially retarded on existing encrusting algae, but after a few months, cover of Ulva equalled that on cleared rock.Most species of algae only grew to maturity inside cages, and remained as a turf of sporelings inside fences. No foliose algae grew to a visible size in open, grazed areas. Grazing thus prevents the establishment of foliose algae above their normal upper limit on the shore, but the effects of physical factors during low tide prevent the growth of algae which become established when grazers are removed. Physical factors thus limit the abundance of foliose algae at mid-tidal levels.The recolonization of cleared areas by Hildenbrandia was not affected by the presence of a turf of sporelings, nor by the shade cast by roofs, but was retarded in cages where mature algae formed a canopy. Even under such a canopy, Hildenbrandia eventually covered as much primary substratum as in open, grazed areas. This encrusting alga is able to escape from the effects of grazing by having a tough thallus, and by its vegetative growth which allows individual plants to cover a lot of substratum, and by the tendency for new individuals to start growing from small cracks and pits in the rock, which are apparently inaccessible to the grazers.Mature foliose algae are removed from the substratum by waves, and many individual plants died during periods of hot weather. Sporelings in a turf were eliminated, after experimental fences were removed, by the combined effects of macroalgal grazers, which invaded the areas, and microalgal grarers which ate the turt from the edges inwards.The results obtained here are discussed with respect to other studies on limits to distribution of intertidal macroalgae, and the role of grazing in the diversity and structure of intertidal algal communities. Some problems of these experimental treatments are also discussed.     

Estuarine intertidal sandflat benthic microalgal responses to in situ and mesocosm nitrogen additions

Benthic microalgal communities are important components of estuarine food webs and make substantial contributions to coastal materials cycling. Nitrogen is generally the limiting factor for marine primary production; however other factors can limit benthic primary producers because of their access to the additional nutrients found in sediment porewater. Field and laboratory experiments were conducted to test the hypothesis that water column nitrogen supply affects estuarine sandflat benthic microalgal community structure and function. Our field and mesocosm experiments assessed changes at both the population and functional group levels. Simulated water column nitrogen additions increased maximum community photosynthesis in most cases (Pb_max from photosynthesis vs. irradiance curves). Additional changes that resulted from nitrogen additions were decreases in porewater phosphate, increases in porewater ammonium, shifts in community composition from N₂ fixing cyanobacteria toward diatoms, and detectable, though not statistically significant increases in biomass (as chlorophyll a). Results from field and laboratory experiments were quite similar, suggesting that laboratory experiments support accurate predictions of the response of intertidal benthic microalgae to changes in water column nutrient conditions.     

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.     

Large-scale coexistence and small-scale segregation of key species on rocky shores

Biodiversity is defined by the scales at which organisms coexist. Coexistence at larger spatial scales may be underpinned by segregation at smaller spatial scales or temporal variability in behaviour and/or distribution. Limpets of the genus Patella are keystone grazers controlling the growth of macro-algae on many rocky shores. This study examined the distribution and abundance of P. vulgata and P. ulyssiponensis in relation to habitat and to each other at a range of spatial scales. Limpets were sampled on four shores on each of the east and west coasts of Ireland, in two different transects (10 m apart) at each of three shore heights in winter 2003 and summer 2004. P. vulgata and P. ulyssiponensis coexist around the coast of Ireland, but on shores where they co-occur, the pattern of coexistence begins to break down at the level of shore height: P. ulyssiponensis were more abundant on the low-shore than mid-shore, and P. vulgata were distributed evenly throughout the low and the mid-shore. At the level of habitat (pools vs. open rock), the two species exhibited a high degree of segregation: P. vulgata of all sizes were more common on open rock and P. ulyssiponensis of all sizes were more common in pools. The presence or absence of P. ulyssiponensis had no effect on the distribution of juvenile P. vulgata with respect to pools. Unexpectedly, however, a greater proportion of adult P. vulgata were found in pools at sites where P. ulyssiponensis were present in the pools than at sites where they were absent. The results of this study emphasize the need for multiple-scale analyses of the distribution and abundance of organisms to understand the organisation of natural assemblages. Handling editor: K. Martens     

Benthic microalgae in coral reef sediments of the southern Great Barrier Reef,Australia

The abundance and productivity of benthic microalgae in coral reef sediments are poorly known compared with other, more conspicuous (e.g. coral zooxanthellae, macroalgae) primary producers of coral reef habitats. A survey of the distribution, biomass, and productivity of benthic microalgae on a platform reef flat and in a cross-shelf transect in the southern Great Barrier Reef indicated that benthic microalgae are ubiquitous, abundant (up to 995.0 mg chlorophyll (chl) a m^–2), and productive (up to 110 mg O₂ m^–2 h^–1) components of the reef ecosystem. Concentrations of benthic microalgae, expressed as chlorophyll a per surface area, were approximately 100-fold greater than the integrated water column concentrations of microalgae throughout the region. Benthic microalgal biomass was greater on the shallow water platform reef than in the deeper waters of the cross-shelf transect. In both areas the benthic microalgal communities had a similar composition, dominated by pennate diatoms, dinoflagellates, and cyanobacteria. Benthic microalgal populations were potentially nutrient-limited, based on responses to nitrogen and phosphorus enrichments in short-term (7-day) microcosm experiments. Benthic microalgal productivity, measured by O₂ evolution, indicated productive communities responsive to light and nutrient availability. The benthic microalgal concentrations observed (92–995 mg chl a m^–2) were high relative to other reports, particularly compared with temperate regions. This abundance of productive plants in both reef and shelf sediments in the southern Great Barrier Reef suggests that benthic microalgae are key components of coral reef ecosystems.Communicated by Environmental Editor, B.C. Hatcher     

Gradients of intertidal primary productivity around the coast of South Africa and their relationships with consumer biomass

The structure of rocky intertidal communities may be influenced by large-scale patterns of productivity. In this study we examine the in situ rates of production by intertidal epilithic microalgae (chlorophyll a production per unit area per month), intertidal nutrient concentrations (nitrates, nitrites, phosphates and silicates), and standing stocks of different functional-form groups of macroalgae around the South African coast, and their relationships to consumer biomass. Clear gradients of in situ intertidal primary production and nutrient concentrations were recorded around the South African coast, values being highest on the west coast, intermediate on the south and lowest on the east coast. Primary production by intertidal epilithic microalgae was correlated with nutrient availability and could also be related to nearshore phytoplankton production. The dominance patterns of different functional forms of macroalgae changed around the coast, with foliose algae prevalent on the west coast and coralline algae on the east coast. However, overall macroalgal standing stocks did not reflect the productivity gradient, being equally high on the east and west coasts, and low in the south. Positive relationships existed between the average biomass of intertidal intertebrate consumers (grazers and filter-feeders) and intertidal productivity, although only the grazers were directly connected to in situ production by epilithic intertidal microalgae. The maximum body size of a widely distributed limpet, Patella granularis, was also positively correlated with level of in situ primary production. The maximal values of biomass attained by intertidal filter-feeders were not related to intertidal primary production, and were relatively constant around the coast. At a local scale, filter-feeder biomass is known to be strongly influenced by wave action. This implies that the local-scale water movements over-ride any effects that large-scale gradients of primary production may have on filter-feeders. The large-scale gradient in intertidal productivity around the coast is thus strongly linked with grazer biomass and individual body size, but any effect it has on filter-feeder biomass seems subsidiary to the local effects of wave action.     

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

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

湛江沿海海洋微藻及其多糖、脂类和蛋白藻株多样性研究

从硇洲岛和徐闻珊瑚礁自然保护区潮间带采集的海水和沉积物样品中分离培养海洋微藻, 筛选其中富含多糖、脂类或蛋白质的藻株。采用形态学观察、18S rDNA序列比较及其系统发育分析法, 对分离培养的海洋微藻及其富含多糖、脂类或蛋白质的藻株进行分类鉴定和生物多样性分析。分离、培养、鉴定并储藏了189株海洋微藻, 归属于65个种, 分布于硅藻门(Bacillariophyta)、绿藻门(Chlorophyta)、定鞭藻门(Haptophyta)和红藻门(Rhodophyta)的9纲、25目、30个科、38个属; 其中多糖含量较高的46株海洋微藻, 分布于25个种, 20个属; 脂类含量较高的46株海洋微藻, 分布于32个种, 15个属; 蛋白含量较高的46株海洋微藻, 分布于28个种, 18个属。结果表明硇洲岛和徐闻珊瑚礁自然保护区潮间带可培养的海洋微藻及其富含多糖、脂类和蛋白质藻株的物种多样性丰富, 在新型药物、活性天然产物、功能食品和饲料及其添加剂的发掘等方面具有良好前景。