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.     

Analysis of competitive interactions between the limpets Patella depressa Pennant and Patella vulgata L. on the northern coast of Portugal

Inter- and intraspecific competitive interactions among the species Patella depressa Pennant and P. vulgata L. were analysed on the northern coast of Portugal, where both species co-occur in similar proportions. Increased (×2, ×4), normal and decreased (×1/2 intraspecific only) densities of limpets were used to test the effects of competition on the growth and mortality of the limpets, and competitive interactions between the different species. Fenced plots of 25×25 cm enclosing marked limpets at various densities were set up at a mid tidal level on the shore. Twelve treatments with three replicates of different combinations of densities and species were established. Mortality was recorded every 15 days and length was measured monthly during the 5 months of the experiment. At the end of the experiment, limpets were collected for biometry, sex determination and gonad staging. Both species of grazing molluscs showed increased mortality and reduced size and weight in increased density treatments. Limpets in decreased density treatments showed lower mortality and higher size. There were no significant differences between the effect of P. vulgata on P. depressa and the effect of P. depressa on P. vulgata on mortality and length. There were, however, decreases in weight due to intraspecific effects for both species. For P. vulgata, interspecific effects were much less than intraspecific effects and for P. depressa, the interspecific effects depended on density. The analysis of the gonad stage at the end of the experiment showed that the lower stages of development were most prevalent at increased densities and the occurrence of neuter individuals only in increased densities single species plots corroborated the evidence of a stronger intraspecific interaction. The availability of food was indirectly assessed by determination of chlorophyll concentration with spectrophotometric analysis of rock chips. Microalgal food assessment in this experiment, however, did not show any clear trends. The results were compared with previous studies of competition in grazing molluscs and discussed in relation to the limits of distribution of P. vulgata on the Portuguese coast.     

Effects of grazer identity on the probability of escapes by a canopy-forming macroalga

Through their grazing activities limpets have an important role in controlling macroalgal abundance and as a result the structure and dynamics of rocky shore assemblages. Using two congeneric limpet species, with different biogeographic distributions, and whose ranges are expected to alter with climatic warming, we separated the magnitude of their grazing activity over time and the subsequent consequence for macroalgal growth.The northern/boreal limpet, Patella vulgata (L.), consistently grazed more than the southern/lusitanian limpet, P. depressa (Pennant), particularly during spring and summer when P. depressa was reproductively active. Individuals of Fucus vesiculosus (L.) that settled during this time were able to grow to a size where they escaped the grazing activities of P. depressa, resulting in mature F. vesiculosus being present in all P. depressa treatments. In contrast, P. vulgata, which was not reproductively active during this period, exhibited no reduction in its grazing activity and prevented macroalgae from growing in experimental treatments. It therefore appears that P. vulgata has a stronger role, than P. depressa, in controlling macroalgal abundance on shores of southwest Britain.We present a conceptual model highlighting the direct and indirect interactions between these two limpet species and F. vesiculosis. If as predicted, under current warming scenarios, P. depressa becomes the dominant limpet on shores of southwest Britain there will be subsequent changes in rocky shore community structure and ecosystem functioning. Our research emphasises that even closely related species with similar ecological niches can exhibit different behaviours that fundamentally alter their biological interactions with other organisms leading to idiosyncratic responses to predicted changes in climate.     

Positive and Negative Effects of Habitat-Forming Algae on Survival,Growth and Intra-Specific Competition of Limpets

Understanding the effects of environmental change on the distribution and abundance of strongly interacting organisms, such as intertidal macroalgae and their grazers, needs a thorough knowledge of their underpinning ecological relationships. Control of grazer-plant interactions is bi-directional on northwestern European coasts: grazing by limpets structures populations of macroalgae, while macroalgae provide habitat and food for limpets. Scottish shores dominated by the macroalga Fucus vesiculosus support lower densities and larger sizes of limpets Patella vulgata than shores with less Fucus. These patterns may be due to differences in inter-size-class competitive interactions of limpets among shores with different covers of Fucus. To examine this model, densities of small and large limpets were manipulated in plots with and without Fucus. Amounts of biofilm were measured in each plot. The presence of Fucus increased survival but hindered growth of small (15 mm TL) limpets, which were negatively affected by the presence of large limpets (31 mm TL). In contrast, large limpets were not affected by the presence of Fucus or of small limpets. This suggests the occurrence of asymmetric inter-size-class competition, which was influenced by the presence of macroalgae. Macroalgae and increased densities of limpets did not influence amounts of biofilm. Our findings highlight the role of interactions among organisms in generating ecological responses to environmental change.     

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.     

Seasonal variation in epifaunal communities associated with giant kelp (Macrocystis pyrifera) at an upwelling‐dominated site

Kelp forests are highly productive and species‐rich benthic ecosystems in temperate regions that provide biogenic habitat for numerous associated species. Diverse epifaunal communities inhabit kelp sporophytes and are subject to variations in the physical environment and to changes experienced by the kelp habitat itself. We assessed seasonal variations in epifaunal invertebrate communities inhabiting giant kelps, Macrocystis pyrifera, and their effects on this seaweed. Six seasonal samplings were conducted over a year at an upwelling‐dominated site in northern‐central Chile where physical conditions are known to fluctuate temporally. More than 30 taxa were identified, among which peracarid crustaceans stood out in both diversity and abundance. Species richness and abundance differed among sporophyte sections (holdfast and fronds) and throughout the year. The frond community was dominated by two grazers (the amphipod Peramphithoe femorata and the isopod Amphoroidea typa), while suspension feeders, grazers, and omnivores (the amphipod Aora typica, the isopod Limnoria quadripunctata, and polychaetes) dominated the holdfasts. Abundances of the dominant species fluctuated throughout the year but patterns of variation differed among species. The most abundant grazer (P. femorata) had highest densities in summer, while the less abundant grazer (A. typa) reached its peak densities in winter. Interestingly, the area of kelp damaged by grazers was highest in autumn and early winter, suggesting that grazing impacts accumulate during periods of low kelp growth, which can thus be considered as ‘vestiges of herbivory past.’ Among the factors determining the observed seasonal patterns, strong variability of environmental conditions, reproductive cycles of associated fauna, and predation by fishes vary in importance. Our results suggest that during spring and early summer, bottom‐up processes shape the community structure of organisms inhabiting large perennial seaweeds, whereas during late summer and autumn, top‐down processes are more important.     

Molluscan grazing and macroalgal zonation on a rocky intertidal platform at Perth,Western Australia

Abstract Intertidal limestone platforms off Perth show a characteristic pattern of algal zonation, with dense macroalgal beds nearshore bounded by a ‘barren zone’ along the seaward edge. Abalone (Haliotis roei) and several species of limpets and chitons are abundant in the barren zone, which is generally devoid of non-coralline macroalgae. The relative importance of abalone versus limpets and chitons in limiting macroalgal abundance in the barren zone was evaluated by manipulating the presence of each group in a factorial experiment. Percentage algal cover was measured photographically in 0.25m² plots at 1–2 month intervals for 9 months. Mean algal cover (mainly the foliose green alga, Ulva rigida) was highest in plots where all grazers were excluded (77–99%), intermediate where only limpets and chitons were excluded (37–85%), and lowest where only abalone were excluded (4–30%) or where no grazers were excluded (2–19%). The effect of limpets and chitons accounted for 55–89% of the variance in total algal cover, whereas the effect of abalone generally accounted for <10% of the variance. Similar results were obtained in terms of the biomass of Ulva rigida at the end of the experiment. Haliotis roei are relatively large and sedentary herbivores, feeding mainly on drift algae. Their effect on algal abundance was mediated both through pre-emption of space, which might otherwise be colonized by algae, and by grazing around their home scar. Limpets and chitons are smaller than abalone, but were much more abundant. Intensive grazing of the reef surface by limpets and chitons precluded the establishment of non-coralline macroalgae, even where abalone were absent.     

Limpet grazing on a physically stressful Patagonian rocky shore

Many theories of consumer control of communities have come from studies conducted in relatively benign, temperate zone rocky intertidal systems. Here, we examine gastropod grazing and the maintenance of bare space on a dry, wind-swept rocky shore of Patagonia, Argentina. Two limpet species are the primary intertidal grazers. Siphonaria lessoni dominates mid and high intertidal zones, while Nacella magellanica dominates the lower zone. In all zones, limpet densities are positively correlated with bare space and the occurrence of cracks. Tethering experiments revealed that: (1) physical stress sets the upper distribution limit of both limpets, (2) predators, such as oyster catchers, regulate Nacella populations and may restrict them to cracks and vertical surfaces, and (3) desiccation stress appears to drive similar crack distribution patterns of Siphonaria in the upper intertidal. Experimental removal of limpets in each intertidal zone indicated that limpets have: (1) no detectable effect in the high intertidal where physical forces dominate community organization, (2) weak impacts at mid-elevations as grazing only limited the abundance of fleshy algae with physical forces again dominating community structure, and (3) relatively stronger, but still weak impacts in the low zone. These results suggest that grazing impacts on Argentine rocky shores are weak in comparison to the physical stresses (e.g. high winds, low humidity) that largely determine structure in this system. The dominance of physical forcing in this system occurs despite having similar grazer densities to other temperate, but comparatively wet, rocky shorelines (e.g. British Isles) where top-down control is strong.     

Impacts of elevated temperature on the growth and functioning of decomposer fungi are influenced by grazing collembola

Elevated temperature has potential to influence the biological mechanisms regulating ecosystem–atmosphere carbon exchange. The relationship between warming and heterotrophic microbial respiration remains poorly understood, not least in terms of the differential sensitivity of microbial groups to temperature and the complexity of interactions with other biota. Cord‐forming basidiomycete fungi are dominant primary decomposers in temperate woodland. Decomposition rates are determined by the composition of the decomposer community, ecophysiological relationships between these fungi and abiotic variables and interactions with other organisms. Amongst the latter, a major determinant is the balance between mycelial growth and removal by soil invertebrate grazers, which can themselves be affected by elevated temperature. We investigated the impact of elevated temperature on fungal foraging and decomposition of beech (Fagus sylvatica) wood in soil microcosms to which the invertebrate grazers, Folsomia candida and Protophorura armata (Collembola), were added in factorial combinations with five basidiomycete fungi. Species‐specific impacts on mycelial development and function resulted from differential sensitivity of fungi to warming and grazing. Temperature impacts on collembola abundance were resource‐specific, causing increased grazing pressure by both species, but on different fungi. Grazing often counteracted warming‐induced stimulation of mycelial growth, but occasionally amplified the temperature effect, with implications for colonization rates of new resources. High grazing pressure did not prevent increased fungal‐mediated decomposition of colonized wood, as fungi utilized more resource‐derived energy to maintain explorative growth. Impacts of elevated temperature on decomposition are likely to depend on local composition of the fungal and invertebrate decomposer community.     

A continental scale evaluation of the role of limpet grazing on rocky shores

It is critical for our knowledge of biodiversity and ecosystem processes to understand how individual species contribute to ecosystem processes and how these contributions vary in space and time. We used a manipulative field experiment in five locations over 17° of latitude [from southern Portugal to the Isle of Man (British Isles)] to determine the relative response of rocky intertidal algal assemblages released from control by the grazing of limpets. Response ratios showed that when limpets were removed there was a trend of effects from north to south. In the north, grazing had a strong effect on algal assemblages, but removing grazers reduced spatial variability in assemblages. In the south, the effect of limpet grazing was far weaker and removal of grazers had a much reduced impact on spatial variability. Here we show a clear trophic control of an ecosystem in that grazing by limpets not only determines macroalgal abundance overall but also modifies ecosystem stability via variability in cover of algae.     

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.     

Effects of interactions between algae and grazing gastropods on the structure of a low-shore intertidal algal community

Summary At low levels on shores in New South Wales, foliose algae are abundant and often occupy all substrata; microalgal grazing gastropods are rare or absent. At higher levels, foliose algae are sparse or absent and grazing gastropods are abundant. Hypotheses for the causes of the lower vertical limits of distribution of these grazers include the effects of increased predation or the deleterious physiological effects of increased period of submergence at lower levels on the shore. Alternatively, the presence of the algae, because they occupy space and deprive the grazers of substratum for feeding, may prevent the downward movement, or survival of the grazers at low levels. Under the first two of these hypotheses, algae are able to colonize and grow in low-shore areas as an indirect result of factors which remove grazers. Under the third hypothesis, the algae are directly responsible for the lack of grazers.Experimental clearings of the low-shore algae and introductions of the mid-shore limpets Cellana tramoserica and Siphonaria denticulata were used to test these hypotheses. C. tramoserica grazes microalgae and removes them from the substratum, preventing colonization. S. denticulata, in contrust, crops the algae, leaving a visible cover of algae on the substratum, which can grow rapidly. Because of its method of feeding, S. denticulata had no measurable impact on the rates of colonization, nor on the dry weights of algae, compared with those of ungrazed areas. C. tramoserica could keep cleared areas tree from foliose algae, but only when the limpets were mainfained in great density (10 per 900 cm²). They were less effective where wave-action was greater.Neither species of limpets could survive when placed onto beds of mature algae, because they had no substratum on which to cling and were swept away by the waves. C. tramoserica did not invade clearings below their lower limit of distribution where they had to move over a bed of foliose algae. Few C. tramoserica moved directly downshore into cleared areas. When placed on bare rock within low-shore beds of algae of different ages, S. denticulata remained amongst the algae and maintained their tissue-weights. Few C. tramoserica remained in areas with well-developed algae, compared with areas having sparse algal growth. Those Cellana which remained amongst well-developed algae lost weight, whereas limpets in areas with less algal growth mammtained their weights. In experimental cages in low-shore beds of algae, where the limpets were inaccessible to potential predators, C. tramoserica lost weight and died. On cleared areas they survived for many weeks, but lost weight and died as algae grew and covered the substratum. In the absence of predation, the micro-algal grazer C. tramoserica could not survive in lowshore areas because algae grew too fast and occupied the substratum, making it inaccessible for the limpets to graze; the algae, once grown beyond small sporelings, are not a suitable food-source for C. tramoserica, and the loss of weight and death of these limpets is attributable to starvation.The lower limit of distribution of C. tramoserica is not due to the direct effects of physical factors associated with prolonged submersion, nor to the impact of predators, but is apparently determined by the presence of rapidly growing, extensive beds of foliose algae at low levels on the shore. The cause of the limit of distribution of S. denticulata is not yet known and predation may prove to be important. Removal of S. denticulata from low-shore algal beds would not, however, affect the domination of substrata by algae. Grazing by S. denticulata at very great density had no effect on algal cover nor weight. In the intertidal community studied, the persistence of a low-shore algal zone, bounded above by abundant grazers is not influenced by the activities of predators, but is a direct result of interactions between the grazers and the algae.     

Geographical variation in the breeding cycles and recruitment of Patella spp.

The effect of geographical differences in breeding cycles on the recruitment variation of the limpets — Patella vulgata L., P. aspera Röding and (to a limited extent) P. depressa Pennant — has been studied over much of their range in Britain and NW. Europe. In spite of considerable annual and local variation in recruitment success, broader patterns can be distinguished, which can be linked to spawning times and factors affecting the survival of newly-settled spat. The breeding cycles of P. vulgata and P. aspera differ across their ranges in that, in both species, spawning begins, and gametogenesis ends, earlier in the north and east than in the south and west. The cause of these differences can be correlated with geographical and annual differences in sea temperature over the potential breeding periods, and can be related to the regional incidence of conditions found experimentally to be necessary for successful settlement and survival of spat during a critical stage of their growth. The significance of this temperature window in determining the littoral and geographical distribution of the species is discussed.     

The effect of zooplankton grazing on estuarine blooms of the toxic dinoflagellate Gonyaulax tamarensis

A series of short-term ^{in situ} experiments was conducted intwo Cape Cod embayments to estimate mortality rates of the toxicdinoflagellate, Gonyaulax tamarensis, resulting from grazingby zooplankton. Rates of grazing by the whole zooplankton communityand by specific zooplankton populations were measured at variouspoints in the G. tamarensis bloom cycle. The planktonic larvaeof the spionid polychaete Polydora ligni and the tintinnid ciliateFavella sp. were important grazers in the systems studied. Gonyaulax-specificclearance rates effected by Polydora ranged from 0.02 to 0.5ml individual^–1 h^–1; for Favella the range was aboutan order to magnitude lower. Peak population densities wereclose to 900 and 400 individuals 1^–1 for P. ligni andFavella, respectively. Whether measured directly or predictedas the product of individual clearance rates and numerical abundance,rates of grazing were often higher than estimated algal divisionrates in years when blooms failed to develop. A simulation modelcorroborated the results of the field study, demonstrating thatgrazing can be a significant source of mortality during blooms,and can suppress bloom development when grazers are abundant.     

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.     

Aerial and underwater metabolism of <Emphasis Type="Italic">Patella vulgata</Emphasis> L.: comparison of three intertidal levels

Intertidal molluscs are known to possess specific respiratory organs that permit aerial breathing during emersion. Patella vulgata is a widely distributed intertidal species found from low-water spring tide to high-water neap tidal level. In order to determine metabolic adaptations to habitat, carbon fluxes associated with respiration and calcification of P. vulgata living at high-shore, middle-shore and low-shore levels were compared. Seasonal aerial respiration was measured using an infrared gas analyser; seasonal underwater respiration and calcification were calculated from dissolved inorganic carbon and total alkalinity. P. vulgata showed net CaCO₃ deposition at all seasons, although the high-shore level limpet annual calcification rate was relatively low due to longer air exposure. Both aerial and underwater respiration rates were highly correlated with seasonal temperature variations and followed the vertical shore gradient, with stronger fluxes for low-shore tidal level limpets and lower fluxes for high-shore level limpets that must limit energy expenditure. P. vulgata appears to be well adapted to aerial exposure, with average hourly respiration fluxes stronger in air than in water. This study demonstrates that P. vulgata calcification and respiration are reduced in upper shore levels and are important factors determining the upper distribution limit of the species.     

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

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

Biomanipulation and food-web dynamics — the importance of seasonal stability

Responses of phytoplankton biomass were monitored in pelagic enclosures subjected to manipulations with nutrients (+N/P), planktivore roach (Rutilus rutilus) and large grazers (Daphnia) in 18 bags during spring, summer and autumn in mesotrophic Lake Gjersjøen. In general, the seasonal effects on phytoplankton biomass were more marked than the effects of biomanipulation. Primary top-down effects of fish on zooplankton were conspicuous in all bags, whereas control of phytoplankton growth by grazing was observed only in the nutrient-limited summer situation. The effect of nutrient additions was pronounced in summer, less in spring and autumn; additions of fish gave the most pronounced effect in spring. The phytoplankton/zooplankton biomass ratio remained high (10–100) in bags with fish, with the highest ratios in combination with fertilization. The ratio decreased in bags without fish to<2 in most bags, but a real grazing control was only observed in bags with addition ofDaphnia. No direct grazing effects could be observed on the absolute or relative biomass of cyanobacteria (mainlyOscillatoria agardhii). The share of cyanobacteria in total phytoplankton biomass was lowest in summer (7–26%), higher in spring (39–63%) and more than 90% in the autumn experiment. The development of the cyanobacterial biomass was rather synchronous in all bags in all the three experiments. A high biomass ofDaphnia gave no increase in the pool of dissolved nutrients in spring, a slight increase in summer and a pronounced increase in autumn. While a strong decrease in the P/C-cell quota of the phytoplankton was observed from spring to autumn, no effect of grazing or nutrient release could be related to this P/C-status. The experiments indicate that such systems, with high and stable densities of inedible cyanobacteria, are rather insensitive to short-term (3–4 weeks) biomanipulation efforts. This is supported by observations on the long-term development of the lake.

     

Activity patterns in Siphonaria populations: Heading choice and the effects of size and grazing interval

Directional choice and the effects of size and grazing frequency on activity were studied in the pulmonate limpets Siphonaria normalis Gould at Enewetak Atoll and S. alternata (Say) in Bermuda. Whenever individual limpets made a second grazing trip on the same tide, they avoided moving in the direction taken on the first trip. When a longer time interval separated two trips, heading choice on the second excursion was random. Headings within groups of siphonariid limpets active on the same tide were also randomly distributed.Infrequent grazers moved further per trip than their less sedentary neighbors at both sites in Bermuda; larger animals moved further than smaller ones at one of these sites. At Enewetak, there were no significant associations between distance and size or grazing interval. Frequency of movement and median time spent off scars remained unaffected by size at all sites, and there was no evidence that grazing frequency affected the length of time that limpets spent off scars.     

Calanoid copepod grazing on phytoplankton: seasonal experiments on natural communities

Calanoid copepods are major components of most lacustrine ecosystems and their grazing activities may influence both phytoplankton biomass and species composition. To assess this we conducted four seasonal, in situ, grazing experiments in eutrophic Lake Rotomanuka, New Zealand. Ambient concentrations of late stage copepodites and adults of calanoid copepods (predominantly Calamoecia lucasi, but with small numbers of Boeckella delicata) were allowed to feed for nine days on natural phytoplankton assemblages suspended in the lake within 1160 litre polyethylene enclosures. The copepods reduced the total phytoplankton biomass of the dominant species in all experiments but were most effective in summer (the time of highest grazer biomass) followed by spring and autumn. In response to grazing pressure the density of individual algal species showed either no change or a decline. There were no taxa which increased in density in the presence of the copepods. The calanoid copepods suppressed the smallest phytoplankton species (especially those with GALD (Greatest Axial Linear Dimension) < µm) and there appeared to be no selection of algae on the basis of biovolume. Algal taxa which showed strong declines in abundance in the presence of the copepods include Cyclotella stelligera, Coelastrum spp., Trachelomonas spp., Cryptomonas spp., and Mallomonas akrokomos. Calanoid copepods are considered important grazers of phytoplankton biomass in this lake. The study supports the view that high phytoplankton:zooplankton biomass ratios and large average algal sizes characteristic of New Zealand lake plankton may, at least partly, be caused by year round grazing pressure on small algae shifting the competitive balance in favour of larger algal species.