The interplay of physical and biological factors in maintaining mid-shore and low-shore assemblages on rocky coasts in the north-west Mediterranean

This study examined the interactive effects of grazing by limpets and inclination of the substratum in maintaining differences between mid-shore and low-shore assemblages of algae in the northwest Mediterranean, at different scales of space and through time. Alternative models leading to different predictions about these effects were proposed and tested. Limpets were excluded by fences from areas of the substratum at mid levels on the shore. The response of algal assemblages to this manipulation was compared with control and enclosure plots at the same level, and with unmanipulated plots in the low shore where limpets are less abundant. The effects of limpets were examined at several replicated sites (0.1–4 km apart) for each slope of the substratum (nearly horizontal vs vertical), at different locations (hundreds of kilometres apart) and at different times. Individual taxa responded differently to limpet exclusion. The percentage cover of the coarsely branched and filamentous algae increased significantly in exclosure plots, in some loser reaching values found on the low shore. These patterns, however, varied greatly from shore to shore and significant effects were found both on horizontal and vertical substrata. Multivariate analyses indicated that grazing by limpets accounted for about 20% of the differences between mid-shore and low-shore assemblages. This effect was independent of substratum inclination and was consistent in space and time, suggesting that physical conditions were not as stressful for macroalgae on vertical substrata as initially supposed. Variable recruitment of algae is proposed as a possible explanation for the lack of consistency in the effects of limpets at the scale of the shore. The results of this study emphasize the need for multiple-scale analyses of the interactive effects of physical and biological factors to understand the organization of natural assemblages. Received: 22 June 1999 / Accepted: 15 November 1999     

An integrated study of the temporal and spatial variation in the supply of propagules, recruitment and assemblages of intertidal macroalgae on a wave-exposed rocky coast, Victoria, Australia

Recruitment is known to influence distributions and abundances of benthic marine organisms. It is therefore important to document patterns of variability in recruitment and how these relate to patterns in established assemblages. This study provides an integrated assessment of the temporal and spatial variation in supply and recruitment of propagules and established populations of several macroalgae. Propagules in water samples from two stages of the incoming tide, recruitment to artificial substrata and percentage cover of species established on the shore were recorded every 2 months from December 1994 to October 1995, in two zones of an intertidal, wave-exposed rocky shore. Variability in recruitment was measured at three spatial scales: 10s cm, 100s cm and 100s m. Availability and recruitment of most taxa were greatest between April and August, although many species had available propagules and recruited throughout the year. Temporal variation in the established assemblages was, however, more species-specific. Differences in established assemblages between zones were reflected in differences in availability and recruitment of propagules between zones. Recruitment could not be predicted directly from supply of propagules, but the two processes were linked. For most species, the greatest variation in recruitment occurred at the smallest spatial scale of 10s cm, although there was also considerable large-scale (between site) variation in recruitment of several species. Results indicate that while pre-and post-settlement mortality are likely to influence macroalgal distribution and abundance, the temporal and spatial variability in supply and recruitment of propagules can explain much of the patchiness in macroalgal assemblages.     

Relative effects of elevated grazing pressure and competition from a red algal turf on two post-settlement stages of Fucusevanescens C. Ag.

In Nova Scotia, Canada, the lower shore of wave-exposed and semi-sheltered portions of the coast is dominated by a dense turf of the red alga Chondrus crispus C. Ag. The mid shore is dominated by three species of rockweed belonging to the genus Fucus, which are also physiologically competent in the low zone, but may be inhibited by pre-emptive competitive exclusion by red algal turf and/or increased grazing pressure from abundant mesoherbivores. A grazer assay showed that herbivore pressure is >50% higher in the Chondrus zone than in the rockweed zone. Fucus can colonize the low shore only through microrecruits which develop from planktonic zygotes. A main objective of this study was to test the hypothesis that early post-settlement stages represent a critical phase for successful recruitment. We, therefore, tested the relative effects of competitive pressure from Chondrus and grazing on small (2 mm) and larger (6 cm) Fucus evanescens plants in a three-way factorial experiment. Red algal turf and grazers independently depressed F. evanescens growth, but only a combination of factors resulted in a significant weight loss in the rockweeds. The effect size of grazing was more than double that of competition from the turf. Small and larger individuals of F. evanescens were equally affected by grazing and competition. The results showed that colonization by rockweeds may only occur where there are gaps in the Chondrus canopy. We hypothesized that grazer effects may decrease with increasing gap size. However, a controlled experiment showed no differences in Fucus recruitment among the range of naturally occurring gap sizes. That is, larger gaps do not promote rockweed recruitment. In concordance with this observation, we found that grazer abundances were similar among gap sizes. Chondrus was shown to be highly resistant to intense physical and herbivore-induced disturbance. This combination ensures the persistence of a dense canopy which largely excludes rockweed recruitment from zygotes. Competitive dominance of Chondrus on the low shore is not obviously related to individual growth rates or canopy height. A positive correlation between growth rate and competitive ability may be expressed only in morphologically similar species which are likely to compete as germlings.     

The influence of canopy algae on vertical patterns of distribution of low-shore assemblages on rocky coasts in the northwest Mediterranean

Canopy-forming algae have been shown to play a fundamental role in the maintenance of understorey assemblages on rocky shores. In the Mediterranean, low-shore habitats are often monopolised by canopy algae of the genus Cystoseira and, in particular, by the species Cystoseira amentacea var. stricta. Alternatively, low-shore habitats are dominated by turf-forming algae and/or mussels. Previous studies showed that loss of Cystoseira, by natural or anthropogenic disturbances, resulted in the increase of turfing algae. Here, we propose that turf-forming algae may act as colonisers from nearby habitats, specifically from lower portions of the shore. The following hypotheses were tested to investigate this proposition: (1) in presence of Cystoseira, the assemblage living under its canopy will be distinct, in terms of composition and structure, from that found lower on the shore, (2) if the canopy of Cystoseira was removed, the differences between the two assemblages would decrease or disappear, (3) the effects of the canopy would be more important than other causes of variability at the spatial scale investigated (hundreds to thousands of metres), and (4) that effects of removal of the canopy would be consistent through time. These hypotheses were tested by means of a canopy removal experiment, involving several spatial and temporal repetitions of the manipulation.The assemblage underneath the canopy of Cystoseira was distinct from that found immediately lower on the shore, but when the canopy was removed there were no differences between the two. The effects of the canopy were consistent at the spatial and temporal scales investigated. Several species living under the canopy disappeared and were replaced by turf-forming species, resulting in a homogenisation of the two habitats and in a drastic loss of habitat diversity. This study shows that Cystoseira can be considered a habitat-forming species, responsible for the maintenance of two distinct low-shore assemblages. In this light, its importance should be taken into account by policies aiming to preserve biodiversity on rocky shores.     

Demography, competitive interactions and grazing effects of intertidal limpets in southern New Zealand

Population dynamics and the effects of intraspecific competition on limpet growth and maintenance of bare patches were investigated for the intertidal limpet Cellana ornata (Dillwyn) at a boulder-dominated site and on a rocky platform near Kaikoura (South Island), New Zealand. Distribution and abundance patterns of C. ornata were described in relation to other biota and tidal height. C. ornata occurred almost exclusively in patches devoid of macroalgae, particularly in the mid-tidal zone. Both adult and juvenile limpets were most abundant on the tops of boulders, where their numbers were positively correlated with barnacle cover, which averaged 77%. The size structure and growth patterns of C. ornata were different between populations. Mark-recapture studies showed that the slopes of annual growth increments regressed on initial sizes were similar at both sites but that the annual increments on the platform were about 6 mm greater than on boulders. Growth virtually ceased at 27 mm for limpets at the barnacle-dominated boulder site and at 40 mm at the platform site. Recruiting cohorts had 20% survival on boulders and 37% on the platform during their first year. The largest size classes at both sites had around 57% annual survival. To test the effects of varying limpet densities on the growth and mortality of limpets and the maintenance of bare patches, densities of C. ornata were experimentally increased at both sites. Beyond a density of 4 per 0.25 m², sizes and survival of limpets were reduced at both sites, but the effect was more pronounced at the boulder site. Limpets at the boulder site were more effective at maintaining bare space than those on the reef platform. Enclosing limpets in plots with and without barnacles showed that C. ornata and a co-occurring species (Cellana denticulata (Martyn)) grazed more effectively and had greater growth in cleared plots. Overall, there was considerable variation in the demographics of C. ornata between populations driven by site-specific characteristics.     

Spatial and temporal distribution patterns of the macrozoobenthos assemblage in the salt marshes of Tejo estuary (Portugal)

The present study focuses on the spatial and temporal distribution of the macroinvertebrate community of the salt marsh areas of the Tejo estuary, based on surveys conducted from autumn 1998 to summer 2000. Samples were collected quarterly in five different intertidal areas along an elevation gradient in: mudflats, creek mouths, creeks, pioneer salt marsh areas and middle marsh areas. A total of 36 benthic invertebrate taxa were identified. Insect larvae were the most well represented group, with 10 taxa identified. Oligochaetes and ostracods were the most numerically abundant taxa, whereas bivalves dominated in biomass. Benthic macroinvertebrate assemblages were dominated, both in number and biomass, by deposit feeders. Three distinct macroinvertebrate assemblages were distinguished along the elevation gradient, based on species presence, density and biomass: the unvegetated muddy areas with a macrobenthic assemblage composed mostly by infauna; the salt marsh pioneer areas of Spartina maritima in which several epibenthic taxa were found, as well as endobenthic taxa characteristic of muddy sediment; and the creek margins, with epifauna taxa such as insect larvae and crustaceans and a low abundance of benthic infauna. Total biomass in the unvegetated and Spartina areas was higher during spring and summer mainly due to the increase in biomass of Scrobicularia plana and Hydrobia ulvae. No decreases in the salt marsh macroinvertebrate biomass values were observed during the highest densities of their potential nektonic predators (summer). This fact might indicate that macroinvertebrates are not a limiting resource for the nektonic species, and that the natural biomass increment of these invertebrate species could be masking the predation/disturbance caused by the nektonic species.     

The effects of grazing on the pollen production of grasses

Samples from moss polsters taken over three consecutive years during a controlled grazing experiment in a nature reserve in the Netherlands were analysed for their pollen content, particularly grass pollen. Samples were also taken in the year following the experiment, with grazing pressure strongly diminished. This yielded data regarding the effect of grazing on the pollen production of grasses in open habitats, plantations, and unproductive and productive woods under different grazing pressures.     

The impact of grazing on plant communities,plant populations and soil conditions on salt marshes

Grazing an abandoned salt marsh causes retrogressive succession, since mid salt-marsh communities change into lower salt-marsh communities. Grazing and mowing are compared in detail. Both management practices enhance species diversity in an abandoned salt marsh. This can be attributed to the removal of litter. The finding that lower salt-marsh species appear more with grazing than with mowing or abandoning is not related to a higher soil salinity as compared to mowing or abandoning, but probably to locally baring of the soil by grazing animals. Only species of pioneer or unstable environments seem to have a persistent seed bank, for other species seed dispersal seems to be a limiting factor for their establishment.Nomenclature follows Heukels & van Ooststroom (1977) for species; Westhoff & den Held (1969) for syntaxa.Mrs R. Rusthoven analyzed the soil samples, Mr E. Leeuwinga drawed the figures, and Mrs J. O'Brien corrected the English text.     

The effects of sheep grazing on seedling establishment and survival in grassland

This paper examines the effects of sheep and insect grazing on the generation of natural gaps in turf and the establishment of seedlings therein. The site is on an old field on limestone. There were five sheep grazing treatments: ungrazed, and grazed briefly in spring, briefly in autumn, in autumn or in summer and autumn. The first three of these treatments also contained plots in which insecticide was applied regularly during the growing season. Pins of increasing size (1.6 mm–12.8 mm diameter) were lowered at grid intersections in permanent quadrats to record gap type and the occurrence of seedlings on seven dates over 15 months.Heavily-grazed plots had more bare ground in spring and more herb seedlings established in them. Seedlings of annual species of Bromus were able to establish in gaps (bare ground and litter) of smaller size than was required by herb seedlings (which had much smaller seeds). The use of an insecticide increased seedling establishment in October in ungrazed and in spring-grazed paddocks but decreased it in autumn-grazed paddocks. This may indicate the greater importance of insect herbivory in making gaps in the matted sward of autumn-grazed plots.Seedlings which established in October 1986 and March 1987 were more likely to die than those establishing earlier in 1986. This may be because of winter-kill and especially vigorous spring vegetation growth respectively. Seedling mortality was not affected by gap size or grazing treatment (unless such effects operated before we detected the seedlings).     

The effects of fire frequency and grazing on tallgrass prairie productivity and plant composition are mediated through bud bank demography

Periodic fire, grazing, and a variable climate are considered the most important drivers of tallgrass prairie ecosystems, having large impacts on the component species and on ecosystem structure and function. We used long-term experiments at Konza Prairie Biological Station to explore the underlying demographic mechanisms responsible for tallgrass prairie responses to two key ecological drivers: fire and grazing. Our data indicate that belowground bud banks (populations of meristems associated with rhizomes or other perennating organs) mediate tallgrass prairie plant response. Fire and grazing altered rates of belowground bud natality, tiller emergence from the bud bank, and both short-term (fire cycle) and long-term (>15 year) changes in bud bank density. Annual burning increased grass bud banks by 25% and decreased forb bud banks by 125% compared to burning every 4 years. Grazing increased the rate of emergence from the grass bud bank resulting in increased grass stem densities while decreasing grass bud banks compared to ungrazed prairie. By contrast, grazing increased both bud and stem density of forbs in annually burned prairie but grazing had no effect on forb bud or stem density in the 4-year burn frequency treatment. Lastly, the size of the reserve grass bud bank is an excellent predictor of long-term ANPP in tallgrass prairie and also of short-term interannual variation in ANPP associated with fire cycles, supporting our hypothesis that ANPP is strongly regulated by belowground demographic processes. Meristem limitation due to management practices such as different fire frequencies or grazing regimes may constrain tallgrass prairie responses to interannual changes in resource availability. An important consequence is that grasslands with a large bud bank may be the most responsive to future climatic change or other global change phenomena such as nutrient enrichment, and may be most resistant to exotic species invasions.     

The distribution and conservation of threatened Sphaeriidae on British grazing marshland

Freshwater sphaeriid bivalves are poorly known ecologically, particularly uncommon species such as Pisidium pseudosphaerium (UK RDB (Red Data Book) staus = ‘rare’). In the UK, this species occupies grazing marsh where conservation opportunities might be shared with other threatened molluscs. We surveyed sphaeriids including P. pseudosphaerium and snails in 106 drainage ditches in SE England in 1999. P. pseudosphaerium occupied over half the ditches surveyed, at slightly elevated BOD (6 ± 7 mg l⁻¹ sd) but reduced calcium (64 ± 31 mg l⁻¹) and nitrate (0.5 ± 1.2 mg l⁻¹). As part of a sphaeriid assemblage comprising Sphaerium corneum, Musculium lacustre and Pisidium obtusale (=Assemblage 1), P. pseudosphaerium occurred in ditches with floating vegetation and abundant snails of conservation importance, co-occurring significantly with either Valvata macrostoma (RDB ‘vulnerable’) or Anisus vorticulus (RDB: ‘vulnerable’). A more diverse sphaeriid assemblage (=Assemblage 2) included species common in wider, deeper ditches with open water. We suggest that traditional ditch management can support both rare and representative snails and bivalves on grazing marsh. Quasi-traditional and rotational ditch clearance will favour common sphaeriids and pioneer snails during early succession; P. pseudosphaerium, V. macrostoma, A. vorticulus during mid-succession; and the snail Segmentina nitida (RDB: ‘Endangered’) in late successional ditches. In common with threatened snails, P. pseudosphaerium will also benefit from reduced eutrophication. We recommend that P. pseudosphaerium retain RDB status to ensure protection and to emphasise the need for improved ecological information.     

The effects of simulated spring goose grazing on the growth rate and protein content of Phleum pratense leaves

The effects of simulated goose grazing on Phleum pratense plants were tested in an Iceland hayfield during the spring goose staging period (19 April–11 May 1997). Plants in an area exclosed from the influence of grazing and the nutrient effects of goose faeces were subject to the removal of the youngest lamina once, three and four times during this period. Clipping three and four times resulted in 25–41% increases in cumulative elongation of youngest laminae compared with unclipped plants. Total cumulative lamina growth of entire plants showed no significant difference between unclipped plants and those clipped three and four times, hence no overcompensation occurred. Sequential clipping elevated the protein content of the youngest laminae from 20% to 27–33%, whereas there was no change amongst shoots clipped only once. Because geese only consume the youngest lamina of each Phleum plant, measurements from this experiment showed that regular physical removal of growing biomass doubled the biomass of preferred tissue available to geese and increased the potential protein intake 3.5 times at experimental clipping frequencies similar to levels of sequential harvesting observed amongst staging geese compared to less frequent harvesting. These increases were achieved without any fertilising effects of goose faeces implicated in such effects in previous studies. Received: 26 January 1998 / Accepted: 20 March 1998     

Conservation implications of grazing practices on the plant and dipteran communities of a turlough in Co. Mayo,Ireland

Turloughs, which are classified as priority habitats under the European Habitats Directive, are seasonally flooded depressions found almost exclusively in Ireland. In 2001, three adjacent fields with different stocking densities were selected and plant/dipteran communities within the same vegetation zone of each field (site) were investigated using quadrats and sweep netting, respectively. There was a significant positive relationship between Diptera morphospecies richness/Diptera abundance and mean vegetation height (P < 0.001). However, no significant relationship between Diptera morphospecies richness and plant species richness was found. Median Diptera morphospecies richness per sweep was lower at the site with the highest stocking density (17) than at the other two sites (22 and 31, respectively). Total species richness of Sciomyzidae was greater at the least grazed site (7) than at the more heavily grazed sites (2 and 1, respectively). The results suggest that an evaluation of turlough management practices based on plant communities alone is not sufficient and that at least some areas within the turlough basin remain ungrazed on a rotational basis to ensure maximum diversity of Diptera.     

The effects of selective fish predation on the horizontal distribution of pelagic Cladocera in a southern French reservoir

A four-year study showed a clear seasonal succession of species within the cladoceran community of the large, oligotrophic Sainte-Croix reservoir (S.E. France). Diaphanosoma brachyurum and Ceriodaphnia pulchella were strictly limited to the warm stratified period (July to October), whereas Bosmina longirostris and Bosmina coregoni were dominant during spring and autumn. Daphnia longispina was the only species to occur throughout the year with higher densities in spring.In spring and late autumn, the discharge of the inflowing river Verdon was high and the abundance of all species showed a gradient over the whole lake with lower densities close to the inflow. During the stratified period, water inflow was very low and species showed different patterns. Densities of the small form Ceriodaphnia pulchella were similar all along the long axis of the lake, whereas Daphnia densities were significantly higher near the dam. The distribution pattern of Diaphanosoma, an intermediate-sized species, showed similar trends to that of Daphnia. The only planktivorous fish in the pelagic zone, the bleak (Alburnus alburnus), fed mostly on large-bodied species (> 1.0 mm) and was more abundant close to the inflow current. A comparison between the length frequency distributions of cladocera upstream and downstream provided a clear demonstration of the effects of size-selective predation on prey populations. Finally, the interactions between spatial heterogeneity and long-term development of the zooplankton community and the indirect effects of predation are discussed.     

The effects of chrysomelid beetle grazing and plant competition on the growth of Rumex obtusifolius

Summary The interaction between grazing by Gastrophysa viridula and interspecific plant competition was investigated for Rumex obtusifolius growing in the field. During an eight-month growing season non-competing R. obtusifolius grew larger than competing plants although herbivore loads (beetle productivity per unit leaf area) were similar for competing and non-competing plants. However, grazing significantly reduced the growth of R. obtusifolius only in competing plants thus demonstrating a synergism between plant competition and invertebrate herbivory. The relevance of these findings for the distribution of plant species is discussed.     

Water salinity effects on performance and rumen parameters of lactating grazing Holstein cows

Eighteen multiparous lactating grazing Holstein cows, 9 ruminally cannulated, average 136.1 ± 14.6 days in milk, were randomly assigned to three treatments consisting of water containing different levels of total dissolved solids (TDS; mg/l): Treatment 1 = 1,000; Treatment 2 = 5,000 and Treatment 3 = 10,000, at the Experimental Dairy Unit at Rafaela Experimental Station (31°11′S latitude) during summer 2005. Animals were arranged in a randomized complete block design with three 28-day experimental periods, with 3 weeks for water adaptation and 1 week for measurements. Feed and water intake, milk production and composition, body weight and condition score and rumen parameters were evaluated. No treatment effects were observed in any of the variables evaluated, with the exception of water intake, which was higher for animals receiving 10,000 mg/l TDS in the drinking water (189 l/day vs. 106 and 122 l/day for cows receiving water with 1,000 and 5,000 mg/l TDS, respectively). Water intake was significantly higher for animals in treatment 10,000 (P < 0.05). It was concluded that the rumen presents a surprising buffer capacity and that consideration of TDS alone is insufficient to characterize drinking water quality.     

The effects of sea urchin grazing and drift algal blooms on a subtropical seagrass bed community

Subtropical seagrass beds can be subject to relatively high levels of direct herbivory and large blooms of drift algae, both of which can have important effects on the floral and faunal components of the community. Caging experiments were used to investigate these factors in a Thalassia testudinum bed in Biscayne Bay, Florida. Abundance of sea urchins, Lytechinus variegatus, and drift algae was manipulated within the cages. Naturally occurring levels of urchin grazing do not appear to affect the T. testudinum population. With experimentally increased urchin densities in the winter, seagrass shoot density and aboveground biomass decreased significantly. Similar effects were not detected in the summer, indicating that the impact of grazing on T. testudinum is lessened during this time of year. Shoot density was more vulnerable to grazing than aboveground biomass. This may be a result of grazing-induced increases in seagrass productivity, in which the remaining shoots produce more or longer leaves. In the winter, drift algal blooms form large mats that cover the seagrass canopy. Under the normal grazing regime these algal blooms do not have significant negative effects on the seagrass. With increased grazing pressure, however, there is a synergistic effect of grazing and drift algae on seagrass shoot density. At intermediate urchin density (10 per m(-2)), cages without algae did not undergo significant decreases in shoot density, while those with algae did. At the high density of urchins, the number of seagrass shoots in cages both with and without algae decreased, but the effect was more pronounced for cages with algae. Invertebrate abundance at the field site was low relative to other seagrass beds. There were no discernible effects, either positive or negative, of urchin and algae manipulations on the sampled invertebrate community.     

Seasonal changes in the abundance and distribution of submerged aquatic vegetation in a highly managed coastal lagoon

The inflow of fresh water into coastal lagoons is a key factor influencing the structure and function of these ecosystems. Biscayne Bay, a coastal lagoon adjacent to the city of Miami, is located downstream of the Everglades ecosystem where the extensive water management system now in place has modified the historical hydrology, replacing groundwater and overland flows with pulsed releases from canals. In areas where canals discharge directly into littoral habitats, an environment with low-mean salinity and high-salinity variability is created. In this study, we characterize the salinity patterns of nearshore habitats (<500 m from shore) and document patterns of seasonal abundance and distribution of submerged aquatic vegetation (SAV) to evaluate the impacts of water management practices. Seagrasses were the principal component of the SAV community during the 2005 dry season (mean cover = 25.5%), while macroalgae dominated during the wet season (mean cover = 33.4%). The distribution and abundance of SAV were directly related to the tolerance of each taxon to salinity patterns. Seagrass species with high tolerance to low and variable salinity such as Halodule wrightii and Ruppia maritima were found only in canal-influenced areas and increased in abundance and spatial distribution in the wet season when freshwater inflow is highest. The dominance of rhizophytic macroalgae during the wet season was determined by the appearance and high abundance of Chara, a taxon commonly associated with freshwater environments. Thalassia testudinum, the most abundant seagrass species, was found throughout the study region, but decreased in abundance in the canal-influenced areas during the wet season when lower, more variable salinity resulted in lowered productivity. The data presented here showed a significant relationship between salinity patterns and the seasonal abundance and distribution of SAV. These findings support the use of SAV as appropriate indicators of changes in water quality resulting from future restoration projects associated with the Everglades Restoration Plan, which will once again modify the delivery of fresh water into littoral habitats with unknown ecological consequences. Handling editor: S. M. Thomaz     

The independent and interactive effects of snail grazing and nutrient enrichment on structuring periphyton communities

We investigated the independent and interactive effects of nutrient enrichment and snail grazing on structuring periphyton communities in a northern temperate lake. Nutrient releasing substrates and grazer enclosures were used to simultaneously manipulate nutrient availability and herbivory. Periphyton was allowed 18 days to accrue before grazers (Elimia livescens = Goniobasis livescens) were introduced.Addition of nitrogen and phosphorus caused a significant increase in biovolume (p < 0.001), whereas grazing had no significant effect on biovolume but resulted in a shift in species composition. Four taxa were largely responsible for the increase in biovolume on the nutrient enriched substrates: Oedogonium sp, Stigeoclonium tenue, Navicula radiosa var. radiosa and Navicula radiosa var. tenella. By the 28th day, nutrient enrichment caused a shift from a community dominated by diatoms (Bacillariophyceae) to a community dominated by green algae (Chlorophyceae). Blue green algae (Myxophyceae) maintained an equal proportion in high and low-nutrient regimes.Grazing had a more pronounced effect on altering community composition on the nutrient enriched substrates than on the unenriched substrates. Grazing caused a decrease in diversity and an increase in dominance by green algae on the nutrient enriched substrates. The relative biovolume of green algae increased from 64% to 93% on grazed substrates, due to the significant increase in relative abundance of Stigeoclonium tenue. This taxon has both prostate basal cells and erect filamentous cells. The ratio of basal: filamentous cells increased from 4.7 to 5.2 with grazing, suggesting that the heretotrichous growth form of Stigeoclonium tenue is adapted to grazing by virtue of the basal cells which are able to adhere to the substratum and resist being grazed.     

The influence of copepod and krill grazing on the species composition of phytoplankton communities from the Scotia Weddell sea

The influence of copepods (mainly Oithona sim-ilis) and krill (Euphausia superba) grazing on the species composition of plankton communities in ship board con tainers was investigated during the spring and post spring period in the Scotia Weddell Sea in the Antarctic ocean. Numbers of grazers were experimentally manipulated in containers with natural phytoplankton assemblages. With ratural levels of copepods but no krill a high (700–950 g C·l¹, ca 30 g chl a·.l¹) phytoplankton biomass developed. In these cultures large diatoms, e.g. Corethron criophilum and chains of Thalassiosira sp., made up 80% of total phytoplankton cell carbon at the end of the experiment. In cultures with elevated numbers of copepods (5X or 10X the natural level) phytoplankton biomass was somewhat reduced (ca 23 g chl a · l¹) compared to cultures with natural copepod abundance, but still high. Phytoplankton species composition was on the other hand greatly influenced. Instead of large diatoms these cultures were dominated by Phaeocystis pouchetii (70%) together with small Nitszchia sp. and Chaetoceros neogracile (20%). In containers with krill (both juveniles and adults), but without elevated numbers of copepods, phytoplankton biomass rapidly approached zero. With 10X the in situ level of copepods, krill first preyed on these before Corethron criophilum and Thalassiosira sp. were grazed. When krill were removed a plankton community dominated by flagellates (60–90%), e.g. Pyramimonas sp. and a Cryptophycean species, grazed by an unidentified droplet-shaped heterothrophic flagellate, developed. These flagellates were the same as those which dominated the plankton community in the Weddell Sea after the spring bloom. A similar succession was observed in situ when a krill swarm grazed down a phytoplankton bloom in a few hours. Our experiments show that copepods cannot control phytoplankton biomass in shipboard cultures even at artificially elevated numbers. Krill at concentrations similar to those in natural swarms have a great impact on both phytoplankton biomass and species composition in shipboard cultures. Both copepods and krill may have an impact on phytoplankton species composition and biomass in situ since the rates of phytoplankton cell division were probably artificially increased in shipboard cultures compared to natural conditions, where lower growth rates make phytoplankton more vulnerable to grazing. A similarity between phytoplankton successions in containers and in situ, especially with respect to krill grazing, supports the conclusion that grazing may structure phytoplankton communities in the Scotia-Weddell Sea.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation