Variation in structure and diversity in Mediterranean grasslands related to trophic status and grazing intensity

Abstract. The spatial organization of Mediterranean grassland in Spain is described, based on samples from 71 sites covering the existing variation in slope exposition and inclination.The whole set can be regarded as representing a trophic gradient, along which gradual variation in soil, species composition, biomass, and coverage were quantified. Corresponding to other studies from varying habitats, maximal species richness, diversity, and heterogeneity were observed on moderately infertile sites. Maximum species richness, over 60 species, occurred on sites with biomass values from 150–350 g / m². Species richness values are much higher and biomass values are much lower than those found in temperate grasslands. The decrease in diversity towards the mosteutrophic communities is stronger than expected, but can be easily explained by the high grazing pressure here. The variation in diversity observed runs parallel with that in heterogeneity. Zones with a high species richness also have a high heterogeneity, meaning a low amount of dominance. Grazing is understood as abiotic form of disturbance. Differences in grazing pressure may modify the relation between richness and fertility. While the greatest grazing pressure coincides with the most eutrophic communities, decreasing progressively towards the oligotrophic ones, the trend predicted by the resource availability theory is maintained; species diversity will be maximal at intermediate levels of disturbance. Absence of grazing in the eutrophic communities would lead to an investment in the soil of the unconsumed organic matter.     

Diversity and guild structure of insect assemblages under grazing and exclusion regimes in a montane grassland from Central Argentina

The effects of grazing disturbance on insect communities were examined at a montane grassland in Central Argentina, by comparing two grazed sites differing in cattle load (heavy and continuous or moderate and discontinuous) and two cattle exclusions differing in age (7 and 19 years). Two aspects of insect diversity (taxonomic and guild structure) and two levels of taxonomic resolution (family and species) were considered. Four monthly samples were taken with a suction sampler in two 1 m² areas at each site. Collected specimens were counted, identified to family (all insects) or species (Coleoptera) level, and allocated to trophic guilds. Abundance, richness, diversity and biomass of the insect assemblages had minimum values in the most intensely grazed habitat, which also differed from the other sites in terms of insect families and Coleoptera species composition. It also showed a distinct guild structure, with fewer secondary consumers, and chewers replacing suckers as the most abundant herbivore group. According to these observations, intense grazing in montane grasslands in Central Argentina could result in taxonomic and guild changes in the associated insect communities, but such effects would not be noticeable with less intensive use. Moreover, using family taxonomic level could be as or even more appropriate than species level in order to characterize insect communities in the studied habitats under varying disturbance regimes.     

Effects of grazing by Diadema antillarum Philippi (Echinodermata: Echinoidea) on algal diversity and community structure

In Discovery Bay, Jamaica W.I., densities of Diadema antillarum Philippi were experimentally manipulated over a range of 0–64 individuals · m^?2 with the aid of enclosures in shallow water. The effects of grazing by this echinoid on the algal community structure of a coral reef were compared under conditions of primary and post-primary succession.Algal biomass decreased as Diadema density was increased, particularly under primary succession conditions. A similar response was noted under post-primary conditions, but here biomass was also dependent on species composition and abundance of algae prior to experimental alterations of Diadema density. Peyssonnelia sp. was found to be particularly resistant to echinoid grazing, surviving even the most intense grazing. Within 11 months, algal diversity decreased significantly in an exponential fashion as Diadema density increased. This occurred under conditions of both primary and post-primary succession. Diversity, irrespective of index, was not maximized at intermediate grazing pressures. No single species of alga dominated the benthic community at even the lowest Diadema densities.Species composition of algae in the benthic community prior to changes in grazing pressure and lack of successful recruitment of a potential dominant were found to have a major influence on the response of the algal community to the treatments. Each of these factors can influence both the relationship between algal diversity and grazing pressure and the rate at which the community approaches an initial plateau of diversity in its earliest stages of succession.     

Effects of grazing,competition, disturbance and fire on species composition and diversity in grassland communities

Abstract. The effects of grazing, physical disturbance, interspecific competition, and fire on populations of individual species and on the structure and diversity of grassland communities were investigated in short-, mid-, and tall-grasslands in the Serengeti National Park, Tanzania. The treatments included deep and shallow artificial disturbances, early dry-season burning, and removal of dominant grass species from 2m² plots located inside and outside of large-mammal-proof exclosures. Species-cover data were collected for five years and analyzed by analysis of variance and diversity indices. In all three communities, protection from grazing and physical disturbance had greater impacts on species cover and diversity than removal of dominant species or fire. More species were significantly affected (positively and negatively) by disturbance and protection from grazing (50 - 100%) than by species removals and fire (< 41%). In most cases, the cover of tall, perennial species increased following protection from grazing while the cover of annual and short, perennial species decreased. Over the five years of the study, vegetatively reproducing species benefited more than sexually reproducing species from protection from grazing, and tall rhizomatous species benefited more than tall stoloniferous species. Disturbance caused annual and short perennial species to increase and tall perennials to decrease in cover. Following species removals, tall species were either unaffected or they increased in cover while some short species increased and others decreased in cover. Species diversity in two of the three communities decreased when the communities were protected from grazing but increased when disturbed, when dominant species were removed, and when burned.     

Plant—herbivore interactions in streams near Mount St Helens

1. In four separate field experiments near Mount St Helens (Washington, U.S.A.) during 1986, the grazing effects of two large benthic herbivores, tadpoles of the tailed frog Ascaphus truei and larvae of the caddisfly Dicosmoecus gilvipes, were investigated using streamside channels and in-stream manipulations. In the experimental channels, abundances of periphyton and small benthic invertebrates declined significantly with increasing density of these larger herbivores. 2. In eleven small, high-gradient streams affected to varying degrees by the May 1980 eruption, in-stream platforms were used to reduce grazing by A, truei tadpoles on tile substrates. Single platforms erected in each tributary and compared to grazed controls revealed only minor grazing effects, and no significant differences among streams varying in disturbance intensity (and, consequently, tadpole density). However, results probably were confounded by high variability among streams in factors other than tadpole abundance. 3. Grazing effects were further examined in two unshaded streams with different tadpole densities, using five platforms per stream. In the stream with five tadpoles m^?2, grazing reduced periphyton biomass by 98% and chlorophyll a by 82%. In the stream lacking tadpoles, no significant grazing effects were revealed. Low algal abundance on both platforms and controls, and high invertebrate density in that stream (c. 30000m^?2) suggests that grazing by small, vagile invertebrates was approximately equivalent to that of tadpoles. 4. The influence of large benthic herbivores on algal and invertebrate communities in streams of Mount St Helens can be important, but reponses vary spatially in relation to stream disturbance history, local environmental factors, and herbivore distributional patterns and abundance.     

Seasonal fluctuations in the diversity and compositional stability of phytoplankton communities in small lakes in upper Bavaria

A study of diversity and compositional stability of phytoplankton communities during one vegetation period was carried out in small lakes in upper Bavaria. Shannon-Weaver diversity index was calculated on the base of number of individuals and on the base of biomass. On average, the diversity (annual mean) was highest in mesotrophic lakes. A comparison of three morphologically different (esp. exposure to wind, surface area and mean depth) lakes (Pelhamer See, Thalersee and Kautsee) sought to find out how the phytoplankton community structure reacts to events of intermediate disturbance, in terms of diversity- or biomass changes. Principal Component Analysis was used to measure the persistence of the phytoplankton association. The examples given in this paper led to these conclusions: High diversity or increase in diversity occur in compositionally instable communities, in high wind-stress events, with small algae and with high grazing. Low diversity or decrease in diversity occur in compositionally stable periods, when conditions select few species, as large forms dominate and as grazing by zooplankton takes effect. Gradual seasonal changes are observed in structurally stable lakes. In lakes exposed to frequent disturbances, seasonal changes may be dominated by intermediate responses.     

中国东北样带草地群落放牧干扰植物多样的变化

放牧干扰是草地群落植物多样性变化的主要影响因素之一。中国东北样带９个草地群落放牧干扰植物多样性性变化的研究结果表明：中牧或重牧阶段Ｓhannon指数达最大值。形成中牧（重牧）＞重牧（中牧）＞轻牧＞过牧的规律。群落物种丰富度、均匀度与多样性的相关分析表明,均匀度变化对多样性变化具有更大的贡献率。而丰富度呈下降趋势,即轻牧（中牧）＞中牧（轻牧）＞重牧＞过牧。生活型功能群多样性也表现出明显的变化。中国东北样带草地群落植物多样性的分布格局是：草甸草原＞典型草原＞典型草原＞荒漠草原＞碱化草甸,并且群落物种丰富度对多样性有更大贡献率。     

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

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

The species richness–biomass relationship in herbaceous plant communities: what difference does the incorporation of root biomass data make?

So far, in all studies on the much-discussed hump-backed relationship between plant community productivity and species richness, productivity has been assessed through plant shoot biomass, i.e. it has been ignored that frequently most of the biomass is produced below ground. We revisited the 27 grassland and forest field-layer communities, studied earlier by Zobel and Liira, to sample root biomass, plant total biomass and root/shoot allocation, and learn how the incorporation of below-ground biomass data would affect the shape of the hump-backed relationship. In order to avoid scaling artefacts we estimated richness as the average count of species per 500 plant ramets (absolute richness). We also included relative richness measures. Relative richness was defined as richness per 500 ramets/size of the actual species pool (the set of species present in the community), relative pool size was defined as size of the actual species pool/size of the regional species pool (the set of species available in the region and capable of growing in the given community).
The biomass-absolute richness relationship was humped, irrespective of the biomass measure used, the hump being most obvious when plant total biomass was used as the independent variable. Evidently, the unimodal richness–productivity curve is not a sampling artefact, as suspected by Oksanen. However, relative richness was not related to community biomass (above-ground, below-ground or total). The hump-backed curve is shaped by the sizes of actual species pools in communities, implying that processes which are responsible for small-scale diversity pattern mainly operate on the community level.
Neither absolute nor relative richness were significantly related to root/shoot allocation. The presumably stronger (asymmetric) shoot competition at greater allocation to shoots appears not to suppress small-scale richness. However, there is a significant relationship between relative pool size and root/shoot allocation. Relatively more species from regional species pools are able to enter and persist in communities with more biomass allocated into roots.     

The stream as a habitat templet for bryophytes: species' distributions along gradients in disturbance and substratum heterogeneity

1. The structure of bryophyte communities in streams in relation to habitat characteristics, especially disturbance, is described. Disturbance in rivers is quantified as movement of the stream bed, whereas in small streams water level fluctuation is used as an indicator of disturbance frequency. 2. Canonical correspondence analysis differentiated frequently disturbed sites from more stable ones. The existence of a disturbance gradient was confirmed in a subset of the study sites using long-term records of discharge variation. A parallel change was detected in the species composition of bryophyte communities with low-statured, potentially fast colonizers dominating the disturbed end and large perennial species the stable end of the gradient. 3. A consistent pattern of zonation of bryophyte species was found along the gradient from continually submersed to persistently dry conditions in small streams and lake outlets. An abrupt increase in species richness occurred at or just above the water line, where facultatively aquatic species tolerant of both conditions formed the bulk of the community. 4. The relationship between species richness and standing crop in stream bryophyte communities was consistent with the hump-backed model of Grime (1979), especially at the within-habitat scale. Quadrats of low and very high standing crop were characterized by low species richness, while peak richness was observed at intermediate standing crops. 5. A few perennial species (e.g. Fontinalis spp. and Rhynchostegium riparioides) capable of monopolizing space dominated the most stable habitats. Species composition in low biomass sites was more variable, yet only one basic growth-form (small-statured species with high allocation to spore production) seemed possible in these highly disturbed streams. In habitats of intermediate biomass, small-scale disturbances apparently allow the formation of a more varied bryophyte community. 6. A habitat templet for stream bryophyte life strategies and community structure is presented. Disturbance is proposed to be the factor filtering out traits unsuitable for a given environment. The potential of stream bryophytes for testing and developing general ecological theory is emphasized.     

Gastropod grazers and nutrients,but not light,interact in determining periphytic algal diversity

The potential interactions of grazing, nutrients and light in influencing autotroph species diversity have not previously been considered. Earlier studies have shown that grazing and nutrients interact in determining autotroph species diversity, since grazing decreases species diversity when nutrients (i.e. N or P) limit autotroph growth, but increases it when nutrients are replete. We hypothesized that increased light intensities would intensify the interactions between grazing and nutrients on algal species diversity, resulting in even stronger reductions in algal species diversity through grazing under nutrient–poor conditions, and to even stronger increases of algal species diversity through grazing under nutrient-rich conditions. We studied the effects of grazing (absent, present), nutrients (ambient, N + P enriched) and light (low light, high light) on benthic algal diversity and periphyton C:nutrient ratios (which can indicate algal nutrient limitation) in a factorial laboratory experiment, using the gastropod grazer Viviparus viviparus. Grazing decreased algal biomass and algal diversity, but increased C:P and N:P ratios of periphyton. Grazing also affected periphyton species composition, by decreasing the proportion of Spirogyra sp. and increasing the proportion of species in the Chaetophorales. Grazing effects on diversity as well as on periphyton N:P ratios were weakened when nutrients were added (interaction between grazing and nutrients). Chlorophyll a (Chl a) per area increased with nutrient addition and decreased with high light intensities. Light did not increase the strength of the interaction between grazing and nutrients on periphytic algal diversity. This study shows that nutrient addition substantially reduced the negative effects of grazing on periphytic algal diversity, whereas light did not interact with grazing or nutrient enrichment in determining periphytic algal diversity.     

中国东北样带草地群落放牧干扰植物多样性的变化

放牧干扰是草地群落植物多样性变化的主要影响因素之一。中国东北样带9个草地群落放牧干扰植物多样性变化的研究结果表明：中牧或重牧阶段Shannon指数达最大值,形成中牧（重牧）>重牧（中牧）>轻牧>过牧的规律。群落物种丰富度、均匀度与多样性的相关分析表明,均匀度变化对多样性变化具有更大的贡献率,而丰富度呈下降趋势,即轻牧（中牧）>中牧（轻牧）>重牧>过牧。生活型功能群多样性也表现出明显的变化。中国东北样带草地群落植物多样性的分布格局是：草甸草原>典型草原>典型草原 >荒漠草原>碱化草原,并且群落物种丰富度对多样性有更大贡献率。     

Influence of grazing and soil conditions on secondary savanna vegetation in India

Abstract. Savanna vegetation and pertinent soil features were studied on 43 sites in a dry tropical forest region of India. Grazing intensity ranged from 0.68 to 0.98. Soil moisture was positively related to the proportion of fine soil particles (< 0.1 mm), and the latter decreased while the proportion of coarse particles (2.0-0.5 mm) increased with increasing grazing intensity. Canopy biomass ranged from 28 to 104 g/m² in grazed communities and from 230 to 337 g /m² in ungrazed communities and was positively related to vegetation cover which ranged between 30–72 % in grazed and 68 - 91 % in ungrazed communities. Vegetation cover was negatively related to grazing intensity. Species richness and diversity were highest at low grazing intensity. Using community coefficients and Detrended Correspondence Analysis, the grazed stands were clustered into six and the ungrazed ones into three communities. The grazed communities were recognised as degradation stages and the ungrazed ones as recovery stages. Only five grass species, in various combinations were able to dominate in one of the different stages. Evidently the harsh climatic conditions (high temperatures, high variability in rainfall and a long dry period) in the region permit only a few species already adapted to these conditions to participate in the succession.     

Development of filamentous green algae in the benthic algal community in a littoral sand-beach zone of Lake Biwa

Temporal changes of biomass and dominant species in benthic algal communities were investigated in a littoral sand-beach zone in the north basin of Lake Biwa from December 1999 to September 2000. Chlorophyll-a amounts of benthic algal communities per unit area of the sandy sediments rapidly increased from late April to June. Increases in biomass of the benthic algal communities are considered to result from the propagation of filamentous green algae Oedogonium sp. and Spirogyra sp. The cell numbers of filamentous green algae and chlorophyll-a amounts of benthic algal communities at depths of 30 and 50cm at a station protected by a breakwater in May were significantly higher than those of a station exposed directly to wave activity. Thus, the biomass accumulation of the benthic algal communities seems to be regulated strongly by wave disturbance. The development of filamentous green algae may contribute to the increase in biomass of the benthic algal community and to the changes in seasonal patterns of biomass in the sand-beach zone of Lake Biwa. We consider that the development of the filamentous green algal community in the littoral zone of Lake Biwa is the result of eutrophication.     

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity–ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m⁻² and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m⁻². One year after seed addition, local plant species richness had increased on average by six species m⁻² (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m⁻² (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump‐shaped model appears to be the limiting outline of the natural diversity–productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.     

Variation in species composition and diversity of wetland communities under different disturbance intensity in the Sanjiang plain

In this study, ditch density was chosen as an indicator of disturbance intensity, and four sites with different ditch density were chosen to compare species composition and diversity among them. Four transects and sixty-seven quadrats were sampled along the ditch density gradient. The results showed that there existed a significant negative relationship between disturbance intensity and species parameters, while species diversity did not differ greatly among sites, but species composition varied considerably. With increasing disturbance intensity, the wetlands tended to be reduced in plant species diversity, and an increasing loss of indigenous wetland species was paralleled with an increasing invasion of upland species. The results also show that the rate of species turnover within communities differed among communities under different disturbance intensities. The communities with intermediate ditch density (0.6-1.2 km/km²) had higher rates than the other communities.     

三江平原不同强度干扰下湿地植物群落的物种组成和多样性变化

以三江平原湿地-农田景观中沟渠的密度作为反映干扰强度的指标,研究了不同沟渠密度下湿地斑块植物群落的物种组成和多样性变化.结果表明:(1)在不同的干扰强度下,湿地植物群落的物种组成差异很大, 随干扰强度增大,原有湿生物种减少,而中湿生、中生物种显著增加;(2)湿地植物群落的多样性随干扰强度的增大,整体上呈降低趋势,群落多样性在中等沟渠密度下(0.6～1.2km/km2)达到最大,但不同沟渠密度下群落间多样性差异基本不显著;(3)位点间的干扰强度差异越大,Whittaker指数值越高;(4)不同的干扰强度下,各湿地斑块内部的物种周转速率同样在沟渠密度中等(0.6～1.2km/km2)时最高.     

Grazing effects in macroalgal communities depend on timing of patch colonization

Rocky macroalgal assemblages are typically composed of patches differing in age and species composition and grazing is generally a very important modifier of such assemblages. We hypothesized that patch colonization time determines its algal community and that grazing effects depend on the colonization time and vary with depth. We created patches by placing empty substrates at two sublittoral depths over five consecutive months, manipulated grazer entry and determined the algal species composition in each patch in the next growing season. Distinct algal colonization periods resulted in different algal assemblages. Although algal communities in our study area consist mainly of opportunistic species, thus being highly dynamic, the resulting macroalgal assemblages differed in species richness, diversity, composition, and total biomass even a year after first colonization. Substrates close to the water-surface supported a higher species richness and diversity than those in the deeper littoral. The community characteristics, total density, total biomass and species richness were only slightly, if at all affected by grazing. However, individual algal species or taxa showed varying and even contrasting responses to grazing, often differently between depths and depending on colonization time. In the deeper littoral, but not close to the water surface, grazing increased the density of filamentous brown algae while reducing the green alga Cladophora glomerata. In these taxa, grazing effects were strongest in patches colonized during the early growing season. Grazing at the colonization stage had lasting consequences for the density of several individual species.     

Vascular plant species richness and rarity across a minerotrophic gradient in wetlands of St. Lawrence County,New York,USA

Thirteen wetlands in St. Lawrence County, NY were sampled to examine the effect of a minerotrophic gradient on vascular plant species richness and rarity. Wetlands ranged from organic soil based poor fens (average conductivity 46.40 microsemens, average Ca 3.55 ppm) to mineral soil based rich fens (average conductivity 342.10 microsemens, Ca 23.00 ppm). Vascular plant species richness was sampled during 1990 in randomly located 1.0 m² quadrats. Specific conductivity, presence or absence of hummocks, and water depth predicted 62% of the variation in richness. Richness increased as conductivity increased until 413 microsemens at which a down trend became obvious. The negative curvilinear relation between conductivity and richness is in accordance with the hump-backed model of Grime but occurs at high rather than intermediate conductivity values. State-listed rare species were found in species-rich wetlands only and had a mean associated richness value of 14.50 species m^-2. This relationship should be taken into consideration when selecting wetlands for protection or managing wetlands for maximum plant diversity.     

Independent and interactive effects of nutrients and grazers on benthic algal community structure

Algal responses to nutrients, grazing by Helicopsyche borealis, and concurrent grazing by Helicopsyche and Baetis tricaudatus were examined in recirculating stream chambers. Alagl communities, dominated by Achnanthes minutissima, Cocconeis placentula, and Synedra ulna, were primarily phosphorus-limited. Algal populations responded after only 6 days of nutrient enrichment. Initially, both the adnate diatom Cocconeis and erect diatom Synedra showed positive response to nutrient enrichment. Accumulation of algal biomass between day 3 and 6 in the P enriched treatment was resulted primarily from the growth of Synedra, an overstory rosette-like diatom colony. Such a shift in dominant growth from adnate to erect diatoms is a general phenomenon in periphyton succession in the absence of disturbance. Algal species showed differential responses to an increase of Helicopsyche densities. The accrual rate of Achnanthes continuously decreased with increasing grazer densities. The accrual rates of both Cocconeis and Synedra declined but reached plateaus between medium and high grazing densities. Baetis effectively and exclusively depressed Synedra and had no significant impact on Cocconeis. After concurrent grazing, algal communities were mainly dominated by Cocconeis (approximately 80% of total algal biovolume). The grazer' s mouth structures, grazing efficiencies, and mobility may account for the differential effects of concurrent grazing on algal communities. Significant interactive effects of P and grazing by Helicopsyche indicated that both nutrient addition and grazing may exert significant impact on algal communities. However, grazing may have a much stronger effect on algae than nutrients. Our results indicate that enhancement of algal biomass by P was dampened by grazing activities and that P had no effect on algal biomass in the presence of grazers.