Disturbance frequency influences patch dynamics in stream benthic algal communities

Disturbance is integral to the organisation of riverine ecosystems. Fluctuating low flows caused by supra-seasonal drought and water management periodically dewater habitat patches, potentially creating heterogeneity in the taxonomic composition and successional dynamics of benthic communities. The frequency of disturbance induced by low flows is contingent upon the topography of the river bed and thus varies among patches. We investigated whether the frequency of patch dewatering influenced the structure and temporal dynamics of benthic algal communities attached to the upper surfaces of stones in stream mesocosms (4 m²). In a 693-day disturbance experiment, we applied short dewatering disturbances (6 days) at high (33-day cycles) and low frequencies (99-day cycles) and compared algal assemblages with undisturbed controls at 21 endpoints. In the absence of disturbance, epilithic space was dominated by the green encrusting alga Gongrosira incrustans. However, drying disturbances consistently reduced the dominance of the green alga, and crust abundance decreased with increasing disturbance frequency, thereby opening space for a diversity of mat-forming diatoms. The response of mat diatoms to disturbance varied markedly during the experiment, from strong reductions in the abundance of loosely attached mats in mid-late 2000 to the exploitation of open space by closely adhering mats in 2001. Contrary responses were attributed to changes in the species composition of mat diatoms, which influenced the physiognomy and hence stress-resistance and resilience of the assemblage. Our results indicate that patchy dewatering of habitat patches during periods of low flow influences the successional dynamics of algae, thereby creating distinctive mosaics on the stream bed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The response of benthic rhizopods to sediment disturbance does not support the intermediate disturbance hypothesis

Disturbance is one of the mechanisms which counteract competitive exclusion of populations in resource-limited communities, thereby facilitating coexistence and maintaining community species diversity. The intermediate disturbance hypothesis predicts maximum diversity at intermediate disturbance intensities and frequencies. This paper reports results of an experimental test of this hypothesis using a coastal benthic community of rhizopods (Protozoa: Rhizopoda), and experimental sediment resuspension as a simulated natural disturbance. We carried out two experiments of 5 d duration which focussed on the effects of resuspension intensity and frequency, respectively, on the abundance, species richness and on the Shannon-Weaver diversity index of rhizopod communities in surface sediments of natural sediment cores from the coastal southern Baltic. Care was taken to adjust the experimental treatments to the natural disturbance regime in this area.
Twenty-four and 28 rhizopod species were present during the intensity and frequency experiment, respectively. Small bacterivorous rhizopods of the Vannellidae, Cochliopodidae, Paramoebidae and Rhizopoda incertae sedis dominated the communities during both experiments. Rhizopod abundance, species richness and diversity increased towards the end of the intensity experiment, but they did not show effects of disturbance intensity. Similarly, no effects of disturbance frequency were found during the frequency experiment. Our results indicate that coexistence and community diversity maintenance in benthic rhizopod communities, and probably in benthic heterotrophic protistan communities in general, may rely on different mechanisms than intermediate disturbance, such as trophic niche separation and high rates of dispersal and colonisation.     

Megafaunal Communities in Rapidly Warming Fjords along the West Antarctic Peninsula: Hotspots of Abundance and Beta Diversity

Glacio-marine fjords occur widely at high latitudes and have been extensively studied in the Arctic, where heavy meltwater inputs and sedimentation yield low benthic faunal abundance and biodiversity in inner-middle fjords. Fjord benthic ecosystems remain poorly studied in the subpolar Antarctic, including those in extensive fjords along the West Antarctic Peninsula (WAP). Here we test ecosystem predictions from Arctic fjords on three subpolar, glacio-marine fjords along the WAP. With seafloor photographic surveys we evaluate benthic megafaunal abundance, community structure, and species diversity, as well as the abundance of demersal nekton and macroalgal detritus, in soft-sediment basins of Andvord, Flandres and Barilari Bays at depths of 436–725 m. We then contrast these fjord sites with three open shelf stations of similar depths. Contrary to Arctic predictions, WAP fjord basins exhibited 3 to 38-fold greater benthic megafaunal abundance than the open shelf, and local species diversity and trophic complexity remained high from outer to inner fjord basins. Furthermore, WAP fjords contained distinct species composition, substantially contributing to beta and gamma diversity at 400–700 m depths along the WAP. The abundance of demersal nekton and macroalgal detritus was also substantially higher in WAP fjords compared to the open shelf. We conclude that WAP fjords are important hotspots of benthic abundance and biodiversity as a consequence of weak meltwater influences, low sedimentation disturbance, and high, varied food inputs. We postulate that WAP fjords differ markedly from their Arctic counterparts because they are in earlier stages of climate warming, and that rapid warming along the WAP will increase meltwater and sediment inputs, deleteriously impacting these biodiversity hotspots. Because WAP fjords also provide important habitat and foraging areas for Antarctic krill and baleen whales, there is an urgent need to develop better understanding of the structure, dynamics and climate-sensitivity of WAP subpolar fjord ecosystems.     

Abrupt change in primary productivity in a littoral zone of Lake Biwa with the development of a filamentous green-algal community

1. Some characteristics of the photosynthesis and primary production of benthic and planktonic algal communities were investigated in a littoral zone covered with gravel in the north basin of Lake Biwa, paying special attention to the recent development of filamentous green algae (FGA) in the benthic algal community.
2. P_max (maximum gross photosynthesis rate) values of the benthic algal community (0.1–1.2 mg C mg chl. a ⁻¹ h⁻¹) obtained from photosynthesis–irradiance (P–I) curves were lower than those of the planktonic algal community (2.4–11.5 mg C mg chl. a ⁻¹ h⁻¹). This is apparently a result of the high degree of self shading in the benthic algal community and its low turnover as compared with that of the planktonic algal community.
3. Relatively high I_k values (150–200 μmol photon m⁻² s⁻¹) were observed in the benthic algal community only in June–July when a FGA, Spirogyra sp., was abundant. This reflected a photosynthetic characteristic of the Spirogyra itself, in which photosynthesis was saturated at high light intensity.
4. The FGA community established in the layer between planktonic and sessile (benthic algae except for FGA) algal communities. It brought about extraordinarily high organic matter production in the littoral zone at the expense of production in the sessile algal community.     

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.     

Quantitative paleoecology of Cambrian (Series 2–Miaolingian) communities from central Sonora,Mexico

This work presents a paleoecological approach to study benthic communities preserved in Cambrian strata exposed at the Sahuaral locality in central Sonora, northwestern Mexico. Based on species richness and abundance, this work analyzed 627 individuals corresponding to 33 species from five distinctive Cambrian formations: the lowermost Proveedora Formation characterized by bioturbation horizons that record the occurrence of a benthic-infaunal biota from coastal shallow seas, followed by the Buelna Formation which contains reef deposits dominated by autotrophic epibenthic communities. The Cerro Prieto Formation was deposited in a high-energy shallow marine environment, dominated by primary consumers-epifaunal and suspensions feeders. The uppermost unit, the El Gavilán Formation, is an intercalation of shale and fossiliferous limestone characterized by the fossil remains of suspension feeders, detritus feeders and nektobenthic organisms of low energy deep water environments. The paleobiota consist of ichnofossils, microbial oncolites, chancellorids, brachiopods, hyolithids, and trilobites in a sequence of increasing diversity and complexity. Paleoecological indices (Simpson Dominance, Shannon-Weaver Index, Evenness Index, and Total Richness) were determined and compared in the study area. In addition, the effective number of species and the true diversities were obtained, which are similar between each station distributed in each formation. However, it is possible that the high degree of dominance is a factor to indicate an inequality between the true diversity and the species richness identified. These results provide new analytical and paleoecological criteria for better understanding depositional environments and fossil associations where the spatial distribution of benthic organisms, richness and abundance provide a valued window to Cambrian biotas. The structure of the marine community of the El Sahuaral area has a great affinity with other Cambrian communities, such as those from the Stephen Formation (Burgess Shale, British Columbia) and the Chengjiang biota of the Yangtze Shelf deposits in South China.     

Algal richness and life‐history strategies are influenced by hydrology and phosphorus in two major subtropical wetlands

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Habitat dynamics,marine reserve status,and the decline and recovery of coral reef fish communities

Severe climatic disturbance events often have major impacts on coral reef communities, generating cycles of decline and recovery, and in some extreme cases, community‐level phase shifts from coral‐ to algal‐dominated states. Benthic habitat changes directly affect reef fish communities, with low coral cover usually associated with low fish diversity and abundance. No‐take marine reserves (NTRs) are widely advocated for conserving biodiversity and enhancing the sustainability of exploited fish populations. Numerous studies have documented positive ecological and socio‐economic benefits of NTRs; however, the ability of NTRs to ameliorate the effects of acute disturbances on coral reefs has seldom been investigated. Here, we test these factors by tracking the dynamics of benthic and fish communities, including the important fishery species, coral trout (Plectropomus spp.), over 8 years in both NTRs and fished areas in the Keppel Island group, Great Barrier Reef, Australia. Two major disturbances impacted the reefs during the monitoring period, a coral bleaching event in 2006 and a freshwater flood plume in 2011. Both disturbances generated significant declines in coral cover and habitat complexity, with subsequent declines in fish abundance and diversity, and pronounced shifts in fish assemblage structure. Coral trout density also declined in response to the loss of live coral, however, the approximately 2:1 density ratio between NTRs and fished zones was maintained over time. The only post‐disturbance refuges for coral trout spawning stocks were within the NTRs that escaped the worst effects of the disturbances. Although NTRs had little discernible effect on the temporal dynamics of benthic or fish communities, it was evident that the post‐disturbance refuges for coral trout spawning stocks within some NTRs may be critically important to regional‐scale population persistence and recovery.     

Biotic disturbance and benthic community dynamics in salmon-bearing streams

1. Organisms can impact ecosystems via multiple pathways, often with positive and negative impacts on inhabitants. Understanding the context dependency of these types of impacts remains challenging. For example, organisms may perform different functions at different densities. 2. Anadromous salmon accumulate > 99% of their lifetime growth in marine ecosystems, and then return to spawn, often at high densities, in relatively confined freshwaters. While previous research has focused on how salmon nutrients can fertilize benthic communities, we examined how an ecosystem engineer, sockeye salmon Oncorhynchus nerka, influences seasonal dynamics of stream benthic communities through their nest-digging activities in south-western Alaska, USA. Benthic invertebrate and algal abundance were quantified every 7-14 days during the open water seasons of 10 streams in riffle and run habitats across multiple years, leading to 25 different stream-year combinations that spanned a large gradient of salmon density. 3. In streams with few or no salmon, benthic algal and insect biomass were fairly constant throughout the season. However, in streams with more than 0.1 salmon m(-2), algal and insect biomass decreased by an average of 75-85% during salmon spawning. Algal biomass recovered quickly following salmon disturbance, occasionally reaching pre-salmon biomass. In contrast, in streams with more than 0.1 salmon m(-2), aquatic insect populations did not recover to pre-salmon levels within the same season. We observed no positive impacts of salmon on algae or insects via fertilization from carcass nutrients. 4. Salmon, when their populations exceed thresholds in spawning density, are an important component of stream disturbance regimes and influence seasonal dynamics of benthic communities. Human activities that drive salmon densities below threshold densities, as has likely happened in many streams, will lead to altered seasonal dynamics of stream communities. Human activities that alter animal populations that are sources of biogenic disturbance can result in shifts in community dynamics.     

井冈山自然保护区蛾类多样性及人为干扰的影响

对井冈山自然保护区蛾类群落的多样性及其受人为干扰的影响进行了研究。结果显示：（1）人为干扰导致井冈山自然保护区蛾类群落构成有较大差别,严重干扰导致蛾类科数明显降低;物种数和个体数呈现出随着人为干扰加强而降低的趋势;蛾类各优势科在不同人为干扰样点的相对多度存在差异。（2）人为干扰导致蛾类主要科的物种数和个体数变化不同：尺蛾科、夜蛾科、毒蛾科和舟蛾科的物种数和个体数在人为干扰下降低;轻度和中度干扰导致螟蛾科和天蛾科的物种数和个体数升高,重度干扰下物种数和个体数降低;人为干扰导致灯蛾科的物种数和苔蛾科的个体数升高。（3）人为干扰导致蛾类群落Shannon多样性指数、Pielou均匀度指数降低,Berger-Parker优势度指数升高。（4）人为干扰对蛾类群落的物种数和个体数的时间动态影响较大,对Shannon指数和Pielou均匀度指数时间动态影响较小,导致Berger-Parker优势度指数时间波动幅度增大。（5）人为干扰导致样点间蛾类群落相似性较低。     

A heat wave and dispersal cause dominance shift and decrease biomass in experimental metacommunities

In experimental metacommunities with marine benthic microalgae, we tested whether heat stress changes effects of connectivity and habitat heterogeneity on metacommunity structure and functioning, by manipulating a simulated heat wave, dispersal frequency and a light intensity gradient. We found that all measures of mean local and regional diversity and community biomass significantly declined after the heat wave and showed no sign of recovery. Additionally, dispersal decreased diversity and increased dominance in both the heat stressed and control communities. Together the heat wave and high dispersal frequency induced a dominance shift by spreading a temperature tolerant but low yielding species from its source patches with low light intensity across the metacommunity, an effect that increased with time. Although different species became dominant at high dispersal frequency with and without the heat wave, the shift towards a temperature tolerant species was not sufficient to maintain total community biomass. Thus, short‐term disturbance may cause longer‐term loss of ecosystem function due to dominance shifts in the composition of communities. This study illustrates the importance of employing multispecies approaches when attempting to predict responses of communities to environmental changes.     

Interaction between disturbance and primary productivity in determining stream invertebrate diversity

Invertebrate diversity patterns were examined in 10 streams that differed in substrate disturbance rates, in Taranaki, New Zealand, between April 1999 and January 2000. Two sites on each stream were sampled, one under native forest canopy where light was postulated to limit periphyton growth and a similar site 225–3800 m downstream in open grassland. Periphyton biomass was considerably higher at open stable sites than at closed or unstable sites. Associated with the higher algal biomass, species number and total abundance of animals were higher at open canopy sites. Species number exhibited a negative linear relationship with disturbance but only at open sites. In contrast, rarefied species richness exhibited a negative linear relationship with disturbance at both open and closed sites. This was a result of communities at the more disturbed sites being numerically dominated by only a few taxa compared to the more evenly distributed communities at stable sites. The observed patterns provide little support for contemporary diversity disturbance models but suggest diversity of invertebrates in streams is a function of time since the last disturbance, mediated through recovery of the food base in autotrophic streams.     

Assessment of littoral algal diversity from the northern Gulf of Mexico using environmental DNA metabarcoding

Traditional methods for algal biodiversity monitoring are costly and time inefficient because they rely on high‐level taxonomic expertise to address species identity problems involving phenotypic plasticity and morphological convergence. These problems are exacerbated in regions such as the Gulf of Mexico, that has a limited history of phycological exploration, but that are economically important or threatened by numerous anthropogenic stressors. Given the high pace of disturbance to natural systems, there is a critical need for expedient and cost‐effective tools for the study of benthic algal communities. Here we document the use of environmental DNA metabarcoding, using the partial LSU rDNA and 23S rDNA plastid molecular markers, to elucidate littoral algal diversity in the Northern Gulf of Mexico. We assigned 73.7% of algal OTUs to genus and 59.6% to species ranks. Our current study detected molecular signals for 35 algal/protist species with no previous reports in the Gulf of Mexico, thus providing an important, molecular‐validated, baseline of species richness for this region. We also make several bioinformatic recommendations for the efficient use of high‐throughput sequence data to assess biological communities.     

Drifting algal mats as an alternative habitat for benthic invertebrates: Species specific responses to a transient resource

Patchy occurrences of benthic drift algae (i.e. loose lying macroalgal mats) may increase habitat complexity on normally bare soft bottoms, but at the same time, extensive amounts of drifting algal mats are known to stress the benthic fauna. This paper presents results of the first detailed study of the fauna associated with drift algal mats in the northern Baltic Sea. In order to assess the importance of drifting algae as an alternative habitat for benthic fauna, benthic drift algal mats were sampled on shallow (2-9 m) sandy soft bottoms in the outer archipelago of the ?land Islands (Finland). Species composition, abundance and biomass of the macrofauna associated with algal mats were recorded. The results show that drifting algae at times can harbour very high abundances of invertebrates (up to 1116 individuals/g algal dryweight), surpassing invertebrate densities recorded in seagrass communities. The algal fauna varied between sites and over time, and factors such as ambient benthic fauna, exposure to wind-wave disturbance, depth, and algal coverage and condition influenced the invertebrate community composition of the algal mats. Abundance increased while individual biomass of the animals decreased over time (summer season; July-October). A series of laboratory experiments were conducted in order to test the ability of a few important benthic species to move up into, and survive in a drifting algal mat. Macoma balthica, Hydrobia spp., Nereis diversicolor and Bathyporeia pilosa were used in the experiments, and significant differences in their survival and mobility within drifting algae were recorded. This study shows that benthic species differ significantly in their ability to utilise the algal mats, with mainly opportunistic and mobile taxa such as Hydrobia spp., Chironomidae and Ostracoda benefiting from the algae, whereas infaunal species such as M. balthica and B. pilosa are negatively affected. The occurrence of eutrophication induced drifting macroalgal mats has increased significantly during the last decade in the northern Baltic Sea. Hence, the importance of drifting algae as a stress factor and as an alternative habitat for benthic fauna increases.     

长白山北坡不同海拔及干扰程度下访花食蚜蝇群落的差异

本研究采用垂直面取样法,对中国长白山北坡不同海拔及干扰程度下访花食蚜蝇群落的物种组成、多度、丰富度及多样性进行了调查研究。我们选择了不同垂直带的两个样地,分别位于800~1 100 m和1 750 ~2 150 m;每个样地选取3种生境类型,每一生境类型采样面积为20 m×20 m。252组调查数据表明,共采到访花食蚜蝇42种2 540个体。不同海拔生境访花食蚜蝇的组成、多度、丰富度及多样性的差异分析结果表明: 低海拔样地内不同生境访花食蚜蝇的群落相似性高于高海拔样地;低海拔样地访花食蚜蝇的多度高于高海拔样地,而物种丰富度低于高海拔样地;不同生境类型访花食蚜蝇多样性指数存在差异,但高海拔样地与低海拔样地多样性差异不显著;过渡林生境（岳桦林带）与次生林生境（针阔混交林带Ⅱ）中访花食蚜蝇的多样性较高。保护珍稀植物物种的人工植物花园访花食蚜蝇多样性仅次于过渡林生境（岳桦林带）与次生林生境（针阔混交林带Ⅱ）,保护作用显著。     

168 Cell Wall Proteomics of the Green Alga Haematococcus Pluvialis (Chlorophyceae)

Coral reefs of US-held islands in the central Pacific Ocean are among the most pristine in the world and represent over 93% of the reef systems under United States jurisdiction. The remote location of many islands has limited past algal research, resulting in incomplete understanding of species diversity, quantity, and ecology. Starting in 2000, the Coral Reef Ecosystem Investigation (CREI) began rapid ecological assessments on many Pacific island reefs to monitor ecological changes in reef biota over time. During the past year, algal efforts have concentrated on the French Frigate Shoals (Northwestern Hawaiian Islands) where we have increased the number of algal species reported by 1000%. Additionally, species new to science, including Acrosymphyton brainardii and Scinaia huismanii, have been described. Quantitative field sampling using a photoquadrat method is revealing species of the green algae Halimeda and Microdicyton to be ecological dominants in many areas during late summer/early autumn. Preliminary analyses with Primer software show species composition and abundance of all benthic organisms to differ significantly between most field sites sampled. Additional benthic habitat mapping of Pacific island reefs by CREI researchers is breaking the long-held paradigm that macroalgal cover is minimal in healthy tropical reef systems. Videotape analyses of benthic communities often find over 50% algal cover from 1 to 20 meter depths in many locations. Common ratios of macroalgae, turf algae, and crustose coralline algae to corals, other benthic organisms and substrate types on US Pacific reefs are being calculated for the first time.     

Bathymetric distribution and diversity of deep water polychaetous annelids in the Sigsbee Basin, northwestern Gulf of Mexico

The deep water polychaete fauna is analyzed in this study particularly regarding its composition and variations with depth in the Sigsbee Basin, northwestern region of the Gulf of Mexico. Samples were taken at 10 stations along a bathymetric gradient with depth ranges from 200 to 3760 m with a USNEL (0.25 m²) corer. A total of 287 individuals were identified, from 21 families and 65 species. The most important families, both in terms of abundance and species richness, were: Paraonidae (65.4 ind./0.25 m², 9 spp.), Cirratulidae (28.93 ind./0.25 m², 7 spp.) and Spionidae (18.07 ind./0.25 m², 7 spp.). In general, density tended to decrease with depth with minima at around 2000 m, although two abundance peaks were detected at 3700 and 3760 m, making the pattern seem an inverted parabolic curve. The Shannon-Wiener diversity values varied from 0.54–0.92 at around 2000 m to 3.39 at 3620 m and 3.34 at 3760 m. These results contrast with what is already reported from the North Atlantic and the Tropical Pacific deep benthic communities, where highest diversities are found at 2000 m. Faunal changes evaluated through Beta Diversity (0.08–0.1) and the low similarity found between the stations, emphasized the high variability in the composition of the fauna in the Sigsee Basin, meaning that the faunal composition is practically different at all the sampling stations. Ten species are newly recorded for the Mexican fauna.     

Seasonal patterns of sediment loading and benthic invertebrate community dynamics in Lake Tanganyika, Africa

1. This study investigated the spatiotemporal dynamics of benthic invertebrate communities located close to the mouths of the Kalambo and Lunzua Rivers, at the southern end of Lake Tanganyika, and subject to varying degrees of sediment load. 2. Metrics of exposure to riverine sediments were associated inversely with abundance and diversity of benthic organisms at sampling locations adjacent to both river mouths, indicating a major effect of riverine sediments on the structure of near‐shore benthic communities. 3. A lower abundance of benthos was found at the mouth of the Lunzua River, which exports significantly higher sediment loads than the Kalambo. 4. Seasonal cycles of abundance and diversity in the benthos varied with distance from river mouths. This has important implications for monitoring programmes investigating benthic communities close to the mouths of rivers. 5. Our findings suggest that recent increases in the sediment loading of Lake Tanganyika, owing to anthropogenic catchment disturbance, impact significantly on biological diversity throughout the lake littoral, which provides habitat for the majority of species in the lake.     

Planktonic communities of melt ponds on the McMurdo Ice Shelf,Antarctica

The planktonic community of 20 melt ponds on the McMurdo Ice Shelf was investigated to determine taxa abundance and diversity and the controlling environmental variables. Grazing rates were measured using fluorescent beads to examine trophic interactions between ciliates, bacteria and phytoplankton. The melt ponds contained a surprisingly varied planktonic community with relatively high abundance compared with Antarctic continental lakes. There was a clear distinction between small, productive ponds dominated by bactivorous small ciliates, hymenostomes and heterotrophic cryptophytes and the larger, less productive ponds where these taxa were less abundant. The benthic mats of cyanobacteria and diatoms were potentially a source of food for some ciliate species but the majority were bacterivores. The lack of large herbivorous ciliates, the heterotrophic capabilities of cryptophytes and the broad ecological tolerances contributed to a planktonic community dominated by cryptophytes.     

Algal species composition and phosphorus recycling at contrasting grazing pressure: an experimental study in sub-Antarctic lakes with two trophic levels

1. To test the response of algal communities to altered grazer abundance in lakes lacking efficient predators on herbivores, we performed field and experimental studies in two sub-Antarctic lakes (South Georgia).
2. The number of algal species in these high latitude lakes is low, and all dominant species have grazer-resistant adaptations, including spines in three dimensions ( Staurastrum sp.), large size ( Tribonema sp.), a mucus sheet allowing viable passage through the gut ( Chlamydocapsa sp., Elakatothrix sp.) or ability to recruit individuals from the sediment surface ( Mallomonas sp.).
3. Algal community composition was only slightly changed by experimentally altered grazer abundance, indicating that it was already adapted for a high grazing pressure. Hence, the diets of herbivores were restricted to vulnerable food organisms such as Mallomonas sp. and heterotrophic flagellates in the water column, and to benthic food sources.
4. At high grazer abundance, the concentration of available phosphorus (PO₄-P) in the water was lower than at low grazer abundances, due to inefficient nutrient regeneration by the copepod herbivores. Hence, in lakes where copepods are dominant grazers, algae suffer both directly from grazing and indirectly from reduced nutrient availability.