Predation risk and the structure of freshwater zooplankton communities

Summary Many predators inflict substantial mortality on their prey. The prey respond to these selective pressures with changes in their spatial and temporal overlap with the predator (density risk responses), or with changes in their vulnerability to the predator (prey vulnerability responses). Here we develop a simple predation model that permits quantification of the basic response types of the prey in nature. We then test the hypothesis that prey response will be proportional to the intensity of the predation mortality relative to all other sources of mortality and decreased natality acting on the prey. A significant regression relationship is obtained for the prey vulnerability response but not for any of the density risk responses. The individual response values and regression statistics are used to interpret the relative importance of the different response types and to assess the predator's influence on prey community structure.Supported by Lehigh University Environmental Studies Center     

Zooplankton community structure driven by vertebrate and invertebrate predators

Summary A zooplankton community was established in outdoor experimental ponds, into which a vertebrate predator (topmouth gudgeon: Pseudorasbora parva) and/or an invertebrate predator (phantom midge larva: Chaoborus flavicans) were introduced and their predation effects on the zooplankton community structure were evaluated. In the ponds which had Chaoborus but not fish, small- and medium-sized cladocerans and calanoid copepods were eliminated while rotifers became abundant. A large-sized cladoceran Daphnia longispina, whose juveniles had high helmets and long tailspines as anti-predator devices, escaped from Chaoborus predation and increased. In the ponds which had fish but not Chaoborus, the large-sized Daphnia was selectively predated by the fish while small-and medium-sized cladocerans and calanoid copepods predominated. In the ponds containing both Chaoborus and fish, the fish reduced the late instar larvae (III and IV) of Chaoborus but increased the early instar larvae (I and II). Small- and large-sized cladocerans were scarcely found. The former might have been eliminated by predation of the early instar larvae of Chaoborus, while the latter was probably predated by fish. Consequently, the medium-sized cladocerans, which may have succeeded in escaping from both types of predator, appeared abundantly. The results suggest that various combinations of vertebrate and invertebrate predators are able to drive various kinds of zooplankton community structure.     

Zooplankton distribution and dynamics in a temperate shallow estuary

The spatial, temporal and tidal dynamics of the zooplanktonic community of the Mondego estuary was studied from January 2003 to 2004. The monthly sampling procedure included the measurement of hydrological parameters (salinity, temperature, Secchi transparency, chlorophyll a and nutrients) and the collection of zooplankton with a Bongo net of 335 μm mesh size. Zooplankton composition, distribution, density, biomass and diversity were determined. The principal component analysis (PCA) revealed the existence of a spatial gradient with the upstream sampling stations, associated to high values of nutrients, in opposition to the downstream stations characterized by higher salinity and transparency values. The Copepoda was the main dominant group and Acartia tonsa revealed to be the more abundant taxon. The spatial and temporal dynamics of zooplanktonic communities analysed by non-metric MDS showed the existence of four assemblages of species-sites, reflecting differences in zooplankton composition between both branches of the estuary. The results suggest that abundance, biomass and diversity of the zooplanktonic community are strongly influenced by the hydrological circulation pattern and by direct or indirect human impacts that occur in each branch. The northern branch is dominated by the river flow suffering from regular dredging activities and the southern branch is dominated by tidal circulation suffering from an ongoing eutrophication process.     

Biotic relations affecting species structure in zooplankton communities

Mathematical modelling and laboratory experiments were used to study how exploitative competition and predation influence the species structure in cladoceran community. For five species of Cladocera (Sida crystallina, Daphnia magna, Simocephalus vetulus, Daphnia longispina, and Diaphanosoma brachyurum), representing a gradient of body size, population characteristics were described as functions of food concentration. Abundance dynamics were simulated in mixed species cultures and invasion experiments under different levels of food supply corresponding to oligo-, meso-, and eutrophic conditions. Separate simulations were also run including and excluding (fish) predation. The competitive ability of each species was estimated as the values of the population equilibrium food concentration. Simulation results showed that for the no-predator scenario, increases in the level of food supply promoted species coexistence while under lower food concentrations only one species remained at the end of the simulation runs. When predation was allowed, the number of species that coexisted at the end of the simulations increased up to four species, indicating that predator pressure facilitated species coexistence because it shortened periods of food depletion. Simulation results were verified in laboratory experiments which suggested that population equilibrium food concentration can be used as an estimate of competitive ability. Finally, species structure and relative abundance in Lake Naroch (Belarus) during the summer of 2004 was found to be consistent with our results from computer simulation and laboratory experiments with regard to competition and predation impacts on zooplankton community. Handling editor: S. I. Dodson     

Calanoid copepod grazing on phytoplankton: seasonal experiments on natural communities

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

Mesozooplankton biomass and indices of grazing and metabolic activity in Antarctic waters

Mesozooplankton abundance, body area spectrum, biomass, gut fluorescence and electron transfer system (ETS) activity were studied in the Antarctic Peninsula during the post-bloom scenario in these waters. Values of abundance and biomass were rather low and decreased sharply from the slope waters to the coastal area. In contrast, specific gut fluorescence and ETS activity were high in the coastal area and decreased through the shelf-break. Large copepods were very scarce, similarly to the post-bloom conditions in phytoplankton where large cells are not abundant and small cells such as flagellates dominate the water column. The vertical distribution showed two well defined layers by day, one at the surface which corresponded to krill organisms and a second at depth (>300 m) formed mainly by the large copepod Metridia gerlachei. During the short night, this layer ascended at the time that krill at the surface migrated to deeper waters as observed from acoustics and net sampling. This observation and the absence of large copepods over the shelf suggest that krill consumption on large phytoplankton cells during the bloom is followed by an increase in predation upon mesozooplankton. It also suggests that krill decrease the abundance and biomass of mesozooplankton over the shelf and continues their predation upon mesopelagic copepods during the post-bloom in Antarctic waters. This behaviour could explain the long ago described impoverishment in mesozooplankton south of the Antarctic Circumpolar Current.     

武汉东湖浮游动物群落结构的研究

对１９９１年武汉东湖不同湖区浮游动物群落结构的研究表明,Ⅰ、Ⅱ、Ⅲ站浮游动物的数量为１２２７８２、７０４１５和１８１８４个·Ｌ（－１）,并随水体富营养化程度增加而增加．生物量分别为８．６３９、５．８７２和２．０８２ｍｇ·Ｌ（－１）。大于１ｍｍ的甲壳动物个体频度为０．１５、０．３０、０．４１．其体长随鱼产量的增加而缩小．最后,对影响东湖浮游动物群落结构的主要生态因子进行了讨论．     

Flagellate predation on a bacterial model community: interplay of size-selective grazing, specific bacterial cell size, and bacterial community composition.

The influence of grazing by the bacterivorous nanoflagellate Ochromonas sp. strain DS on the taxonomic and morphological structures of a complex bacterial community was studied in one-stage chemostat experiments. A bacterial community, consisting of at least 30 different strains, was fed with a complex carbon source under conditions of low growth rate (0.5 day(-1) when nongrazed) and low substrate concentration (9 mg liter(-1)). Before and after the introduction of the predator, the bacterial community composition was studied by in situ techniques (immunofluorescence microscopy and fluorescent in situ hybridization), as well as by cultivation on agar media. The cell sizes of nonspecifically stained and immunofluorescently labeled bacteria were measured by image analysis. Grazing by the flagellate caused a bidirectional change in the morphological structure of the community. Medium-size bacterial cells, which dominated the nongrazed community, were largely replaced by smaller cells, as well as by cells contained in large multicellular flocs. Cell morphological changes were combined with community taxonomic changes. After introduction of the flagellate, the dominating strains with medium-size cells were largely replaced by single-celled strains with smaller cells on the one hand and, on the other hand, by Pseudomonas sp. strain MWH1, which formed the large, floc-like forms. We assume that size-selective grazing was the major force controlling both the morphological and the taxonomic structures of the model community.     

The plankton community of Lake Matano: factors regulating plankton composition and relative abundance in an ancient,tropical lake of Indonesia

Recent evidence reveals that food webs within the Malili Lakes, Sulawesi, Indonesia, support community assemblages that are made up primarily of endemic species. It has been suggested that many of the species radiations, as well as the paucity of cosmopolitan species in the lakes, are related to resource limitation. In order to substantiate the possibility that resource limitation is playing such an important role, a study of the phytoplankton and zooplankton communities of Lake Matano was implemented between 2000 and 2004. We determined species diversity, relative abundances, size ranges, and total biomass for the phytoplankton and zooplankton, including the distribution of ovigerous individuals throughout the epilimnion of Lake Matano in three field seasons. The phytoplankton community exhibited very low biomass (<15 μg l^?1) and species richness was depressed. The zooplankton assemblage was also limited in biomass (2.5 mg l^?1) and consisted only of three taxa including the endemic calanoid Eodiaptomus wolterecki var. matanensis, the endemic cyclopoid, Tropocyclops matanensis and the rotifer Horaella brehmi. Zooplankton were very small (<600 μm body length), and spatial habitat partitioning was observed, with Tropocylops being confined to below 80 m, while rotifer and calanoid species were consistently observed above 80 m. Less than 0.1% of the calanoid copepods in each year were egg-bearing, suggesting very low population turnover rates. It was concluded that chemical factors as opposed to physical or biological processes were regulating the observed very low standing crops of phytoplankton which in turn supports a very minimal zooplankton community restricted in both species composition and abundance. As chemical factors are a function of the catchment basin of Lake Matano, it is predicted that resource limitation has long played an important role in shaping the unique endemic assemblages currently observed in the food web of the lake.     

Effects of flowering plant's patch size on species composition of pollinator communities,foraging strategies,and resource partitioning in bumblebees (Hymenoptera: Apidae)

Summary In 4 common Middle-European mainly bumblebee-pollinated plant species (Impatiens glandulifera, Echium vulgare, Aconitum napellus, Symphytum officinale) the influence of patch size on species composition of the pollinator community was studied. Short-tongued species were most dominant in large patches, while small patches were frequented by middle- and long-tongued bumblebees. This phenomenon was extremely obvious in Symphytum officinale and Aconitum napellus, where short-tongued species had bitten a hole in nearly every flower of large patches. Long-tongued species were forced to small patches, where nectarrobbing occurred only exceptionally. In small patches visitationrate (Number of visits per flower per hour) was not lower but either equal or even higher then in large patches. Nectar measurements in Echium vulgare showed, that not only the mean quantity of nectar but also the variance was lower in small patches. As a result, the possible gain can be predicted much more precisely in a small patch than in a large one, and bumblebees have less difficulties in making the right foraging decisions. According to this, foraging strategies depend on patch size. This was confirmed by a computer simulation. The conclusion can be drawn, that many bumblebee species are able to share the same resource by using different patch sizes. Since large flower patches occur mainly in man-made habitats, the dominance of short-tongued species in many bumblebee communities studied by other authors may be unnatural.Supported by the Landesgraduiertenförderung Baden-Württemberg     

Group size structure affects patterns of aggression in larval salamanders

The potential importance of intrapopulation phenotypic variabilityto population-level ecology has been demonstrated in both theoreticaland field studies. One way to connect individuals to the dynamicsof populations they compose is to study behavioral response(an individual-level characteristic) to variability in conspecificphenotypes (a population-level characteristic). We examinedeffects of variation in size of individuals on patterns of aggressionin larval tiger salamanders (Ambystoma tigrinum nebulosum) byobserving aggressive behavior in groups of three larvae in alaboratory experment. We assessed effects of variability insize of conspecifics independently of mean larval size and larvaldensity Overall levels of aggression were generally higher ingroups in which all individuals were similprly sized than ingroups of variably sized individuals. Medium-sized individualsexhibited significantly higher levels of aggression and wereattacked significantly more often when in groups consistingonly of similarly sized larvae as compared to groups composedof larvae representing a wider range of body sizes. Activitylevels of larvae were also generally lower when all individualswere the same size, resulting in a negative correlation betweenactivity and levels of iggressititi. These results suggest thatgioups of similarly sized individuals experience a more aggressivesocial environment than groups of variably sized individuals,and they suggest a promising avenue of research for connectingindividual behavioral and physiological responses to size structure(phenotypic variability) with population dynamics.     

Warming alters community size structure and ecosystem functioning

Global warming can affect all levels of biological complexity, though we currently understand least about its potential impact on communities and ecosystems. At the ecosystem level, warming has the capacity to alter the structure of communities and the rates of key ecosystem processes they mediate. Here we assessed the effects of a 4°C rise in temperature on the size structure and taxonomic composition of benthic communities in aquatic mesocosms, and the rates of detrital decomposition they mediated. Warming had no effect on biodiversity, but altered community size structure in two ways. In spring, warmer systems exhibited steeper size spectra driven by declines in total community biomass and the proportion of large organisms. By contrast, in autumn, warmer systems had shallower size spectra driven by elevated total community biomass and a greater proportion of large organisms. Community-level shifts were mirrored by changes in decomposition rates. Temperature-corrected microbial and macrofaunal decomposition rates reflected the shifts in community structure and were strongly correlated with biomass across mesocosms. Our study demonstrates that the 4°C rise in temperature expected by the end of the century has the potential to alter the structure and functioning of aquatic ecosystems profoundly, as well as the intimate linkages between these levels of ecological organization.     

Zooplankton feeding on algae and bacteria under ice in Lake Druzhby,East Antarctica

The feeding of the cladoceran Daphniopsis studeri on algae and bacteria was investigated under ice in an ultra-oligotrophic Antarctic lake from late autumn (May) to early spring (October) in 2004. D. studeri fed on both algae and bacteria with estimated filtering rates of 0.048 and 0.061 l ind⁻¹ day⁻¹), respectively. Algae seemed to be the major food resource for the D. studeri population, however at times of low algal densities the bacterioplankton represented an important alternative food resource. The D. studeri grazing impact on the algal and bacterial standing stock was in general low (0.6–4.6% removed per day), but during the winter period this organism can remove up to 34% of the bacterial production (BP). At times D. studeri grazing can temporarily have a significant impact on the BP rates, though their impact was relatively low when compared to viral-induced bacterial mortality in the lake.     

Resource availability affects the structure of a natural bacteria-bacteriophage community

Antagonistic networks are known to be structured in the wild, but knowledge on how this structure may change as a response to environmental perturbations is scarce. We describe a natural bipartite network between bacteria and lytic bacteriophages, and investigate how it is affected by environmental productivity in the form of different resource levels for the bacteria. We report that low amounts of resource decrease phage generality and lead to less robust and less stable communities. We discuss how resource levels in nature may alter the structure of complex communities.     

Effect of grazing on community structure and productivity of a Uruguayan grassland

Grasslands and their grazers provide some of the most compelling examples for studying the relationship between diversity, productivity, and disturbance. In this study, we analyzed the impact of grazing-induced changes in species composition and community structure upon the productivity of a grassland in the Campos region, Uruguay. We compared three treatments: a continuously grazed area, a 9-year old exclosure to domestic herbivores, and grazing-simulated plots inside the exclosure, which were clipped so that their standing biomass resembled that of the grazed area. We studied the community composition of the grazed and ungrazed situations, and determined biomass and above-ground net primary production (ANPP) of the three treatments during 1 year. Grazed plots had higher species richness and diversity than the exclosure. Grazing resulted in the replacement of some cool-season, tussock grasses by warm-season, prostrate grasses. ANPP was 51% higher under grazing than in the exclosure, but the grazing-simulated plots inside the exclosure were the most productive treatment, 29% higher than the grazed plots. Thus, two components of grazing effect may be postulated for this grassland. The structural component resulted in higher ANPP, probably due to the elimination of standing dead biomass. The species composition component resulted in lower ANPP once the structural component was controlled, probably due to the shift to warm-season phenology and prostrate habit. Our findings contrast with a similar experiment carried out in the neighbouring Flooding Pampa region, which suggests that the relationship between grazing and community structure and function is difficult to generalize.     

Plankton community structure and size spectra in the Georgian Bay and North Channel ecosystems

Patterns in the size distribution and taxonomic composition of phytoplankton and zooplankton communities for 1974 in Georgian Bay and the North Channel are described. The Diatomeae predominate the phytoplankton in both areas. Copepods, particularly Calanoida, comprise the greatest fraction of the zooplankton biomass. Normalized plankton biomass spectra for both ecosystems are typical of those found in Lake Superior and offshore Lake Huron. The plankton communities of Georgian Bay and the North Channel are thus similar to the most oligotrophic of the Laurentian Great Lakes.     

Size structure of the metazoan community in a Piedmont stream

We characterized the size structure of virtually the entire metazoan community in a fourth order, sandybottomed Piedmont stream during late summer. Our study, the first to sample across all habitat types and sizes of metazoans in an aquatic ecosystem, indicates that at the community level, stream size spectra may be bimodal for the benthos or trimodal when fish are included. Animals spanning 10 orders of magnitude in dry mass (from gastrotrichs to fish) were quantitatively collected from nine habitat types. The bimodal benthic size spectrum was characterized by a meiofaunal component (mostly oligochaetes and micro-crustacea) and a macrobenthic component (mostly the introduced asiatic clam, Corbicula fluminea). Insects contributed little to overall standing crop. Size-specific contribution to whole-community metabolism was assessed using allometric equations for respiration, and we found a distinctly bimodal distribution across the entire metazoan size range, with peaks in the meiofaunal and benthic macrofaunal size ranges. Our bimodal benthic size spectrum is similar to that observed for marine benthos but not to other freshwater benthic systems, possibly because the entire range of habitat types and/or animal sizes were not sampled in the latter. Numerous factors may influence size spectra in stream ecosystems, including local geomorphic (habitat) conditions, water level fluctuations, species introductions, and predation processes.     

Increasing community size and connectance can increase stability in competitive communities

The relationship between community complexity and stability has been the subject of an enduring debate in ecology over the last 50 years. Results from early model communities showed that increased complexity is associated with decreased local stability. I demonstrate that increasing both the number of species in a community and the connectance between these species results in an increased probability of local stability in discrete-time competitive communities, when some species would show unstable dynamics in the absence of competition. This is shown analytically for a simple case and across a wider range of community sizes using simulations, where individual species have dynamics that can range from stable point equilibria to periodic or more complex. Increasing the number of competitive links in the community reduces per-capita growth rates through an increase in competitive feedback, stabilising oscillating dynamics. This result was robust to the introduction of a trade-off between competitive ability and intrinsic growth rate and changes in species interaction strengths. This throws new light on the discrepancy between the theoretical view that increased complexity reduces stability and the empirical view that more complex systems are more likely to be stable, giving one explanation for the relative lack of complex dynamics found in natural systems. I examine how these results relate to diversity-biomass stability relationships and show that an analytical solution derived in the region of stable equilibrium dynamics captures many features of the change in biomass fluctuations with community size in communities including species with oscillating dynamics.     

The interactive effects of temperature,food level and maternal phenotype on offspring size in Daphnia magna

Invertebrate offspring are usually larger in colder environments. To test for possible effects of covariates (e.g. maternal phenotype and feeding conditions) on this pattern, we performed a laboratory experiment to look at the effect of temperature on newborn weight in the planktonic crustacean Daphnia magna. Three tempèratures (12°C, 16°C and 22°C) and two food levels (10,000 cells ml^–1 and 150,000 cells ml^–1) were used, and offspring were examined from the first five clutches of mothers that had been maintained under the constant experimental conditions for three generations. Preliminary analysis suggested that newborn weight was significantly affected by temperature although patterns in the data were not clear cut. In addition, the covariates mother weight and clutch size were positively and negatively correlated with newborn weight, respectively; and later clutches tended to contain heavier offspring. Therefore, in an effort to control for the effects of the covariates, repeated-measures analysis of covariance was performed using ratio values of newborn weight/mother weight (relative newborn weight) as the dependent variable, clutch size as the covariate and clutch number as the repeated measures term. Now, temperature as a main effect in an ANCOVA model did not significantly influence relative newborn weight. The repeatedmeasure term clutch number also became nonsignificant, indicating that when differences in mother weight due to age were accounted for there were no overall differences in relative newborn weight between clutches from a particular mother. Temperature effects on relative newborn weight were only significant as part of interaction terms with food concentration and with clutch number. Thus there were different weight responses to temperature within food levels, and between clutch numbers within food levels. Under the low-food conditions newborn were heaviest at 16°C, lightest at 12°C and intermediate at 22°C. Conversely, under the high-food condition newborn were lightest at 16°C, heaviest at 12°C and again intermediate at 22°C. However, newborn tended to be heavier under the low food condition, and food concentration was highly significant as a main effect. Mother growth rate showed no significant relationship with newborn weight. It is concluded that direct temperature effects on relative newborn weight are marginal and nonsignificant. Temperature effects through interactions with food concentration and clutch number are important determinants of newborn weight, but relatively speaking account for only a small proportion of observed variance in newborn weight (25%), compared with the direct effect of food concentration (67%).