Invasibility of a reservoir to exotic Daphnia lumholtzi: experimental assessment of diet selection and life history responses to cyanobacteria

SUMMARY 1. In reservoirs of the south-central United States, the exotic cladoceran Daphnia lumholtzi is common during warm midsummer conditions, when cyanobacteria are abundant and native Daphnia are rare. In the current study, we employed surveys, field experiments, and a life table experiment to investigate the role of food quality in explaining the distribution and phenology of D. lumholtzi , relative to two native species ( Daphnia parvula and Daphnia mendotae ).
2. During May–September 2000 in eutrophic McDaniel Lake, Missouri U.S.A., cyanobacteria (primarily Oscillatoria ) first appeared at 6-m depth and then became abundant throughout the epilimnion.
3. During the May field experiment, D. lumholtzi , D. parvula and D. mendotae all consumed a similar diet of algae, showing positive selection for small greens (chlorophytes and cryptophytes <20 μm). During the July experiment, when the epilimnion exceeded 25 °C and cyanobacteria were common in the lake, D. lumholtzi consumed significantly more total algae and more cyanobacteria than the two native species. Although the Daphnia selected against cyanobacteria, all three species consumed about 25% of this food in their diet.
4. A life table experiment compared the responses of D. lumholtzi and D. parvula with variation in density of high-quality food ( Ankistrodesmus ) and concentration of a toxic strain of cyanobacteria ( Anabaena flos-aquae ). Both Daphnia species showed reduced survivorship, fertility and intrinsic rates of increase in response to elevated concentrations of cyanobacteria, particularly at the higher food level.
5. The results suggest that D. lumholtzi shows similar inhibition from cyanobacteria as does the native Daphnia . However, their continued high in situ feeding rates imply that D. lumholtzi is less affected by midsummer conditions in warm-water reservoirs than are native Daphnia .     

Differing effects of suspended sediments on the performance of native and exotic Daphnia

1. Although large cladocerans are usually uncommon in large rivers, Daphnia lumholtzi (an exotic species in North America) is widespread and occasionally abundant. We collected zooplankton on the Illinois River (Illinois, U.S.A.) in 1995 and 1996 and found that the peak density of D. lumholtzi (25 L⁻¹) typically exceeded that of all native species combined. Maximum density occurred during warm periods (up to 27 °C) when concentrations of inorganic suspended sediments were high (>50 mg L⁻¹).
2. Using a life table experiment, we examined the effects of variation in suspended sediment (0 and 80 mg L⁻¹) and food (10⁴ and 10⁵ Ankistrodesmus cells mL⁻¹) on fitness of D. lumholtzi and the native Daphnia parvula. Daphnia lumholtzi had greater survivorship than D. parvula in most treatments and higher life-time fertility than D. parvula in all treatments. Both species achieved their fastest intrinsic rates of growth in treatments with high food, but their responses to suspended solids differed. The growth rate of D. lumholtzi in high food was slightly increased by higher turbidity, whereas that of D. parvula was depressed.
3. Results suggest that the ability of D. lumholtzi to tolerate suspended solids is an important factor contributing to its success in invading North American rivers.     

Influence of Temperature on Exotic Daphnia lumholtzi and Implications for Invasion Success

We investigated the effects of water temperature on the exoticcladoceran Daphnia lumholtzi in a eutrophic Kansas reservoir(USA) and under laboratory conditions. Daphnia lumholtzi demonstrateda distinct late summer appearance in the reservoir at temperaturesbetween 26 and 31°C, which corresponded with steep declinesin the densities of native Daphnia spp. Laboratory life-tableexperiments confirmed that D. lumholtzi performs well at elevatedwater temperatures. The intrinsic rate of increase (r), netreproductive rate (R_o), age at first reproduction, survivorship(l_x) and molting rates all demonstrate that D. lumholtzi hasa high temperature optimum between 20 and 30°C. A comparisonof literature-reported r-values indicates that the reproductiverate of D. lumholtzi is comparable with other Daphnia spp. between20 and 25°C, but also implies that D. lumholtzi may out-performsome Daphnia spp. at temperatures >25°C. Collectively,these results suggest that D. lumholtzi may be taking advantageof a late summer thermal niche, and that this invader may continueto colonize lakes and reservoirs in the southern US. However,life table data also indicate that D. lumholtzi performs poorlyat temperatures < 10°C, which may inhibit the range expansionof this invader into northern waters.     

Invasibility of plankton food webs along a trophic state gradient

Biological invasions are becoming more common, yet the majority of introduced exotic species fail to establish viable populations in new environments. Current ecological research suggests that invasion success may be determined by properties of the native ecosystem, such as the supply rate of limiting nutrients (i.e. trophic state). We examined how trophic state influences invasion success by introducing an exotic zooplankter, Daphnia lumholtzi into native plankton communities in a series of experimental mesocosms exposed to a strong nutrient gradient. We predicted that the attributes of nutrient-enriched communities would increase the likelihood of a successful invasion attempt by D. lumholtzi . Contrary to our original predictions, we found that D. lumholtzi was often absent from mesocosms that developed under high nutrient supply rates. Instead, the presence of D. lumholtzi was associated with systems that had low nutrients, low zooplankton biomass, and high zooplankton species diversity. Using generalized estimating equations (GEE) and multivariate species data, we found that the presence–absence of D. lumholtzi could be explained by variations in zooplankton community structure, which was itself strongly influenced by nutrient supply rate. We argue that the apparent invasion success of D. lumholtzi was inhibited by the dominance of another cladoceran species, Chydorus sphaericus . These results suggest that the interaction between trophic state and species identity influenced the invasion success of introduced D. lumholtzi .     

An analysis of seasonal dynamics of a mixed population of Daphnia, and the associated cladoceran community

SUMMARY. The dynamics of a mixed population of Daphnia (D. ambigua and D. parvula) are analysed over 9 months in a small eutrophic lake. A spring peak of D. ambigua occurred shortly after ice break-up. Decline of D. ambigua coincided with a high occurrence of sexual reproduction, the appearance of D. parvula which did not become abundant, and a substantial increase in Ceriodaphnia quadrangula . A third cladoceran, Bosmina longirostris , was abundant before and after the period of numerical dominance by D. ambigua , but declined to very low numbers coincident with the increase in D. ambigua.
Estimates of egg hatching rate at various temperatures demonstrate that these three cladocerans have different rate responses to temperature, and that seasonal succession of species is predictable from temperature alone. The importance of temperature versus species interactions in controlling species succession is discussed.     

Predation by inland silversides on an exotic cladoceran, Daphnia lumholtzi, in Lake Texoma, U.S.A.

In the early 1990s, Lake Texoma was invaded by Daphnia lumholtzi , a large, spiny, exoticcladoceran. From April 1994 to April 1995 D. lumholtzi was most abundant in the lake in mid-summer, after native zooplankton declined in early June. Inland silversides Menidia beryllina selectively preyed on D. lumholtzi but appeared to prefer the large native zooplankton when they were present. Daphnia lumholtzi was an important prey item for silversides during summer when large native zooplankton was scarce. The invasion of D. lumholtzi into Lake Texoma may benefit some zooplanktivorous fishes by increasing foraging opportunities during a time of low prey availability.     

Factors which affect the abundance of an invasive cladoceran, Daphnia lumholtzi, in U.S. reservoirs

1.  Daphnia lumholtzi is a subtropical, Old World species which is rapidly spreading throughout reservoirs in the southern U.S.A. It was first recorded in Lake Texoma (Oklahoma–Texas) in September 1991.
2. Southern U.S. reservoirs typically have strong spatial and temporal gradients in temperature, conductivity, turbidity and in the distribution of organisms. Therefore, the present experiments examined the reproductive and moulting rates, and survival of D . lumholtzi in relation to extremes of food concentration, temperature, conductivity and turbidity.
3. Increases in temperature (range = 15–29 °C) and decreases in turbidity (range = 0.6–470 NTU) increased reproductive and moulting rates, whereas food concentration only affected the rate of reproduction, while conductivity had no effects. Survival was affected only by temperature.
4.  Daphnia lumholtzi tolerates high temperatures (27–30 °C) at which other cladocerans in Lake Texoma disappear. Therefore, D . lumholtzi may exploit resources in midsummer, when there are few potential competitors.     

Two new species of Keratella (Rotifera: Monogononta: Brachionidae) from Inner Mongolia,P.R. China

The relationship of species abundance to eight environmental variables was tested for 24 common species of crustacean zooplankton collected in the Atchafalaya River Basin during the summer of 1994. Stepwise regressions (α = 0.05) revealed significant relationships between zooplankton abundance and at least one environmental variable for 18 species ( R2 = 0.14-0.61, p < 0.0435-0.0001). The majority of these species' peak abundances were correlated with variables indicative of seasonal changes in floodplain habitat, as the Atchafalaya river receded, water temperature increased, and/or phytoplanktonic photosynthesis increased. Surface water temperature and the percent saturation of dissolved oxygen showed the most significant relationships, but specific conductance, current velocity, and Secchi disk depths were also related to abundance patterns of certain taxa. A principal components analysis of species abundances provided further insight into the partitioning of temporally-distinct zooplanton assemblages, showing that several species ( Bosmina longirostris, Daphnia parvula, Eurytemora affinis, and Ceriodaphnia quadrangula) predominated during early summer, and were supplanted by a distinct late-summer assemblage ( Diaphanosoma birgei, Moina micrura, Mesocyclops edax, and Daphnia lumholtzi) as time progressed. The transitional assemblage was dominated by Simocephalus serrulatus, Macrocyclops albidus, Microcyclops rubellus, and Thermocyclops inversus, all of which were most abundant in the hypoxic conditions characteristic of the latter stages of the Atchafalaya River flood-pulse. This revised version was published online in July 2006 with corrections to the Cover Date.     

Embryological aspects of inducible morphological defenses in Daphnia

Many cases of predator-induced morphological plasticity in daphnids are well studied examples of inducible defenses. However, little is known about the early development of these sometimes conspicuous traits. We compared for the first time in five different Daphnia species the embryonic development of predator-induced and noninduced animals using scanning electron microscopy (SEM). We observed significant morphological changes in the last embryonic stage in helmet formation in Daphnia cucullata and in neck-pedestal development in Daphnia pulex. In contrast, no morphological changes could be found during embryogenesis between induced and noninduced Daphnia lumholtzi, D. longicephala, and D. ambigua. Strategies for initiating the defensive traits differ among Daphnia species because of trade-offs between environmental requirements and developmental constraints. Some general features of Daphnia embryonic development are described using SEM. All Daphnia embryos have to shed at least three different membranes before leaving the brood pouch of the mother. After the embryos shed the third membrane, chemosensillae that are likely able to detect predator-released chemicals are exposed to the olfactory environment.     

Resource-mediated demographic variation during the midsummer succession of a cladoceran community

SUMMARY. 1. The midsummer succession of cladoceran zooplankton in Lake Texoma was examined by demographic characterization of lake populations and a parallel set of life table experiments in which the dominant cladoceran species were raised on naturally occurring lake phytoplankton at lake temperatures. Diaphanosoma leuchtenbergianum occurred throughout the study period (mid June-mid August 1981) but Daphnia parvula, D. galeata mendotae and Ceriodaphnia lacustris were rarely detected after 15 July.
2. Analyses of size structure and size-specific fecundity of the cladoceran populations showed that (1) size structure shifted to smaller individuals in mid July, (2) size-specific fecundity was relatively constant in the smaller size classes and was constant or declined in larger females of each species, and (3) size at maturation did not vary significantly during the study of any population.
3. Individual growth rates of the two Daphnia species and Ceriodaphnia in the life table experiments were significantly lower in mid July when populations were declining and when size structure of each population was more dominated by smaller Individuals. The reduction in individual growth rates was short-lived in the life table experiments; growth rates returned to pre-decline levels by the end of July.
4. Two effects of temporary reductions in individual growth rates when size-specific fecundity and size at maturation are constant are (1) that size structure is temporarily shifted to smaller individuals, and (2) the potential rate of population growth is reduced because of delays in reproduction. Because data on size structure and size-specific fecundity are commonly used to evaluate the relative effects of predators and resource changes on populations it is noteworthy that erroneous conclusions about the mechanism of population decline may be reached when individual growth rates vary.     

Range expansion and potential dispersal mechanisms of the exotic cladoceran Daphnia lumholtzi

. Recently the exotic cladoceran Daphnia lumholtzi has invadedfreshwater systems throughout the southern and midwestern UnitedStates. We conducted regional surveys of eastern Kansas reservoirsto document the range expansion of D.lumholtzi. Daphnia lumholtziwas found in five of 35 reservoirs sampled in 1994, and 11 ofthe 35 reservoirs when re-sampled in 1997. In addition, we sampled40 small ponds inaccessible to recreational boats, within thewatershed of an invaded reservoir. We did not find D.lumholtziin any of these ponds, suggesting that non-human dispersal mechanismsplay an insignificant role in the range expansion of D.lumholtzithroughout the United States. Further experimentation, however,is needed to determine if the absence of D.lumholtzi from thesesmall ponds is due to insufficient dispersal mechanisms or theinability of this invader to successfully colonize followingarrival. Daphnia lumholtzi has broad limnological tolerances.However, invaded reservoirs tend to be larger in area, havehigher Secchi disk depths, and lower total phosphorus, totalnitrogen and chlorophyll a levels relative to non-invaded reservoirs.Analyses of pre-invasion zooplankton communities indicate thatD.lumholtzi may be invading reservoirs in which native Daphniaspecies are rare. While the long-term effects of the invasionof D.lumholtzi are unknown, it has the potential to dominatelate summer zooplankton communities in eastern Kansas reservoirs.Therefore, we need to continue to survey and monitor invadedreservoirs to document the range expansion of D.lumholtzi anddetermine the long-term implications of the introduction ofthis invader.     

Zooplankton-phytoplankton interactions: the case for refining methods, measurements and models

Many of the classic experiments on the interactions between animals and their food supply were performed using laboratory cultures of Daphnia but comparable predator-prey cycles have seldom been recorded in the field. In this paper, we report the results of a field experiment designed to examine the effect of seasonal variations in the supply of edible algae on the population dynamics of Daphnia hyalina var lacustris. The experiment was performed in a large (45 m diameter) enclosure that was fertilised at regular intervals to sustain phytoplankton production. The results demonstrate that predator prey interactions of the kind recorded in culture can be detected in the field if large numbers of samples are collected and the results analysed using appropriate methods. Analyses using the numbers of Daphnia collected at a single station and chlorophyll a as a measure of food availability produced inconclusive results. Similarly, Daphnia birth rates calculated using the total number of individuals provided a poor measure of the animals reproductive response to changes in the supply of food. The clearest evidence of systematic plant-herbivore interactions was obtained when improved estimates of Daphnia numbers and adult birth rates were combined with estimates of edible rather than total algal biomass. Daphnia birth rates were then positively correlated with the estimated amount of ingestible carbon (r = 0.77) and the proportion of gravid adults decreased dramatically when the concentration of food fell below 0.10 mg C l^-1. Measurements and models of phytoplankton loss rates were then used to assess the impact of Daphnia grazing on the growth and decline of three species of edible algae. The results demonstrated that most species were able to sustain positive growth rates despite short-term increases in the population filtration rate. For much of the summer, the Daphnia appeared to behave as non-interactive grazers and had very little effect on the growth rate of their main phytoplankton food. Their grazing activity did, however, arrest the growth of a late summer population of Chlorella when the Daphnia were estimated to be filtering all the water contained in the enclosure in less than a day.     

Effects of turbidity on life history parameters of two species of Daphnia

SUMMARY. 1. Life table experiments were carried out in a range of turbidities on D. pulex , representing a 'clear-water', and on D. barbata , representing a 'turbid-water' species.
2. In contrast to predictions based on seasonal occurrence patterns, D. pulex had a higher intrinsic rate of increase ( r _c)s over most of the turbidity range than D. barbata. Consistent differences in life history between the two species were found, but life history characteristics were rather insensitive to turbidity levels. Although turbidity per se appears to have little direct effect on life history parameters, it could influence the seasonal succession of these two species by interacting with other factors.
3. It is suggested that visual predation by fish rather than turbidity per se probably influenced the seasonality of Daphnia species. During the warmer months, selective removal of D. pulex by visually foraging fishes may allow D. barbata to gain dominance. Effects of temperature and nutrition also merit further study.     

Larval fish-induced phenotypic plasticity of coexisting Daphnia: an enclosure experiment

1. The indirect effects of predators on lower trophic levels have been studied without much attention to phenotypically plastic traits of key food web components. Phenotypic plasticity among species creates phenotypic diversity over a changing environmental landscape. 2. We measured the indirect effects of planktivorous larval walleye (Stizostedion vitreum) on phytoplankton biomass through their effects on the dominant herbivore species, Daphnia pulicaria and D. mendotae. 3. Fish had no effect on phytoplankton biomass or overall Daphnia density. We observed a compensatory response to predation by functionally comparable species within a trophic level in the form of shifting dominance and coexistence of Daphnia species. We hypothesized that this phenotypically plastic response to predation decoupled a potential trophic cascade in this freshwater pelagic system. Daphnia pulicaria density decreased over time with fish predation, but D. mendotae density increased over time with fish predation. 4. Phenotypically plastic life history trait shifts and reproductive rates differed between species in fishless and fish enclosures, accounting for population trends. Daphnia pulicaria were also proportionally higher in walleye larvae stomachs than in the enclosures, indicating that walleye preferred to feed on D. pulcaria over D. mendotae. The resultant shift in dominance may partially explain the overall benign effect of fish on grazers and supports the hypothesis that trophic level diversity can decouple a trophic cascade.     

The biogeography of AustralianDaphnia: clues of an ancient (>70 m.y.) origin for the genus

The AustralianDaphnia (D. occidentalis, D. jollyi, D. lumholtzi, D. carinata, D. nivalis, andD. cephalata) include representatives from three distinct groups within the genus. The worldwide distribution of species within these three groups is consistent with a Gondwanan or pre-Gondwanan origin for each group. These data suggest an even more ancient origin for the genusDaphnia. The finding thatD. cephalata must have been separated fromD. carinata since Gondwanan times is consistent with recent biochemical data on the phylogenetic relations of AustralianDaphnia, but contrasts with electrophoretic evidence of continuing hybridization between these two taxa. It is suggested that continued gene exchange between otherwise discrete species, or clonal complexes, ofDaphnia may be maintained for millions of years, partly as a result of the varying degrees of cyclical parthenogenesis, and partly as a result of the highly fragmented population structures, found in these taxa.     

Relative vulnerabilities of natural rotifer and ciliate communities to cladocerans: laboratory and field experiments

SUMMARY. 1. Two experiments with plankton communities from Storrs Pond (NH), one conducted in the laboratory and one in field enclosures, assessed the impact of different cladocerans on rotifers and ciliated protozoa.
2. The smallest cladoceran, Bosmina longirostris , did not depress rotifer or ciliate growth rates while the intermediate sized dadoceran, Daphnia galeata mendotae , reduced ciliate growth rates in the enclosure experiment but had only a marginal effect in the jar experiment. D. galeata mendotae had no effect on any of the rotifers in either experiment.
3. In both experiments the largest cladoceran, Daphniapulex , depressed the growth rates of ciliates and those rotifers known to be vulnerable to interference competition. Polyarthra vulgaris , previously shown to be resistant to cladoceran interference, was the only rotifer unaffected by D. pulex in the field experiment but was depressed by the much higher densities of this cladoceran in the laboratory experiment.
4. Cladocerans did not affect phytoplankton or bacterioplankton abundance in either experiment. Therefore the mechanism most likely to be responsible for the suppressive effect of cladocerans on rotifers and ciliates in these experiments is direct mechanical interference or predation, rather than exploitative competition.     

Linking biotic interactions and climate change to the success of exotic Daphnia lumholtzi

1. Warmer temperatures may increase cyanobacterial blooms in freshwater ecosystems, yet few ecological studies examine how increases in temperature and cyanobacterial blooms will alter the performance of non‐native species. We evaluated how competitive interactions and interactions between these two drivers of ecological change influence the performance of non‐native species using the native zooplankton Daphnia pulex and the non‐native zooplankton Daphnia lumholtzi as a model system. Based on the literature, we hypothesised that D. lumholtzi would perform best in warmer temperatures and in the presence of cyanobacteria. 2. Laboratory competition experiments showed that in the absence of competitors, growth rates of D. pulex (but not D. lumholtzi) were reduced at higher temperatures and with the cyanobacterial foods Anabaena flos‐aquae and Microcystis aeruginosa. In the presence of competitors, however, D. pulex emerged as the superior resource competitor at both temperatures with cyanobacterial food. We therefore predicted that, if competitive interactions are important to its establishment, D. lumholtzi would perform best in the absence of cyanobacteria in heated environments. 3. As predicted, when both species were introduced at low densities in field mesocosms, D. lumholtzi performed best at high temperatures without added cyanobacteria and worst at ambient temperatures with added cyanobacteria, indicating that competitive interactions are likely to be important for its establishment. 4. Taken together, these studies indicate that, while D. lumholtzi may benefit from increases in temperature, associated increased cyanobacterial blooms may hinder its performance. Thus, our findings underscore the importance of considering biotic interactions such as competition when predicting the future establishment of non‐native species in response to climate warming.     

Salamander predation and vertical distributions of zooplankton

SUMMARY. 1. The impact of different predation regimes of zooplanktivorous larval tiger salamanders, Ambystoma tigrinum nebulosum , on the vertical temporal distributions of Daphnia pulex, Diaptomus nudus and larval Chaoborus flavicans was studied in limnetic enclosures in a pond in east-central Arizona. Zooplankton densities were monitored 1-2 h before sunset, at sunset, and 1-2 h after sunset at three depths (0-0.4, 1.0 and 2.0 m) in four salamander treatments (0, 20, 40 salamanders per enclosure, and unenclosed limnetic areas).
2. Densities and body size of Daphnia pulex were consistently greater in salamander-free enclosures than in other treatments. More Daphnia were generally nearer the surface (0-1.0 m) than along the bottom before and at sunset than after sunset in enclosures without salamanders. After sunset, Daphnia was by far most dense along the bottom. Densities were uniform among depths and sample times in treatments with salamanders. Salamanders moved throughout the water column. Frequencies of D. pulex spined morphs (with prominent dorsal crest) did not differ among salamander treatments.
3. Chaoborus densities were positively correlated with D. pulex distributions among depths and sample times in treatments. Spatio-temporal distributions of Chaoborus were influenced more by their daphnid prey than by risk of predation from larval salamanders.
4. Densities of Diaptomus nudus were highest in open water controls and lowest in enclosures without salamanders. Diaptomus was generally more dense between 0 and 1 m than at 2.0 m depths in all treatments, and vertical distributions were not dependent on salamander density.     

Predictors of invasion success by Daphnia species: influence of food, temperature and species identity

Non-indigenous North American Daphnia ‘pulex’ has recently invaded lakes and reservoirs in South Island, New Zealand, that formerly contained only native Daphnia carinata. New Zealand is characterised by a wide range of freshwater ecosystems and low species diversity of planktonic crustaceans, particularly Cladocera. The potential success of a species to invade and establish in a new community is likely to be predicted more accurately when reproductive response norms and fitness of key resident species, as well as the non-indigenous species, have been established under a range of relevant environmental conditions. Based on the results of experiments to test aspects of reproduction and fitness of the invader, D. ‘pulex’, and D. carinata when grown together at a range of relevant temperatures (8–23 °C) and related photoperiods, I predict the species-specific potential of D. ‘pulex’ to be dispersed and colonise New Zealand lentic habitats, and the potential of D. carinata to persist with the invader in these habitats. Larger population densities of D. ‘pulex’ compared to D. carinata at higher temperatures and food level, and larger densities of D. carinata at low temperatures, imply a potential for both species to coexist in New Zealand lakes, facilitated by seasonal succession; increased water temperature and nutrient input associated with climate and land use changes appear likely to promote the wider establishment of D. ‘pulex’, with both negative and positive implications for the conservation and management of New Zealand’s freshwater ecosystems.     

Changes in food quality of the green alga Scenedesmus induced by Daphnia infochemicals: biochemical composition and morphology

1. The effects of Daphnia infochemicals on the morphology and biochemical composition of Scenedesmus were studied and subsequently their influence on Daphnia life history. Three species of Scenedesmus ( S. acutus, S. obliquus and S. subspicatus ) were tested for Daphnia -induced colony formation. Life history experiments were performed with Daphnia cucullata and D. magna feeding on unicellular or colonial S. acutus.
2. Colony formation was promoted when S. acutus and S. obliquus were exposed to filtered water from a D. magna culture. S. subspicatus did not form colonies when exposed to culture water, but showed some colony formation when exposed to live D. magna.
3. No clear differences were found in total lipid, protein or carbohydrate content between the unicellular and colonial ecomorphs of the three Scenedesmus species. However, fatty acid (FA) concentration and composition were changed. Total FA as a percentage of frozen dry weight (DW) were increased in colonies. The ratio of monounsaturated (MUFA) and polyunsaturated fatty acids (PUFA) and the ratio of ω3 : ω6 fatty acids decreased in colonies.
4. A short-term grazing experiment with three size classes of Scenedesmus revealed lower clearance rates for small Daphnia when feeding on large colonial Scenedesmus.
5. Life history parameters of both Daphnia species were influenced by food type. The intrinsic rate of population increase ( r) was significantly lower for daphnids feeding on colonies.
6. The negative influence of colonial Scenedesmus on population growth of Daphnia seems to be determined by algal morphology rather than biochemical composition.