Diel vertical migrations of invertebrate predators (Leptodora kindtii ,Thermocyclops taihokuensis, andMesocyclops sp .) in a shallow, eutrophic lake

We studied vertical distribution patterns of three invertebrate predators – Leptodora kindtii, Mesocyclopssp., and Thermocyclops taihokuensis– in a shallow eutrophic lake, Lake Suwa , Japan. From June to October in 2000 and 2001, we collected samples in the lake center in order to examine the vertical distribution patterns and the densities of the predators in the water column during the day (0900) and at night (2330). We also examined phototactic behavior of Leptodora in the laboratory. The three invertebrate predators showed clear migration patterns. Leptodora and Thermocyclops displayed a typical migration, avoiding the surface and maintaining a high abundance in deeper water during the day, and being distributed uniformly during the night. Mesocyclops, on the other hand, showed no clear vertical distribution pattern in the water column. However, Mesocyclops showed higher densities in the water column during the night than during the day. It suggests that they stayed just above the bottom during the day and migrated upward during the night. Leptodora also showed such a density difference between day and night. In the laboratory, Leptodora showed strong negative phototactic behavior. The observed density changes between day and night in Leptodora and Mesocyclops suggests the possible underestimation of their population density by usual sampling methods, and thus the impact of predation on populations of prey zooplankton species may also be underestimated in shallow water bodies.     

Leptodora kindti: efficient predator and preferred prey item in Neusiedler See,Austria

In the Neusiedler See, Leptodora kindti appeared in the plankton in April/May; the growing season lasted until November. Its optimum conditions prevailed during the summer months, when densities of >500 ind. m^–3 were recorded. First, males appeared by the mid/end of August, while maximum resting egg production occurred in September. At temperatures below 10 °C the population declined.At high population levels, L. kindti caused suppression of its preferred prey item, the juveniles of Diaphanosoma mongolianum. Twenty-five to 45% of juvenile Diaphanosoma can be eliminated by Leptodora. This predation impact is a pronounced but short-term event. During the periods of high Leptodora densities, a significant relationship between Leptodora predation and the death rate of Diaphanosoma is found.The older developmental instars of L. kindti are themselves endangered by the impact of planktivorous fish (i.e. Pelecus cultratus, Alburnus alburnus), which show a preference for Leptodora larger than 6 mm. Frequently, elimination by fish corresponds with high death rates of Leptodora. During the periods when adult Leptodora occurred in numbers >50 ind. m^–3, 55% of the variance connected with its death rate can be attributed to fish predation. The success of this population of L. kindti is discussed within the framework of predator-prey interactions.     

Food and feeding behaviour of a planktivorous cyprinid, Pelecus cultratus (L.), in a shallow eutrophic lake, Neusiedler See (Austria)

Pelecus cultratus (razor fish), a cyprinid fish, has become a dominant species in Neusiedler See. Gut content analyses of 400 specimens collected in 1989 and 1990 showed that Pelecus fed mainly on large zooplankton (Diaphanosoma, Leptodora and Arctodiaptomus), although their diet also included Insecta (larvae, pupae and adults) and Arachnida (spiders), occasionally small fishes. Comparison of the relative abundance of the zooplankton species in the stomach to the lake indicated that Pelecus strongly selected cladocerans over copepods, and fed mostly on large-sized individuals of Diaphanosoma, Leptodora and Arctodiaptomus. The fish showed a significant positive selectivity only for individuals of Diaphanosoma > 1.0 mm and Leptodora > 4.0 mm. In contrast, selectivity increased continuously in relation to the diameter of the compound eye of both prey species. This suggested that prey visibility was a key factor in determining the prey selectivity by Pelecus. It also seems likely that the persistence of the Leptodora population in Neusiedler See can be attributed to negligible predation pressure on the smaller sized individuals of this species.     

Effect of temperature on development and growth of the raptorial cladoceran Leptodora kindtii under laboratory conditions

1. Leptodora is a key species in many temperate freshwater systems, but so far its role in the food web could not be properly evaluated because detailed information about its secondary production was lacking. As we wanted to estimate the secondary production of Leptodora, we measured its development and growth rates in the laboratory. 2. Employing improved methods to estimate growth and instar durations, we cultured Leptodora kindtii in the laboratory at four constant temperatures (15, 17.5, 20 and 25 °C). Growth in length and development times of eggs and instar stages were assessed. 3. Growth rates at 15, 17.5 and 20 °C were similar, but at 25 °C growth was distinctly faster. At 17.5 °C we observed seven juvenile instar stages before the first adult instar stage was reached.     

Leptodora kindti and Flexible Foraging Behaviour of Fish – Factors behind the Delayed Biomass Peak of Cladocerans in Lake Hiidenvesi

In the eutrophic L. Hiidenvesi, the spring biomass maximum of cladoceran zooplankton is missing and the highest biomass takes place in July–August. The factors behind the delayed biomass peak were studied in four different basins of the lake with concomitant data on cladocerans assemblages, density of the predatory cladoceran Leptodora kindti and food composition of fish. In all the basins, the abundance of Leptodora peaked in June, being highest (up to 800 ind. m^–3) in the two most shallow basins (max depth < 4 m). The duration of the high population density was short and in July‐August Leptodora density stayed below 200 ind. m^–3, although the water temperature was still favourable. The collapse of the Leptodora population coincided with the change in the feeding habits of fish. In early summer, fish predation was targeted mainly on copepods and zoobenthos, while in high summer Leptodora was one of the main preys of perch, white bream and bleak. The biomass of herbivorous cladocerans was below 10 μg C l^–1 in June, and climbed to a maximum in August in the two most shallow basins (34 and 76 μg C l^–1), in July in the deepest basin (27 μg C l^–1), and in September in the intermediate basin (55 μg C l^–1). In the two most shallow basins, the death rate of the dominating cladoceran, Daphnia cristata, closely followed the food consumption rate by the Leptodora population. In the deeper basins, the agreement was not so close, smelts (Osmerus eperlanus) and chaoborids being important predators of herbivores. The duration of the period of high Leptodora density thus depended on the predation pressure by fish, while the increased fish predation on Leptodora in July–August allowed the elevation of the biomass of herbivorous cladocerans.     

Effects of <Emphasis Type="Italic">Bythotrephes longimanus</Emphasis> (Crustacea,Cladocera) on the abundance,morphology, and prey community of <Emphasis Type="Italic">Leptodora kindtii</Emphasis> (Crustacea,Cladocera)

We hypothesized that native Leptodora kindtii would be shorter and have smaller feeding baskets in central Ontario lakes with greater abundances of small-bodied zooplankton prey, and that differences in zooplankton size among lakes could be attributed to the invasive cladoceran Bythotrephes longimanus. We evaluated these conjectures by comparing size metrics of Leptodora and the size of their preferred cladoceran prey in lakes invaded or not by Bythotrephes. Leptodora was less abundant in invaded lakes, but were smaller bodied with smaller feeding baskets only in lakes with long invasion histories. Small cladoceran abundance was greater in non-invaded lakes and was directly related to Leptodora abundance although not to Leptodora size. Mean Leptodora body size declined with increasing abundance of Bythotrephes. We evaluated three possible explanations for these patterns in Leptodora—(a) competition with Bythotrephes for zooplankton prey, (b) direct predation by Bythotrephes, and (c) size-selective predation by fish. While we were unable to unequivocally distinguish among these hypotheses, our observations are most consistent with predation by Bythotrephes changing zooplankton community composition and size structure in a manner that is detrimental to Leptodora. Our results indicate that Bythotrephes invasion may trigger more complex and subtle changes in food webs than previously thought.     

Shifting invertebrate zooplanktivores: watershed-level replacement of the native <Emphasis Type="Italic">Leptodora</Emphasis> by the non-indigenous <Emphasis Type="Italic">Bythotrephes</Emphasis> in Canadian Shield lakes

The abundance of the native, pelagic macroinvertebrate predator, Leptodora kindtii, is negatively correlated with the abundance of a new invasive competitor, Bythotrephes longimanus, in a small number of Canadian Shield lakes. However, we do not yet know if Bythotrephes is replacing Leptodora on a regional scale. We determined the distribution of both species in 166 lakes in the District of Muskoka, south-central Ontario, Canada—the watershed with the longest history and largest prevalence of Bythotrephes invasions in North America. The frequency of occurrence of Leptodora was substantially reduced (twofold) in the presence of Bythotrephes. We argue that Bythotrephes is responsible for this dramatic reduction in the frequency of occurrence of Leptodora. Lakes in which both species co-occurred could not be distinguished from invaded lakes without Leptodora, suggesting a pattern of species replacement at a watershed level. We believe this is the first account of the widespread replacement of a native, pelagic macroinvertebrate predator by Bythotrephes in North America, and it does not bode well for Leptodora given the rapid, ongoing spread of Bythotrephes.     

Conochilus in Lake Washington

The rotifer Conochilus unicornis appeared in Lake Washington sporadically and usually in small numbers during a total of 28 years of observation since 1933. Conochilus hippocrepis was present even less frequently until the 3 year period 1977–1979, when it became extraordinarily abundant. The abundances of food organisms and known predators have been examined.The only consistent correlation found was between the seasonal maximum of Leptodora and the decrease of C. hippocrepis from its peak abundance each year. Leptodora was generally less abundant during 1977–1979 than during most of the preceding years and the years following the C. hippocrepis episode. No such correlations were found with Epischura, Cyclops or edible algae.In July 1985, there was a small resurgence of C. hippocrepis preceded by a small population of C. unicornis. Leptodora was less abundant than in the immediately preceding years. Observations were made on stomach contents of both animals and colony size of C. hippocrepis. Recommendations are made for programs of new field studies and laboratory experiments to establish the actual requirements for successful populations of Conochilus.     

Predation impact of Leptodora kindtii on population dynamics and morphology of Bosmina fatalis and B. longirostris in mesocosms

1. We assessed the impact of predation by the invertebrate predator Leptodora kindtii on Bosmina longirostris and B. fatalis, which show seasonal and reciprocal succession patterns in Lake Suwa, in a mesocosm experiment using 20‐L tanks with or without the predator under different food conditions. We also analysed morphological responses of the two Bosmina species to the predator in the tanks. 2. Bosmina fatalis dominated B. longirostris regardless of predator presence under high food density. However, the presence of Leptodora induced the dominance of B. fatalis more rapidly than its absence. On the contrary, no dominance of B. fatalis was observed in tanks with low food density, irrespective of the presence of the predator. Only B. fatalis showed morphological changes in response to the presence of Leptodora. 3. Mucrone length and antennule shape (angle between body and antennule and angle between antennules) showed marked responses at both high and low food densities, but antennule length responded only at high food density. Mucrone length seems to be a more effective defence against Leptodora. 4. The results suggest that B. fatalis is a superior competitor against B. longirostris and is more resistant to Leptodora predation, especially in good food conditions. The repeatedly observed seasonal succession of the two Bosmina species in the eutrophic Lake Suwa – the replacement of B. longirostris by B. fatalis following the occurrence of abundant Leptodora– seems to be caused by the selective predation of Leptodora on B. longirostris as well as the competitive ability of B. fatalis.     

The ecological niches of <Emphasis Type="Italic">Bythotrephes</Emphasis> and <Emphasis Type="Italic">Leptodora:</Emphasis> lessons for predicting long-term effects of invasion

We here exploit two large datasets on zooplankton in Norwegian lakes, spanning a wide range of geographical, physical, chemical and biological properties, to assess the ecological niches and habitats of Bythotrephes longimanus and Leptodora kindtii. The species overlapped geographically, yet co-occurred only in a limited number of lakes. Bythotrephes inhabited virtually all types of lakes, except alpine localities and productive lakes dominated by cyprinid communities where the hyaline Leptodora was most abundant. The zooplankton communities also differed in Bythotrephes and Leptodora lakes, probably both reflecting different predatory regimes, but also water quality and other lake-specific properties. We found no evidence for species being excluded by the presence of Bythotrephes, rather the diversity in general was higher in lakes with these predators present compared with those without. We found, however, a very close association between Bythotrephes and Daphnia galeata and to some extent also between Bythotrephes and D. longispina, suggesting that these species also may benefit from Bythotrephes invasion. Both Bythotrephes and Leptodora species occur naturally in this region, and knowledge about the ecological preferences and the zooplankton community composition in Bythotrephes—and Leptodora lakes will provide valuable information about the long-term effects of Bythotrephes invasion and potential interaction with of Leptodora as top invertebrate predator.     

Investigations on fecundity of Bythotrephes longimanus in Lake Lucerne (Switzerland) and on Niche Segregation of Leptodora kindti and Bythotrephes longimanus in Swiss lakes

In Lake Lucerne, Switzerland, the predaceous cladocerans Leptodora kindti and Bythotrephes longimanus segregate along spatial and temporal dimensions. In spring (April–May/June), Bythotrephes longimanus occurs below 0–20 m, while Leptodora is absent. In summer and early autumn (July–September/October), when Leptodora dominates during daytime in the 0–20 m depth, Bythotrephes longimanus also lives in deeper zones. Food competition and fish predation pressure may be the cause of differences in ecology of Leptodora and Bythotrephes acquired during evolution. Due to its transparency and tolerance of higher temperature, Leptodora could avoid fish predation and, therefore, competes with Bythotrephes longimanus successfully. In addition, the differences between the two species may account for the spatial and temporal niche segregation in oligotrophic Swiss Lakes. But spatial niche segregation is less important in mesotrophic lakes with high prey density than in oligotrophic lakes with low prey density. In small, eutrophic lakes importance of temporal niche segregation also decreases, and Bythotrephes is seldom or not present. The preference of Bythotrephes to live in deeper water to avoid fish predation during summer may be the cause of its difficulties to establish itself in small and eutrophic lakes with high prey densities, where the hypolimnion is missing or anoxic.In the spring, Bythotrephes exhibits r-strategy (smaller body size and a higher fecundity), the female is already fertile after the first molt. In the summer, a K-strategy prevails (larger body length and lower fecundity than in the spring), and female Bythotrephes are fertile only after the second molt. Shortage of prey (biomass of Bosmina and Daphniadecreased after June especially in the surface layers) and the maximum fish predation pressure in summer may change the life strategy of Bythotrephes: while fecundity decreases from generation to generation, body length increases. Enhanced prey densities (e.g. during mesotrophic conditions in L. Lucerne) lead to larger individuals in summer and autumn.     

Leptodora kindtii survival in the laboratory

Leptodora kindtii, a pelagic predatory cladoceran, suffers high mortality on transfer to laboratory, which makes the experimental work difficult. We investigated the causes of high mortality, using four variables: water volume, animal density, light intensity, and origin of water for culturing, i.e., water from native or a non-native lake. For the experiments we used Leptodora and water from Lake Loosdrecht and Lake Maarsseveen (The Netherlands). Water was found to be the most important factor; the animals did not necessarily do better in lake water from which they were collected. Water volume and animal density were of limited importance, and light intensity did not affect survival.     

Contrasting life histories of the predatory cladocerans Leptodora kindtii and Bythotrephes longimanus

The predatory cladocerans, Leptodora kindtii (Focke, 1844) andBythotrephes longimanus (Leydig, 1860), express markedly differentlife-history traits. Leptodora produce small-bodied neonatesthat mature at small sizes but continue to grow throughout life.Bythotrephes produce larger neonates in both relative and absoluteterms that grow rapidly to a large size at maturity whereuponthey curtail somatic growth and divert resources mainly to reproduction.Despite their remarkable differences, the sets of life-historytraits of both species appear to be solutions to the same basicselection pressures imposed by visually discriminating gape-limitedfishes and foraging constraints imposed by prey size. Leptodorastresses pre-contact (transparency) while Bythotrephes stressespostcontact (caudal spine) modes of morphological defense againstfishes. Mounting these disparate modes of defense has consequencesfor selection on timing and allocation to body growth that mayunderlie competitive imbalance between the species. Owing tothe production of large-bodied neonates that grow rapidly, Bythotrephesquickly attain body sizes that both admit them to a broaderprey base in size and taxonomic variety, and allow shorter preyhandling times, in comparison to Leptodora. This provides Bythotrepheswith a wider and more exploitable prey base from an earlierage and may explain why Leptodora has declined in density followingBythotrephes invasion into some North American lakes. The divergentsets of life-history traits expressed by Leptodora and Bythotrephesparallel two dominant life-history strategies evolved by phytoplanktivorousspecies of the order Cladocera.     

The development of Hexarthra spp. in a shallow alkaline lake

In Neusiedler See, a shallow alkaline lake with fluctuating water level and salinity, four species of Hexarthra occur: H. mira, H. fennica, H. jenkinae (occasional) and H. polyodonta. The analysis of longterm data reveals a general phenological pattern which does not change from year to year. They first occur in May, develop a maximum in June/July, sometimes a second one in August/September and disappear in October. But the species succession is different in the various years, occasionally only one species (H. mira or H. polyodonta) being present. There is a fairly consistent relation between the chemical conditions and the prevalent species; an increase in salinity favours the development of H. polyodonta. Low temperature and wind generated suspended particles have a negative influence on the development of the Hexarthra populations. Smaller populations of Hexarthra are in a relation to the occurrence of Leptodora indicating predation pressure of the latter species. In Neusiedler See the Hexarthra populations seem to be controlled to a great extent by abiotic factors, but predation by Leptodora and most probably by young fish seems to play an important role too.     

Inhibition of development of anti-predator morphology in the small cladoceran Bosmina by an insecticide: impact of an anthropogenic chemical on prey–predator interactions

1. The combined effect of the insecticide carbaryl and the predator (Leptodora kindtii) kairomone was assessed on the development of protuberant morphology in the small cladoceran Bosmina fatalis, a feature which evolved originally as a response to the kairomone. The experiment showed that Bosmina changed its morphology in response to the kairomone, but development was inhibited by carbaryl at a sub‐lethal concentration even in the presence of the kairomone. At the same time, reduction of fecundity was observed in animals exposed to carbaryl. 2. A short‐term feeding experiment (B. fatalis versus Leptodora) indicated that such low concentrations of the insecticide had no impact on predation by Leptodora. Thus, it is suggested that the inhibition of development of anti‐predatory defences in Bosmina can increase its vulnerability to predation. 3. Such disturbance of chemical communication by the insecticide reduces the individual survival rate of prey Bosmina in the environment with high predator density. On the other hand, reduction of fecundity may result in decreased population growth rate of animals. 4. The impact of the insecticide on the anti‐predator morphology in Bosmina (inhibition) was opposite to that in Daphnia (enhancement). This suggests that biochemical induction processes in the development of anti‐predatory morphology are evolutionally different between Daphnia and Bosmina.     

The Influence of Invertebrate Predators on Daphnia Spatial Distribution and Survival in Laboratory Experiments: Support for Daphnia Horizontal Migration in Shallow Lakes

The behavioural response of Daphnia cucullata to the presence of the pelagic invertebrate predator Leptodora kindtii, and the predation rate of littoral dragonfly nymphs on this species were investigated under laboratory conditions. Results of this study revealed a strong hiding response of Daphnia cucullata in the presence of the predatory cladoceran, L. kindtii, which was similar to the response of Daphnia in the presence of juvenile perch. This suggests that pelagic invertebrate predators may cause Daphnia to hide in the littoral zone which could result in increased exposure to predation by littoral invertebrates. A strong influence of dragonfly nymphs on D. cucullata, both in the presence and absence of macrophytes, was found. The average predation rate of Odonata larvae was about 5 prey ind^–1 h^–1 and did not differ significantly between treatments. Quantification of dragonfly pressure on Daphnia populations will require cross‐verification with field experiments since in the natural conditions Daphnia seeks a shelter in the vegetation stands against predation by Leptodora, despite the occurrence of odonates. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Long-term changes in the community of planktonic crustaceans in Lake Glubokoe in relation to predation and competition

Studies on the planktonic Crustacea in Lake Glubokoe (Moscow region) from 1897 to 1982 have been reviewed. After 1965, when most of the swamp water inflow was diverted from the lake, certain populations have penetrated into deeper layers, while the average overlap in the community has dropped. The summer dominant Mesocyclops leuckarti disappeared and Chydorus sphaericus appeared in mass quantities; this development was not associated with biomass changes of blue-greens. The Cladocera/Calanoida ratio and the relative abundance of certain species showed no considerable changes. The effect of Chaoborus, Cyclops and Leptodora in the 70's on the crustacean populations was insignificant in spite of the increasing abundance of these genera after 1965. Analysis of the dynamics of the clutch size and the density of lake populations in the 1970's and 1980's revealed food limitation and competition in herbivorous Cladocera. In laboratory experiments with lake water they always competed, though the intensity of their interaction varied depending on the species combinations.     

Biogeography and evolution of the Holarctic zooplankton genus Leptodora (Crustacea: Branchiopoda: Haplopoda)

Aim To reconstruct the phylogeographic history of the Holarctic carnivorous genus Leptodora (Crustacea: Cladocera: Haplopoda). Location We studied the DNA of between one and five specimens each from 28 populations distributed across the Holarctic, but with emphasis on Eurasia. Methods We sequenced a mitochondrial (cytochrome c oxidase subunit I) and a nuclear (elongation factor‐1α) gene, and combined this molecular information with geological and palaeoclimatological data. Haplotype networks and phylogenetic trees were constructed using a Bayesian and maximum likelihood approach. A molecular clock was applied. Results Leptodora consists of three clades (Leptodora kindtii in Europe, Leptodora richardi in China and Japan, and Leptodora sp. in North America), with insular subclades in Japan and in the eastern Mediterranean. The North American clade was not studied in detail. Leptodora richardi is the more thermophilic of the three. It extends from the Tropic of Cancer in the south to the Heilong Basin in the north. The western European L. kindtii is more cold‐water adapted than the eastern Mediterranean subclade. ‘West European’ and ‘Chinese’ clades are broadly separated by a hybrid zone in Siberia and European Russia as far west as the Volga. These hybrids have the mitochondrial DNA of L. kindtii, the nuclear DNA of L. richardi and the low‐temperature preference of L. kindtii, and may have formed as recently as the Holocene hypsithermal. A pure L. kindtii population in the Upper Irtysh catchment, east of the Dzungarian Gates, has been sequestered in endorheic Lake Wulungu, Xinjiang, since the mid‐Pleistocene. Main conclusions Application of a molecular clock places the most recent common ancestor of the North American, East Asian and European populations in the mid‐Miocene. The North American taxon is still living in isolation, while the Eurasian taxa, separated by the Alpine folding, made contact again in the Pleistocene, when the cold‐stenothermic L. kindtii repeatedly moved eastwards across Siberia and back. The population in Xinjiang is a relict of an early wave coming from western Europe: it crossed the Dzungarian Gates during a humid mid‐Pleistocene event, probably corresponding to the Apsheron transgression in the Caspian Basin. Later aridity isolated it there, and it started accumulating private haplotypes. The Holocene Euro‐Siberian hybrid zone may eventually engulf all European populations.     

The transfer of fatty acids in a freshwater planktonic foodweb of the Kuibyshevskoe reservoir (middle reaches of the Volga)

The fatty acid composition of seston (small-size fraction, < 50 m) and of Daphnia galeata, Bythotrephes longimanusand adult individuals of Leptodora kindtii was analysed in the summer of 1997. When comparing the eicosapentaenoic acid (EPA) content of seston with Daphnia, Daphnia with Bythotrephes; Daphniawith Leptodora, we found similarities in the dynamics of EPA accumulation, in most cases. The content of EPA in seston was found to be significantly correlated with numerical abundance of small diatoms (r =0.662).Maximally the % EPA increased from seston (traces – 4% of total fatty acids) to Daphnia (traces – 12.2%). Both the lower and upper relative EPA contents were higher in Bythotrephes (10.8–16%), whereas the maximum value was lower in Leptodora (0.4–6.3%) compared to Daphnia. Correlation coefficients between the EPA content of the organisms at different trophic levels were not significant. The existence of species-specific differences in accumulation and /or transformation of polyunsaturated fatty acids in freshwater crustaceans is proposed.     

Foraging behavior of the predaceous cladoceran, Leptodora kindti, and escape responses of their prey

Using silhouette video photography we have made the first quantitativeobservations of foraging behavior in Leptodora kindti, a predaceouscladoceran (Haplopoda). Leptodora swims with a mean velocityof 13.4?4.0 mm s^–1 and initiates an attack only upon directcontact with potential prey. The attack sequence is as follows:Leptodora swims randomly through the water column with all fivepairs of thorac appendages spread to form a ‘feeding basket’and, seemingly by chance, encounters prey. Shortly after preymake contact with any part of Leptodora's body (usually ventral),the abdomen is rapidly pulled forward, clamping itself underthe feeding basket so that the telson closes it at the posteriorend. The duration of this movement is always the same and weconclude that it is an indiscriminate reflex. If the prey isencountered anywhere but a short distance directly in frontand slightly below the Leptodora, it is not captured. The speedof copepod escape responses effectively allows them to avoidcontact with the predator. Daphnia's escape response, particularlythat of juveniles, is slower and leaves them far more susceptibleto Leptodora predation.