Taxon-specific reaction norms to predator cues in a hybrid Daphnia complex

1. Previous studies have shown that interspecific hybridisation is common among taxa from the Daphnia galeata/hyalina/cucullata species complex. We investigated the influence of predator kairomones on the morphology and life histories of nine clones belonging to three taxa (pure D. galeata, F₁ hybrids between D. galeata and D. hyalina, and backcrossed D. hyalina) of this species complex. Predators exerting positive (fish) and negative (Chaoborus larvae) size‐selective predation were tested. 2. The most responsive traits were size at maturity and size of neonates. Despite large between‐clone variation, discriminant analysis revealed that the three taxa were distinct from each other in key life‐history traits. F₁ hybrids did not react in an intermediate way compared to the other taxa: the multivariate distances between F₁ hybrids and either taxon were larger than between pure D. galeata and backcrossed D. hyalina. 3. The average plasticity (calculated across all traits) was similar for all three taxa. With regard to the size at maturity and neonate body size, the strength of the response was a function of the intrinsic values of these traits expressed in the control. For example, for size at maturity, smaller individuals showed a significantly stronger reaction to Chaoborus kairomones than larger ones. 4. Finally, we monitored seasonal changes in body size, egg number and population density of pure D. galeata and F₁ hybrids in Greifensee (Switzerland). The two taxa experienced similar seasonal changes in body size but, on some sampling dates, they differed in mean egg number. The observed seasonal changes in Daphnia body size were consistent with what would be expected if the predator assemblage shifted from fish to Chaoborus over the course of the summer. The fluctuations in the frequencies of Daphnia taxa, however, were not related to seasonal variation in Daphnia body size. 5. Experimental data suggest that temporally heterogeneous predation regimes might be an important condition stabilising the co‐occurrence of Daphnia hybrids with parental taxa. Predation regimes, however, cannot solely explain dynamic changes in taxon frequency in Greifensee.     

Daphnia hybridization along ecological gradients in pelagic environments: the potential for the presence of hybrid zones in plankton

The relative homogeneity of pelagic environments has been regarded as the reason for the absence of hybrid zones for hybridizing planktonic Daphnia (Crustacea: Cladocera); occasional dominance of interspecific hybrids over parental species was explained by their temporal superiority in fluctuating environments. However, water bodies with spatially varying environmental conditions might facilitate the formation of hybrid zones in plankton. We studied the distribution of species and hybrids of the Daphnia longispina complex in 11 canyon-shaped reservoirs, localities characterized by horizontal environmental gradients (particularly of food supply and size-selective predation); we also analysed patterns of carapace size and fecundity among coexisting taxa. Spatial distribution of taxa agreed with their ecological characteristics; those showing different affinities along longitudinal reservoir profiles differed in size according to the presumed fish predation gradient. Only hybrids of Daphnia galeata with Daphnia cucullata and D. longispina (=hyalina) were recorded. The latter two species preferred opposite ends of gradients, such spatial segregation probably explaining the absence of their hybrids. Distributional patterns were relatively stable in two consecutive summers, apart from a substantial decline of D. galeata X cucullata in the second year. The observed pattern of a hybrid-dominated zone in intermediate conditions suggests that local Daphnia hybrid zones may indeed form within reservoirs.     

Contrasting patterns of body size for Daphnia species that segregate by habitat

Summary We investigate how body size of two coexisting Daphnia species varies among 7 lakes that represent a gradient of predation risk. The two species segregate vertically in stratified lakes; D. galeata mendotae is typically smaller and more eplimnetic than D. pulicaria. The extent of vertical habitat partitioning, however, varies seasonally within and among lakes in apparent response to predation intensity by epilimnetic planktivorous fishes. Daphnia pulicaria uses the epilimnion at low levels of fish predation but is restricted to the hypolimnion under high fish predation, whereas D. galaeta mendotae always utilizes the epilimnion. The species display contrasting patterns of genetic variation in neonate size and size at maturity. D. pulicaria is larger in lakes with higher fish and Chaoborus densities whereas D. galeata mendotae is smaller. This contrast in body size in lakes with high predation is associated with greater habitat segregation in those lakes. In lakes with low predation risk, the two species are similar in body size at birth and maturity.Authorship order alphabetical     

Differential life history responses of several pelagic Daphnia clones differing in migratory behaviour

Females of the hybrid Daphnia galeata × hyalina were isolated from a natural population in Lake Maarsseveen and from these individuals clones were cultured in the laboratory. Some clones were assumed to be migrating and some non-migrating. Life history experiments were performed with these clones in the presence and absence of fish kairomones. Results show that clones differed in life history strategy in concert with their presumed migration strategies which suggests a link between life history and behaviour. However, no sharp border exists between migrating and non-migrating clones since a gradual response pattern was found for the traits investigated. In the absence of fish kairomones, the relation between length at maturity and number of eggs was found to differ between the clones of the non-migrating and the migrating group. The latter had a lower number of eggs at similar length values. In the presence of fish kairomones, the length–clutch size relation was the same. However, non-migrating clones were smaller at the age at maturity and had fewer eggs. Although clones of both groups were thus found to differ significantly, overlap existed. Age at maturity was found to be the same.     

Alternative antipredator responses of two coexisting Daphnia species to negative size selection by YOY perch

Our study showed for the first time in nature that two coexistingDaphnia adopted alternative life history and behavioural strategiesto cope with negative size-selection predation by gape-limitedyoung-of-the-year (YOY) perch. We evaluated the phenotypic plasticityin life history and behavioural traits of two coexisting Daphniaspecies, D. pulicaria (2 mm) and D. galeata mendotae (1.4 mm),in response to seasonal changes in predation by YOY yellow perch(Perca flavescens) in a mesotrophic lake. We expected that thelarge-sized D. pulicaria, the most likely subjected to size-selectivepredation by YOY perch, will show stronger antipredator responsesthan the small-sized D. galeata mendotae. To test this hypothesis,we examined changes in life history and behavioural traits injuveniles and adults of both species during four YOY fish predationperiods that were selected based on the presence of YOY perchin the pelagic zone and the relative abundance of Daphnia preyin their gut contents. Our study supports the scenario of negativesize-selective predation by gape-limited YOY perch on both Daphniaspecies. The electivity index indicated that no daphnids witha body length > 1.75 mm were predated by YOY yellow perch.Coexisting Daphnia exhibited phenotypic plasticity in theirantipredator defenses based on their vulnerability to seasonalchanges in size-selective predation of YOY perch. Juvenile Daphniawere the targeted prey and they responded by a decreased bodylength. Behavioural defenses were the dominant strategy usedby both adult Daphnia populations to withstand high predation.A decreased size at maturity was not employed by Daphnia, exceptat the very end of the predation period. Behavioural defensesare short-term strategy adopted to avoid predation. Both antipredatordefenses became unnecessary expenses and were no longer sustainedafter the predation period.     

Inter-clonal variation in phototactic behaviour and key life-history traits in a metapopulation of the cyclical parthenogen Daphnia ambigua: the effect of fish kairomones

We quantified genetic variation for ecological relevant traits in the presence and the absence of fish chemicals of eleven Daphnia ambigua clones that were isolated from six interconnected ponds that differed in water transparency. In a cohort life table experiment, we tested whether genetic variation for a set of key life history traits was present among these clones. In addition the phototactic behaviour of these clones cultured in the presence and the absence of fish kairomones was quantified using a biotest. We detected a significant effect of fish kairomones on the phototactic behaviour and a highly significant genetic variation among clones for this trait in D. ambigua clones isolated from ponds in De Maten. Differences in size at maturity among D. ambigua clones in De Maten were highly significant, whereas differences in spina length among D. ambigua clones in De Maten were not significant. The presence of fish chemicals did not affect the studied life-history traits. We observed a significant positive relationship between average phototactic behaviour for each population and size at maturity both in the presence as in the absence of fish kairomones. Most of the genetic differences could be attributed to a clone isolated from one clearwater pond that is not directly connected to the remainder of the pond complex.     

Impact of fish predation on coexisting Daphnia taxa: a partial test of the temporal hybrid superiority hypothesis

Fish predation was tested as a factor mediating the coexistence of Daphnia taxa in the shallow, hypertrophic Lake Blankaart. Naturally co-occurring populations of D. galeata and the hybrid D. galeata x cucullata were subjected to different levels of fish predation in in situ enclosures. In control enclosures without fish, the largest taxon D. galeata rapidly became dominant over the intermediate sized D. galeata x cucullata, mainly as a result of higher birth rates. In enclosures with fish, population densities of D. galeata dropped relative to D. galeata x cucullata, due to higher mortality rates. These results are in concordance with the `temporal hybrid superiority hypothesis', and can be explained by a higher vulnerability of the large and more conspicuous D. galeata to the size selective predation exerted by visually hunting planktivorous fishes. After approximately one month, however, population growth rates of D. galeata and D. galeata x cucullata in the enclosures with fish converged, due to a relative reduction in the mortality rate of D. galeata. This suggests that, in the presence of fish, D. galeata may co-exist with hybrids due to a decrease in its relative vulnerability to visual predation with time. Indeed, both D. galeata and the hybrid showed strong reductions in adult body size in the enclosures with fish, but this size reduction tended to be stronger in D. galeata than in D. galeata x cucullata. In addition, turbidity increased in the enclosures with fish and may additionally have reduced the relative advantage of D. galeata x cucullata with regard to mortality caused by visual predation.     

Life-history variation in a hybrid species complex ofDaphnia

Summary Life-history variation was examined among members of theDaphnia longispina group, which consists ofD. galeata, D. hyalina, D. cucullata, and hybrids. Factorial experiments were conducted at two temperatures (14° and 20° C) and two food concentrations (0.2 and 1.0 mg Cl⁻¹). Differences in life-history features (size at maturity, age at first reproduction, size of first clutch, offspring size in first clutch) under the different environmental conditions were assessed among eightDaphnia clones, which represented members of this species complex. Significant differences between parentals and hybrids for most life-history features were observed under various treatments; generally, hybrid clones showed intermediate life-history traits when compared with parentals. When comparisons were made among clones within a given species (i.e.D. galeata, D. galeata xcucullata, D. cucullata), clonal differences were also noted for certain life-history traits. The data are discussed with reference to the formation and maintenance of hybrid species complexes in nature. This paper is dedicated to the memory of Hans Georg Wolf, who died suddenly in May 1990 at the age of 39 years.     

Facing multiple enemies: parasitised hosts respond to predator kairomones

During their lifetime most organisms are exposed to various enemies influencing their victims in multiple direct and indirect ways. Most studies concentrate on the effects of one enemy at a time, thereby not taking into account that in nature organisms are often simultaneously exposed to more than one enemy. We conducted a life-history experiment to investigate the simultaneous effects of predators (fish, Leuciscus idus) and parasites (microparasite, Caullerya mesnili) on their victim (Daphnia galeata). D. galeata were exposed to predator kairomones, parasites or both. D. galeata are able to sense the presence of fish predators via chemical cues (= kairomones). Both fish predator kairomones and microparasite infections influence the life history of Daphnia. Some of the effects of fish predator kairomones are directly opposed to microparasite effects; fecundity, for example, is increased in the presence of fish kairomones and decreased in Daphnia parasitised with C. mesnili. We investigated the influence of both threats on age at maturity, body size at different adult instars, fecundity and survival of one D. galeata clone. In the presence of fish kairomones, all D. galeata matured significantly earlier and increased the number of eggs in the second brood significantly. Parasitised D. galeata matured significantly earlier than non-parasitised ones in the absence and presence of fish kairomones. An infection with the microparasite C. mesnili led to significantly lower clutch sizes at the second adult instar, to significantly smaller body sizes from adult instar three onwards and to significantly reduced survival. No significant interaction effect between the responses to fish presence and to parasite infection was found for any of the investigated life-history traits. The lack of interaction effects between the exposure to predator kairomones and parasite infection was most likely due to the different timing of the effects. Fish kairomones affected D. galeata early in its life history whereas C. mesnili increased in its effects over time. Our results show that parasitised D. galeata are able to exhibit life-history responses to fish predator presence early in their lives. Thus, D. galeata parasitised with C. mesnili have a similar chance as non-parasitised D. galeata to escape from fish predation via life-history changes. Since older parasitised D. galeata are smaller, they may have an even better chance to escape visual predators under actual predation.     

Effects of neonate size and food concentration on the life history responses of a clone of the hybrid Daphnia hyalina × galeata to fish kairomones

1. The influence of fish kairomones on the life history parameters of a clone of the hybrid Daphnia galeata × hyalina was measured in laboratory experiments with small and large neonates at one food concentration. The influence of fish kairomones on neonates of the same size at two food concentrations was also investigated.
2. Neonate size (maternal investment per egg) changed the influence of fish kairomones on life history parameters. Daphnids that were small at birth showed greater differences between fish (kairomones present) and no-fish (kairomones absent) treatments for age at maturity and number of eggs in the first clutch than animals that were large at birth. On the other hand, animals that were large at birth differed more in size at maturity between fish and no-fish treatments than small neonates.
3. The food concentrations used in the experiments (0.25–0.75 mg C l^–1) hardly affected the influence of fish kairomones on the life history characteristics studied.
4. The effect of fish kairomones on the size of neonates from the first clutch was similar for large and small daphnids. In the no-fish treatment significant differences were found between small and large neonates for age at maturity and number of eggs in the first clutch. In the fish treatment size at maturity differed significantly between small and large neonates.
5. In life history experiments, neonate size has to be taken into account when measuring the effects of fish kairomones because size at birth influences the results.     

Effects of neonate size and food concentration on the life history responses of a clone of the hybrid Daphnia hyalina × galeata to fish kairomones

1. The influence of fish kairomones on the life history parameters of a clone of the hybrid Daphnia galeata × hyalina was measured in laboratory experiments with small and large neonates at one food concentration. The influence of fish kairomones on neonates of the same size at two food concentrations was also investigated.
2. Neonate size (maternal investment per egg) changed the influence of fish kairomones on life history parameters. Daphnids that were small at birth showed greater differences between fish (kairomones present) and no-fish (kairomones absent) treatments for age at maturity and number of eggs in the first clutch than animals that were large at birth. On the other hand, animals that were large at birth differed more in size at maturity between fish and no-fish treatments than small neonates.
3. The food concentrations used in the experiments (0.25–0.75 mg C l^–1) hardly affected the influence of fish kairomones on the life history characteristics studied.
4. The effect of fish kairomones on the size of neonates from the first clutch was similar for large and small daphnids. In the no-fish treatment significant differences were found between small and large neonates for age at maturity and number of eggs in the first clutch. In the fish treatment size at maturity differed significantly between small and large neonates.
5. In life history experiments, neonate size has to be taken into account when measuring the effects of fish kairomones because size at birth influences the results.     

Mechanisms regulating zooplankton populations in a high-mountain lake

SUMMARY 1. We studied the seasonal succession of phyto- and zooplankton and the potential impact of predation by salmonids on zooplankton population dynamics in a high-mountain Swiss lake.
2. A comparison of patterns in the abundance, body length, fecundity and age structure in the Daphnia galeata population strongly suggests that trout predation had little impact on the population and was not the cause for a decline in summer.
3. The dominance in the lake of adult trout that feed mainly on benthic prey may buffer the effect of predation on the larger zooplankton. Further, the relatively high amount of phytoplankton after spring thaw could be important for sustaining the Daphnia population under moderate fish predation.
4. Partial correlation analyses proved circumstantial evidence for both exploitative and interference competition between some zooplankton taxa. D. galeata depressed performance of other plankton species through exploitative competition.
5. Our study shows that the impact of fish on zooplankton in high-mountain lakes depends strongly on food web structure and trophic state of the lake. Where fish predation is weak, invertebrate predation combined with competition for food may be responsible for the dominance of large-bodied zooplankton species.     

Preliminary experiments on resource competition between a migrating and a non-migrating clone of the hybrid D. galeata×hyalina

Competition experiments between two clones of thehybrid D. galeata × hyalina differing invertical migration behaviour were carried out in thelaboratory, in a set-up which separated the clonesspatially. The influence of fish kairomones,artificial predation and the imitation of dielvertical migration circumstances were investigated. Inthe presence of fish kairomones, the non-migratingclone outnumbered the migrating one. The introductionof artificial predation reduced this effect in such away that the migrating clone had a competitiveadvantage over the non-migrating clone. It wasconcluded that competition results do not contradictlife history studies performed with the same twoclones, since the presence of fish kairomones enhancesthe reproduction of the non-migrating clone more thanthat of the migrating one.     

The combined effects of hypoxia and fish kairomones on several physiological and life history traits of Daphnia

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Temperature effects on chemical signalling in a predator–prey system

SUMMARY 1. We investigated the effect of temperature on chemical signalling in a predator–prey model system (planktivorous fish and Daphnia galeata ). Life-history changes in Daphnia in response to chemical cues (kairomones) derived from fish have become a paradigm for chemically induced anti-predator defences.
2. As temperature can affect both predator and prey, we carried out two experiments to disentangle these effects. In order to test for temperature effects on the predator, we kept prey at a single temperature and exposed them to kairomones from fish exposed to two different temperatures. Daphnia exhibited a higher intrinsic rate of population increase ( r ) when exposed to fish kairomones produced at high rather than low temperature. Assuming a positive correlation between r (because of an earlier maturation and/or increased clutch sizes) and kairomone concentration, our results suggest that kairomone production increases with rising temperature.
3. In the second experiment, to study the influence of temperature on the prey, Daphnia were kept at two different temperatures and exposed to fish kairomones produced at one constant temperature. We found no interaction between the effects of fish kairomone and temperature on Daphnia life history, suggesting that temperature does not directly alter life-history responses to fish kairomones.
4. Our results suggest that temperature influences Daphnia life history through its effects on fish kairomone concentration, but that temperature does not affect the strength of the response of Daphnia to the presence of fish.     

The prevalence of diapause response to risk of size-selective predation in small- and large-bodied prey species

The prevalence of diapause response to the simulated threat of fish predation was compared in three species of planktonic crustaceans of the genus Daphnia (D. magna, D. pulicaria and D. longispina), which due to their different body size vary in vulnerability to fish predation pressure in natural conditions. Higher incidence of diapause response was presumed in the larger-bodied species, which due to their higher conspicuousness and higher energetic content experience the greatest size-selective pressure from visually foraging fish. Small-bodied species were expected to utilize less costly yet less effective active defences, e.g. lower conspicuousness achieved due to slower body growth when facing risk of size-selective predation. Proportions of females forming diapausing structures as well as females body size at the maturation period were compared in experimental beakers containing or not containing fish kairomones (chemical cues of fish predation) in a few clonal lineages of 3 species of Daphnia derived from a single lake inhabited by fish. The highest incidence of diapause response to fish kairomones was observed in D. magna (when measured both as proportion of sensitive individuals and as proportion of inductive clones), the largest of the tested Daphnia species. The lowest proportion of individuals and clones that employed diapause was reported in the smallest tested species, D. longispina. In addition, the large-bodied Daphnia (D. magna) showed a greater reduction in body size in response to fish kairomones than the small-bodied species (D. longispina). The results of the present study support the assumption of higher prevalence of diapause response to the threat of selective predation in larger and more vulnerable prey species.     

Spatial distribution of the Daphnia longispina species complex and other planktonic crustaceans in the heterogeneous environment of canyon-shaped reservoirs

Canyon-shaped reservoirs are often characterised by longitudinalgradients of environmental factors (including trophic level,phytoplankton and zooplankton biomass and abundance of planktivorousfish) affecting the taxonomic composition of the pelagic community.We tested the hypothesis that the spatial distribution of differentspecies and interspecific hybrids of the Daphnia longispinaspecies complex is non-random under such conditions. Duringthe summer stratification, we sampled crustacean zooplanktonfrom 11 reservoirs, covering both longitudinal (upstream, middle,dam) and vertical (epi-, meta- and hypolimnion) environmentalgradients. Allozyme electrophoresis was used to discriminateamong different Daphnia taxa. All three frequently hybridizingEuropean species of the complex (galeata, cucullata, longispina= hyalina) and hybrids with Daphnia galeata were commonly recorded.Smaller-bodied Daphnia cucullata and its hybrids, when present,preferred mostly the nutrient- and food-rich upstream regions;D. longispina and its hybrids were more commonly found in thedownstream part, and often dominated in the meta- or hypolimnion.Redundancy analyses confirmed significant differences in theDaphnia taxon composition, as well as in spatial distributionof other crustacean species, along both gradients. For the firsttime, we demonstrate regular patterns in the horizontal distributionof Daphnia species and hybrids within a water body, thus acceptingour hypothesis. Such spatial distributional patterns may stronglyimpact local hybridization processes.     

The possibility of predator avoidance by Lake Michigan zooplankton

Low densities of Diaptomus ashlandi, Diacyclops thomasi, and Daphnia galeata mendotae were measured at depths where Mysis relicta formed nighttime aggregations. Calculations suggest that mysid predation can not account for the rarity of prey animals at these depths, which further suggests that the prey avoided the mysids. Unlike D. galeata mendotae, Daphnia pulicaria was common in mysid aggregations. The somewhat larger size of D. pulicaria may reduce its vulnerability to mysid predation, and consequently may explain the vertical distribution differences between the two congeners. Vertical distributions of Limnocalanus macrurus and copepod nauplii showed no obvious relationships to the mysid distributions. These were the only two taxa with distributions that were correlated with chlorophyll a concentrations. All crustacean taxa were rare in the epilimnion at night when sonar recorded a dense fish school.     

Behavioural versus physiological mediation of life history under predation risk

Predator-generated variation in prey energy intake remains the dominant explanation of adaptive response to predation risk in prey life history, morphology and physiology across a wide range of taxa. This "behavioural hypothesis" suggest that chemical or visual signals of predation risk reduce prey energy intake leading to a life history characterized by a small size and late age at maturity. However, size-selective predation can induce either smaller size-early age or large size-late age life history. The alternative "physiological hypothesis" suggests that size-selective cues decouple the relationship between energy and life history, acting instead directly on development. Here we use a series of experiments in a fish-daphnid predator-prey system to ask whether size-selective predator cues induce a physiological mediation of development, overshadowing behaviourally based changes in food intake. We found fish chemical cues reduce the net energy intake in Daphnia magna, suggesting a behaviourally mediated reduction in energy. Experimental manipulation of food levels show further that reductions in food lead to later but smaller size at maturity. However, in line with the physiological hypothesis, we show that D. magna matures earlier and at a smaller size when exposed to fish predation cues. Furthermore, our data shows that they do this by increasing their development rate (earlier maturity) for a given growth rate, resulting in a smaller size at maturity. Our data, from a classic size-selective predation system, indicate that predator-induced changes in this system are driven by physiological mediation of development rather than behavioural mediation of energy intake.