Genetic differentiation in life history between Daphnia galeata populations: an adaptation to local predation regimes?

Species of the water flea Daphnia exhibit constitutive as wellas phenotypically inducible anti-predator defence strategies,involving life history, morphological and behavioural traits.We explored the hypothesis of genetic differentiation in anti-predatordefence strategies using Daphnia clones originating from twodifferent water bodies: Tjeukemeer (the Netherlands) and FishPond (Belgium). Both water bodies are inhabited by zooplanktivorousfish. In contrast to Tjeukemeer, Fish Pond is also inhabitedby larvae of the phantom midge Chaoborus. The life history responsesof the two sets of clones to kairomones from fish (Perca), tokairomones from Chaoborus, and to a mixture of both were compared.Clones from Tjeukemeer and Fish Pond showed strong responsesto the presence of fish kairomone, with reductions in adultand neonate body length, in age at first reproduction and inthe total number of neonates produced during the first threeadult instars. Responses to Chaoborus kairomone were much lesspronounced, although there was a tendency towards an increasein the number of neonates in the first brood. Significant inter-populationgenetic differences were found for all the investigated traits.However, there was no indication for genetic adaptation of theFish Pond clones to negative size-selective predation by Chaoborus.Compared to Tjeukemeer clones, Fish Pond clones had a lowersize at first reproduction, produced smaller neonates and produceda higher number of neonates in their first brood. This suggestsadaptation to positive rather than to negative size selectivepredation. Genetic differences between populations were observedmainly for constitutive life history traits, rather than forphenotypic shifts in response to the presence of predator kairomones.     

Phenotypic plasticity of Daphnia life history traits: the roles of predation, food level and toxic cyanobacteria

1. We studied the life history responses of Daphnia pulex under different biotic conditions. In a factorially designed experiment, we tested the impacts of water conditioned by the invertebrate predator Chaoborus, low and high food level (10 000 and 100 000 Scenedesmus cells ml^–1), and exposure to toxic Microcystis (5000 cells ml^–1) on twelve D. pulex clones originating from different habitats. Our aim was to compare the phenotypic plasticity of different clones, and to study the interactions among biotic factors.
2. Individuals cultured in Chaoborus -conditioned water started to reproduce at a larger size than individuals cultured in water not conditioned by the predators. We found interactions between food level and Chaoborus -conditioned water on age at first reproduction and total offspring number. In addition, the impact of the Chaoborus treatment on the size at first reproduction was reduced by Microcystis exposure.
3. Clonal differences were found in the degree of phenotypic plasticity of different life history traits. However, there was no obvious connection between the original habitat of the clone and the degree of plasticity.     

Components of additive variance in life-history traits of Daphnia hyalina: seasonal differences in the response to predator signals

We attempted to separate the additive variance of life-history parameters (size at first reproduction (SFR), proportion of total production allocated to reproduction) of Daphnia hyalina in a mesotrophic lake during different seasons into genetic components and phenotypic plasticity. Every month we randomly isolated juveniles of 20 clones from the lake and measured their life-history parameters immediately after isolation, i.e., under the influence of all modifying factors in the lake. The measurements were repeated after the clones had been kept in the laboratory for at least six generations without predator signals (controls) and with the addition of fish and Chaoborus kairomones. SFR increased monotonously from July to December in fresh field isolates. Laboratory controls showed always much larger SFR, but approached field value in December. Most of the size difference was due to maternal effects. Reproductive allocation showed a different pattern than SFR with a minimum in September both in field samples and laboratory controls. Fish kairomone reduced SFR in July and August when fish predation is high, but not later in the year. On the contrary, Chaoborus kairomone increased SFR in September and October when Chaoborus larvae are abundant in the lake, but not in summer. This indicates a seasonal shift in the clonal composition of D. hyalina populations towards clones adapted to the specific predation pressure. Reproductive allocation changed in response to fish kairomone in July and August, but not in response to Chaoborus kairomone. In this mesotrophic lake, fish have a stronger direct and indirect impact on Daphnia life history than Chaoborus larvae.     

Environmental correlates of the intrinsic rate of natural increase in primates

Morphological and life history traits of two clones of the cladoceran Daphnia pulex were measured in the presence and absence of size-selective insect predators, the midge larva Chaoborus flavicans, which preys on small Daphnia, and the water bug Notonecta glauca, which preys on large Daphnia. The aim was to detect predator-induced phenotypic changes, particularly the effect of simultaneous exposure to both types of predators. Other work has shown that in the presence of Chaoborus americanus, Daphnia pulex produce a socalled neck spine which may carry several teeth. The morphological modifications are supposed to serve as an anti-predator device. Furthermore, females exposed to Chaoborus often delay their maturation; this has been interpreted as a cost that balances the benefits of the neck teeth. In this investigation, females of both clones produced fewer but larger offspring than control animals when reared in the presence of Chaoborus flavicans. The offspring showed the typical neck spine and delayed first reproduction. In the presence of Notonecta glauca, one of the clones produced more and smaller offspring, and maturation occurred at earlier instars. The other clone also produced more offspring than the control but there was no size difference. When both predators were present, in most cases the reactions of the daphnids were similar to those in the Notonecta experiment. The response to Chaoborus appeared to be suppressed. The observed modifications are interpreted as evolved strategies that reduce the impact of size-selective predation. They are consistent with predictions of life-history theory.     

Waterborne metals impair inducible defences in Daphnia pulex: morphology, life-history traits and encounters with predators

1.  Inducible defences may be temporary and favoured where predation is intermittent and have been demonstrated in several invertebrates and vertebrates when prey detect chemical cues (kairomones) released by predators. Daphnia pulex (a water flea) exposed to Chaoborus (midge larvae) kairomones produce small neckteeth on the dorsal surface of the head as a defence against this gape-limited predator and survive better in the presence of Chaoborus . Recent studies have shown that waterborne copper (Cu) impairs the induction of neckteeth which could lead to lower survival.
2.  Here, we examined the effects of Cu on morphological changes and shifts in life-history traits in D. pulex exposed to kairomone from Chaoborus americanus . We exposed D. pulex mothers to chemical cues of C. americanus fed on either D. pulex neonates or on brine shrimp Artemia salina , the same Chaoborus cues combined with an environmentally relevant concentration of copper (10 μg L⁻¹), or dechlorinated tap water. We examined several morphological characteristics of neonates and life-historical characteristics of adults as well as assessing survival of neonates by staging encounters with predators.
3.  Neonates from mothers exposed to kairomone plus copper had fewer and shorter neckteeth than neonates from mothers exposed to kairomone alone. Moreover, neonates exposed to Cu had lower survival during encounters with predators than neonates exposed to kairomone without Cu.
4.  Adult female Daphnia exposed to kairomones released more neonates within the first 24 h of brood release and emptied their brood pouches quicker than mothers not exposed to kairomones, irrespective of the presence of Cu.
5.  Impairment by metals of morphological defences in zooplankton could lead to a decline in population density and alter community structure.     

Size distribution of Daphnia longispina in the vertical profile

D. longispina of the meromictic lake El Tobar is a round-headed form. It never has a helmet, but in summer a small proportion of immature individuals (0.9–1.2 mm females and males) have one or two neck teeth. The size structure of this Daphnia population, as well as the vertical distribution and migration of different size-classes, were studied in September and November of 1991 and April of 1992. The large variation in mean size and size at first reproduction, as well as the occurrence of different patterns of vertical migration are interpreted as responses to different predator situations. At the end of April, when Daphnia mortality by visually hunting predators is dominating, a typical nocturnal migration is adopted and size distribution is biased to smaller size classes. In November, when mortality is mainly attributed to the nocturnally migrating Chaoborus, Daphnia shows a reversed migration pattern. In September, when the population of Daphnia is responding to both visual (fish) and non-visual predators (Chaoborus), it adopts a pattern of twilight migration. The presence of neck teeth in vulnerable size classes in September might be an additional adaptative response to Chaoborus predation. In September, the size structure of the Daphnia population is shifted to larger classes and the vertical distribution of size classes shows a pronounced segregation between juveniles and adults. Juveniles are found closer to the surface, while adults dwell predominantly in the rich, deep waters near the oxicline. This suggests that an additional advantage of the ascent of the adult Daphnia exploiting those deep resources is the release of young in more oxygenated and warmer waters. The Daphnia population of lake El Tobar is known to be clonally diverse, and the changing frequency of genotypes could play an important part in the observed seasonal differences in behaviour and size.     

Costs of predator‐induced morphological defences in Daphnia

1. Inducible defences are advantageous because they protect the prey while limiting associated fitness costs. The presence of these costs is an essential component of this conditional strategy, since their absence would favour constitutive (fixed) defences. In some cases, however, these costs have been difficult to measure because of complex interactions between the defences themselves, resultant life history changes and the organism’s environment. 2. The pond‐dwelling water flea, Daphnia pulex, forms defensive neck spines in response to kairomones released by predatory larvae of the phantom midge, Chaoborus. This predator–prey interaction and the formation of these inducible defences have been well studied, but costs associated with the development of neck spines remain unclear. In this study, I address this problem by analysing the effect of Chaoborus kairomones on the life history responses (and fitness costs associated with these responses) of two clones of D. pulex that are from the same pond population, but differ greatly in their degree of neck spine development. 3. Both D. pulex clones exhibited the same predator‐induced shifts in life history: larger size at birth, reduced juvenile growth rate (producing a smaller size at maturity), delayed reproduction and a reduction in the number of neonates produced after the first clutch. Relative fitness decreased significantly and to the same degree (c. 10% reduction in r) in each clone. This observed fitness cost was not directly related to the neck spines per se since the cost was the same in both clones, despite their considerable differences in neck spine development. Rather, it appears to be indirectly related to this antipredator morphology via a combination of delayed reproduction and a set of life history trade‐offs (decreased growth rate, decreased reproduction after the first clutch) for increased neonate body size, which is necessary for neck spines to be effective defences. This suite of induced responses is probably a result of local adaptation of these two D. pulex clones to their common pond environment. 4. Costs of inducible defences do not always entail direct allocation costs associated with forming and maintaining a defence, but may also involve indirect life history responses that are specific to particular environmental situations. This local adaptation would explain the highly variable life history responses observed among D. pulex clones from different pond environments.     

Effect of vertebrate and invertebrate kairomones on the life history of Daphnia magna Straus (Crustacea: Branchiopoda)

The history of selection of Daphnia magna populations living in North African temporary ponds may differ from populations inhabiting permanent ponds. Laboratory experiments were conducted to examine the effect of fish Gambusia holbrooki and invertebrate Notonecta glauca kairomones on the life history traits of the freshwater Cladocera Daphnia magna Straus. With fish kairomones, Daphnia reproduced early and had a significantly smaller size at first reproduction (SFR) and a smaller size of neonates compared to control. In contrast, daphnids reared in water treated with Notonecta glauca had no effect on the age at first reproduction but females were also smaller and produced smaller neonates.     

Influence of food density on the effects of a Chaoborus-released chemical on Daphnia ambigua

SUMMARY 1. Daphnia ambigua was reared individually at four different food levels in water conditioned by a predator, Chaoborus flavicans. Daphnia ambigua developed spike-like helmets during the first four instars in each treatment with the Chaoborus-conditioned water, but only during the first instar in untreated water. The helmet development at instars 2–4 was probably induced by a chemical released from the Chaoborus .
2. The helmets decreased in size with decreasing food levels, but never ceased to form, even at a very low food concentration.
3. Daphnids cultured in the Chaoboras -conditioned water exhibited reduced growth, reproduction and survival rates at low food levels. These results might be induced by the energy loss associated with producing the helmets.
4. It is suggested that the predator releases a chemical, which reduces the tolerance of the cyclomorphic Daphnia to food deficiency, thus accentuating the summer decline in the Daphnia population caused by food shortage.     

Predator-induced phenotypic plasticity in the exotic cladoceran Daphnia lumholtzi

1. The exotic cladoceran Daphnia lumholtzi has recently invaded freshwater systems throughout the United States. Daphnia lumholtzi possesses extravagant head spines that are longer than those found on any other North American Daphnia. These spines are effective at reducing predation from many of the predators that are native to newly invaded habitats; however, they are plastic both in nature and in laboratory cultures. The purpose of this experiment was to better understand what environmental cues induce and maintain these effective predator‐deterrent spines. We conducted life‐table experiments on individual D. lumholtzi grown in water conditioned with an invertebrate insect predator, Chaoborus punctipennis, and water conditioned with a vertebrate fish predator, Lepomis macrochirus. 2. Daphnia lumholtzi exhibited morphological plasticity in response to kairomones released by both predators. However, direct exposure to predator kairomones during postembryonic development did not induce long spines in D. lumholtzi. In contrast, neonates produced from individuals exposed to Lepomis kairomones had significantly longer head and tail spines than neonates produced from control and Chaoborus individuals. These results suggest that there may be a maternal, or pre‐embryonic, effect of kairomone exposure on spine development in D. lumholtzi. 3. Independent of these morphological shifts, D. lumholtzi also exhibited plasticity in life history characteristics in response to predator kairomones. For example, D. lumholtzi exhibited delayed reproduction in response to Chaoborus kairomones, and significantly more individuals produced resting eggs, or ephippia, in the presence of Lepomis kairomones.     

Life-History Characteristics and Fitness in Descendents of Parthenogenetic and Ex-Ephippio Females of Daphnia Magna

Daphnids of ex-ephippial and parthenogenetic origin differ substantially in life-history. Possible maternally transmitted ex-diapause effects of differing female origin on the fitness of their offspring were studied across multiple clones in Daphnia magna. Ex-ephippial daphnids responded in egg size to different constant food concentrations with the same pattern as parthenogenetic females. Significant differences in egg characteristics between females of different origin were only found for the first clutch produced under high food. The smaller size of eggs in ex-ephippial females, however, did not translate into size differences of first-clutch neonates. A trend for lower mass density per body volume was detected in offspring from ex-ephippial females in comparison with parthenogenetic daphnids. Hence, an ex-diapause effect transmitted by ex-ephippial mothers to their parthenogenetic offspring is likely. However, there was no difference in life-histories and fitness between offspring produced by females of ex-ephippial and parthenogenetic origin at both, high and low food concentrations across multiple clones. Thus, a significant ex-diapause effect on fitness in successive parthenogenetic generations may not be expected in D. magna at the population level.     

CHAOBORUS PREDATION AND LIFE-HISTORY EVOLUTION IN DAPHNIA PULEX: TEMPORAL PATTERN OF POPULATION DIVERSITY,FITNESS, AND MEAN LIFE HISTORY

The effect of predation by the aquatic dipteran larva Chaoborus americanus on genetic diversity and life-history evolution in the cladoceran Daphnia pulex was investigated in large replicate laboratory populations. Instantaneous daily loss rates of clonal diversity and genetic variance for fitness indicate that 93–99% of initial genetic diversity can be removed from populations during the 8–12 generations of clonal reproduction that occur each year in natural populations. In the absence of predation, the principal evolved changes in mean population life history were smaller immature body size and increased and earlier fecundity. In the presence of size-selective Chaoborus predation, populations evolved toward larger body size and increased and earlier reproduction. The difference between these two trajectories is an estimate of the direct additive effect of Chaoborus predation. This effect was manifested as evolution toward larger body size with a trend toward earlier and increased reproduction.     

Life history variation among low-arctic clones of obligately parthenogenetic Daphnia pulex: a diploid-polyploid complex

Summary Laboratory life table experiments were conducted using nine clones of obligately parthenogenetic Daphnia pulex that were collected from a site in the Canadian low-arctic. Two of the nine clones were diploids, while the other seven clones were polyploids. Significant clonal differences in age at first reproduction, size at first reproduction, number of offspring in each of the first three broods, offsrring sizes for the first two broods, and intrinsic rates of natural increase were detected. Differences in life histories were evident between polyploids and diploids. Generally, polyploid clones reached maturity at later ages, matured at larger sizes, produced smaller broods, and larger offspring than the diploid clones. The data are discussed in reference to potential biotic (i.e. invertebrate predation) and abiotic factors (i.e. physicochemical gradients) that may influence life history variation in this clonal assemblage.     

Resource partitioning in two coexisting cladocerans,Daphnia magna and Scapheloberis kingi (Anomopoda: Daphniidae), in north-eastern Algeria

Life-history traits of two coexisting cladocerans, Daphnia magna Straus and Scapheloberis kingi Sars, inhabiting a temporary pond in north-eastern Algeria were monitored in 2013 under laboratory conditions. Their life histories were compared for differences in traits such as age and size at first reproduction, size of neonates, brood size, number of broods per female, total life span and intrinsic rate of increase (r_m). Data were recorded during their entire life cycle. Daphnia magna, the larger species, could possibly be more successful in colonising temporary habitats than S. kingi because it allocates more energy to reproduction. Scapheloberis kingi, the smaller species, starts reproduction early, resulting in a smaller brood size and a shorter life span with fewer broods. Consequently, S. kingi produces a smaller number of neonates during its lifetime and has a lower r_m. In contrast, D. magna delays reproduction but produces a larger brood size and, because it is longer-lived, produces more broods and so produces a greater number of neonates and has a higher r_m over its lifetime.     

Phenotypic plasticity of Daphnia life history traits in response to predator kairomones: genetic variability and evolutionary potential

Cladoceran populations can respond to changingpredation regimes by a phenotypical response as wellas by shifts in genotype frequencies. In this study,we investigated the phenotypic plasticity exhibited bylife history traits of D. galeata in response tothe presence of predator kairomones, as well as theextent to which natural selection may act on thesetraits and their phenotypic plasticity. In alife-table experiment, seven clones of a natural D. galeata population were subjected to kairomonesfrom fish (Perca), from an invertebrate predator(Chaoborus) or a mixture of both. Life historytraits were affected by the kairomones of bothpredators, but effects of Chaoborus wereneutralised by Perca in the kairomone mix. Noapparent trade-off was found between growth- andreproduction related traits: although daphnids fromthe Chaoborus treatment grew faster thandaphnids from the other treatments, no reduction inthe reproductive output was observed. Broad-senseheritabilities were found to be relatively high forsome life history traits (size at maturity, neonatesize, number of neonates) as well as for thephenotypic plasticity response of these traits. Thisreflects the evolutionary potential of life historytraits and their phenotypic response to predatorkairomones in the D. galeata population.Publication number 2334 of The Netherlands Institute of Ecology, Centre for LimnologyPublication number 2334 of The Netherlands Institute of Ecology, Centre for Limnology     

Interactions Between Predator Kairomone and Food Level Complicate the Ecological Interpretation of Daphnia Laboratory Results

The effect of food concentration on the phenotypic responseof life history traits to two predator kairomones was investigatedin Daphnia. For the experiment, one clone of Daphnia galeatawas used as the prey organism and solutions containing infochemicals(kairomones) of Chaoborus (phantommidge) and Perca (perch) simulatedthe presence of potential predators. The combined effect offood level and predator kairomone on phenotypic plasticity ofDaphnia life history is complex. At low food levels, the responsesin life history traits to kairomones are enhanced. For example,in the Perca treatment, this leads to a younger age at firstreproduction. The presence of Chaoborus kairomone decreasessurvival so that death often occurs before reproduction is reached.This results in a strong decrease in the intrinsic rate of populationincrease and might contribut to the pronounced summer declineof some Daphnia populations during periods of food shortage.     

Transgenerational effects of poor elemental food quality on Daphnia magna

Environmental effects on parents can strongly affect the phenotype of their offspring, which alters the heritability of traits and the offspring’s responses to the environment. We examined whether P limitation of the aquatic invertebrate, Daphnia magna, alters the responses of its offspring to inadequate P nutrition. Mother Daphnia consuming P-poor algal food produced smaller neonates having lower body P content compared to control (P-rich) mothers. These offspring from P-stressed mothers, when fed P-rich food, grew faster and reproduced on the same schedule as those from P-sufficient mothers. In contrast, offspring from P-stressed mothers, when fed P-poor food, grew more slowly and had delayed reproduction compared to their sisters born to control mothers. There was also weak evidence that daughters from P-stressed mothers are more susceptible to infection by the virulent bacterium, Pasteuria ramosa. Our results show that P stress is not only transferred across generations, but also that its effect on the offspring generation varies depending upon the quality of their own environment. Maternal P nutrition can thus determine the nature of offspring responses to food P content and potentially obfuscates relationships between the performance of offspring and their own nutrition. Given that food quality can be highly variable within and among natural environments, our results demonstrate that maternal effects should be included as an additional dimension into studies of how elemental nutrition affects the physiology, ecology, and evolution of animal consumers.     

Resource quality effects on Daphnia longispina offspring fitness

The present study addressed the question: is the fitness ofDaphnia longispina neonates related to the quality (species)of the resource their mothers consumed? A 2 2 factorial designwith two maternal and neonate resource types was employed, withRhodomonas and Microcystis used as high- and low-quality resources,respectively. Neonate fitness was assessed using demographicparameters obtained from a life-table study. The resource typeneonates consumed always had a stronger effect on the neonatesthan did the maternal type. Neonate consumption of Rhodomonas.relative to Microcystis. increased fitness, as estimated bythe instantaneous population growth rate, by 25–28%. whilematernal consumption of Rhodomonas increased fitness by 6-8%.Neonate Rhodomonas consumption also reduced the age at reachingthe primiparous instar; increased brood size at the primiparousinstar and body length immediately after the primiparous instar;increases average dutch size, number of clutches produced, totalegg production. and mean body length up to and on age 30 days.Regardless of resource type, maternal Rhodomonas consumptionresulted in neonates which reached maturity at an earlier ageand had a larger individual size immediately after reachingmaturity. The mean clutch size, total egg production, and lengthup to and at age 30 days were all significantly affected bymaternal diet when neonates consumed Microcystis; however, thesevariables were not influenced by maternal type when neonatesconsumed Rhodomonas. This is the first study to document maternallymediated effects of resource quality on the fitness of crustaceanzooplankton neonates.     

Testing hypotheses for maternal effects in Daphnia magna

Maternal effects are widely observed, but their adaptive nature remains difficult to describe and interpret. We investigated adaptive maternal effects in a clone of the crustacean Daphnia magna, experimentally varying both maternal age and maternal food and subsequently varying food available to offspring. We had two main predictions: that offspring in a food environment matched to their mothers should fare better than offspring in unmatched environments, and that offspring of older mothers would fare better in low food environments. We detected numerous maternal effects, for example offspring of poorly fed mothers were large, whereas offspring of older mothers were both large and showed an earlier age at first reproduction. However, these maternal effects did not clearly translate into the predicted differences in reproduction. Thus, our predictions about adaptive maternal effects in response to food variation were not met in this genotype of Daphnia magna.     

Neck spine protects Daphnia pulex from predation by Chaoborus, but individuals with longer tail spine are at a greater risk

We tested the prey preference of Chaoborus sp. on Daphnia pulexwith different defensive morphologies. The protective functionof inducible morphological defences, such as neck spine andlonger tail spine, was evaluated Second instar D.pulex individualsof two clones differing in their strength of neck spine inductionwere offered as prey to both Chaoborus obscuripes and Chaoborusflavicans. We used logistic regression analysis to evaluatethe effect of morphometry on the vulnerability of Daphnia. Thepresence of a neck spine and increased total length protectedD.pulex from Chaoborus predation. However, individuals witha longer tail spine were more vulnerable to Chaoborus predationChaoborus obscuripes was able to eat daphnids with a neck spinedue to the larger gape size of this chaoborid. The smaller speciesC.flavicans almost always ate prey with no neck spine.