Responses to injured conspecifics and heterospecifics by water fleas,Chydorus sphaericus

Cladocerans have been shown to exhibit a variety of responses to chemical stimuli associated with predation risk, including those from predators and injured conspecifics. In this study, we tested the hypothesis that individuals of Chydorus sphaericus would respond with avoidance behavior to chemical cues from both injured conspecifics and injured heterospecifics. Avoidance behavior was seen in response to cues from C. sphaericus and Daphnia magna. Responses to cues from Daphnia pulex were intermediate between those of the other cues and the water control. We propose that a concentration effect may be responsible for the mixed responses to heterospecific cues and speculate that responses to a broad range of cues may help explain the expansive range of C. sphaericus.     

Sperm number assessed by flow cytometry in species of Daphnia (Crustacea,Cladocera)

Daphnia magna and Daphnia pulex are two important model species in ecotoxicology. In daphniids, studies of the effects of contaminants have mostly focused on female life history traits, yet it would also be important to examine male reproductive traits, particularly in relation to endocrine disruptors. In this study, we developed a protocol that uses flow cytometry to measure sperm number in individual males of different species of Daphnia. We tested our protocol on 114 males from several clones of three common species of Daphnia. Sperm count varied widely among individuals and reached high numbers (up to 1.45 × 10⁵). Positive relationships between male length and sperm number were observed in D. pulex and Daphnia pulicaria, but not in D. magna. Important inter‐clonal differences in sperm production were observed in all species, with some clones producing very little sperm. Duplicated sperm samples showed on average only 6% difference in sperm counts. Sperm counts were stable at least over a 2‐hr period and up to 5 hr for most samples. This sperm isolation protocol and flow cytometric enumeration approach will be of major interest to ecotoxicologists.     

The Effects of Density on the Learned Recognition of Heterospecific Alarm Cues

Numerous species, both aquatic and terrestrial, use alarm cues to mediate predation risk. These cues may be either intentionally or inadvertently released, and may be received by either conspecifics or heterospecifics. In aquatic systems, alarm cues are often chemical in nature and are released when an organism is disturbed or damaged by a predator. In some cases the recognition of alarm cues from conspecifics, or closely related heterospecifics, is innate, while the recognition of alarm cues from distantly related species must be learned. Many studies have documented the use of heterospecific alarm cues, but few have explored the manner in which these cues come to be recognized as an indication of predation. In the current study, we examined the fathead minnow (Pimephales promelas)/brook stickleback (Culaea inconstans) alarm system. We tested the effect of density on the ability of minnows to learn to recognize stickleback alarm cues as a threat. We hypothesized that the ability of minnows to learn to recognize stickleback alarm cues should increase with increasing stickleback density because there would be more opportunity for minnows to associate the heterospecific alarm cue with the threat. To test this hypothesis we stocked minnows into large outdoor pools with no stickleback, low numbers of stickleback, or high numbers of stickleback. All pools contained a predator (pike, Esox lucius) known to the minnows. Following a 14 d conditioning period, minnows were tested for a response to skin extract from stickleback, minnow, and an unknown heterospecific (swordtail, Xiphophorus helleri). Minnows from pools with large numbers of stickleback learned to respond to stickleback alarm cues while minnows from pools with low numbers of stickleback, or no stickleback, did not.     

Temperature affects selectivity of Chaoborus larvae-eating Daphnia

In ponds, a chemical produced by predaceous Chaoborus (Insecta, Diptera) larvae changes the development of juvenile Daphnia pulex (Crustacea, Branchiopoda) so the juveniles grow spines (neckteeth) on the back of their head. It is generally assumed that the spined phenotype is (or is an indicator of) a morphological predator defense. The research reported here tests the hypothesis that the induced neckteeth do in fact increase Daphnia survivorship, over a range of temperatures. Predation experiments were conducted over a range of temperatures from 6 to 22 °C using fourth instar Chaoborus americanus larvae as the predator. The prey were a mixture of spined (induced necktooth phenotype) and unspined (uninduced) juvenile Daphnia pulex. At 6 and 11 °C, Chaoborus selected the unspined phenotype over the spined phenotype, as expected. However, at 22 °C, the selectivity was reversed: significantly fewer on the spined survived compared to the unspined phenotype. These results suggest that the spined phenotype may either increase or decrease Daphnia pulex survival, depending on temperature and clone.     

Ecological causes of morphological evolution in the three‐spined stickleback

The central assumption of evolutionary theory is that natural selection drives the adaptation of populations to local environmental conditions, resulting in the evolution of adaptive phenotypes. The three‐spined stickleback (Gasterosteus aculeatus) displays remarkable phenotypic variation, offering an unusually tractable model for understanding the ecological mechanisms underpinning adaptive evolutionary change. Using populations on North Uist, Scotland we investigated the role of predation pressure and calcium limitation on the adaptive evolution of stickleback morphology and behavior. Dissolved calcium was a significant predictor of plate and spine morph, while predator abundance was not. Stickleback latency to emerge from a refuge varied with morph, with populations with highly reduced plates and spines and high predation risk less bold. Our findings support strong directional selection in three‐spined stickleback evolution, driven by multiple selective agents.     

The role of calcium and predation on plate morph evolution in the three‐spined stickleback (Gasterosteus aculeatus)

While the genetic basis to plate morph evolution of the three‐spined stickleback (Gasterosteus aculeatus) is well described, the environmental variables that select for different plate and spine morphs are incompletely understood. Using replicate populations of three‐spined sticklebacks on North Uist, Scotland, we previously investigated the role of predation pressure and calcium limitation on the adaptive evolution of stickleback morphology and behavior. While dissolved calcium proved a significant predictor of plate and spine morph, predator abundance did not. Ecol. Evol., xxx, 2014 and xxx performed a comparable analysis to our own to address the same question. They failed to detect a significant effect of dissolved calcium on morphological evolution, but did establish a significant effect of predation; albeit in the opposite direction to their prediction.     

Daphnia predation on the amphibian chytrid fungus and its impacts on disease risk in tadpoles

Direct predation upon parasites has the potential to reduce infection in host populations. For example, the fungal parasite of amphibians, Batrachochytrium dendrobatidis (Bd), is commonly transmitted through a free‐swimming zoospore stage that may be vulnerable to predation. Potential predators of Bd include freshwater zooplankton that graze on organisms in the water column. We tested the ability of two species of freshwater crustacean (Daphnia magna and D. dentifera) to consume Bd and to reduce Bd density in water and infection in tadpoles. In a series of laboratory experiments, we allowed Daphnia to graze in water containing Bd while manipulating Daphnia densities, Daphnia species identity, grazing periods and concentrations of suspended algae (Ankistrodesmus falcatus). We then exposed tadpoles to the grazed water. We found that high densities of D. magna reduced the amount of Bd detected in water, leading to a reduction in the proportion of tadpoles that became infected. Daphnia dentifera, a smaller species of Daphnia, also reduced Bd in water samples, but did not have an effect on tadpole infection. We also found that algae affected Bd in complex ways. When Daphnia were absent, less Bd was detected in water and tadpole samples when concentrations of algae were higher, indicating a direct negative effect of algae on Bd. When Daphnia were present, however, the amount of Bd detected in water samples showed the opposite trend, with less Bd when densities of algae were lower. Our results indicate that Daphnia can reduce Bd levels in water and infection in tadpoles, but these effects vary with species, algal concentration, and Daphnia density. Therefore, the ability of predators to consume parasites and reduce infection is likely to vary depending on ecological context.     

Notes on the systematics and ecology of Daphnia pulex LEYDIG in Northern Canada

Daphnia populations from the region of Yellowknife (N. W. T.) on the Great Nave Lake, from Tuktoyaktuk (N. W. T.) on the coast of the Beaufort Sea east of the Mackenzie-Delta, and from Shingle Point (Y. T.) west of the Mackenzie-Delta were described. Population diagrams were given for the beginning of September 1974. The populations found at the above mentioned sites were compared with Daphnia pulex from Spitsbergen and the German island of Spiekeroog. Though some characteristics of Daphnia middendorffiana FISCHER occurred, mainly in individuals from very northern ponds, these were always found in individuals which also possessed some of the characteristics of Daphnia pulex. In every pond, specimens with characteristics of Daphnia middendorffiana lived together with other individuals resembling Daphnia pulex. Males were found in three out of the four visited ponds on the shore of the Arctic Sea, and were apparently Daphnia pulex. All the material sampled in northern Canada was regarded as belonging to Daphnia pulex LEYDIG 1860 emend. RICHARD 1896. There was evidence, that characteristics of Daphnia middendorffiana develop under the influence of environmental conditions, such as food supply. Daphnia magna STRAUS was found far north of the arctic circle.     

Morphological anti‐predator defences in the nine‐spined stickleback: constitutive,induced or both?

Environmental differences among populations are expected to lead to local adaptation, while spatial or temporal environmental variation within a population will favour evolution of phenotypic plasticity. As plasticity itself can be under selection, locally adapted populations can vary in levels of plasticity. Nine‐spined stickleback (Pungitius pungitius) originating from isolated ponds (low piscine predation risk, high competition) vs. lake and marine populations (high piscine predation risk, low competition) are known to be morphologically adapted to their respective environments. However, nothing is known about their ability to express phenotypic plasticity in morphology in response to perceived predation risk or food availability/competition. We studied predator‐induced phenotypic plasticity in body shape and armour of marine and pond nine‐spined stickleback in a factorial common garden experiment with two predator treatments (present vs. absent) and two feeding regimes (low vs. high). The predation treatment did not induce any morphological shifts in fish from either habitat or food regime. However, strong habitat‐dependent differences between populations as well as strong sexual dimorphism in both body shape and armour were found. The lack of predator‐induced plasticity in development of the defence traits (viz. body armour and body depth) suggests that morphological anti‐predator traits in nine‐spined stickleback are strictly constitutive, rather than inducible. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Predator-induced diapause in Daphnia magna may require two chemical cues

The production of diapausing eggs by Daphnia magna stimulated by fish exudates can be explained as an anti-predator defence ensuring genome protection in periods of high risk from fish predation. The combined effects on the induction of D. magna diapause of an “alarm” chemical originating from injured conspecific prey and fish kairomones were tested. The results of the experiment showed that the cues when present together promote both the production of ephippial eggs and male formation, indicating their role in the synchronization of the entire mode of Daphnia sexual reproduction. Ephippial eggs were only produced in the presence of both fish kairomone and conspecific alarm chemicals, while male offspring occurred in the treatments where both, one or none of the cues were present. However, production of males was the highest when both cues were provided. D. magna responded similarly to the tested cues whether or not the hypothetical alarm substance associated with predator odour came from Daphnia specimens actually eaten by fish or from crushed conspecific individuals. However, chemicals from crushed chironomid larvae combined with fish kairomones did not induce a similar response in D. magna. The relative advantage of utilization of alarm cues or predator kairomones in the induction of defence responses in prey organisms is discussed. Received: 8 June 1998 / Accepted: 11 January 1999     

Seasonal variation of egg size and number in a Daphnia pulex population

Seasonal variation of egg size and number was examined in a Daphnia pulex population inhabiting a vernal pond. In this population, size at maturity declines at midseason, probably as an adaptive response to size-selective predation by larvae of the salamander Ambystoma. The larger early season individuals produce more and larger eggs than the smaller late season individuals. Age at maturity does not vary between seasons. Laboratory experiments indicate that temperature may affect egg size, egg number and size at maturity. However, field data suggest that temperature accounts for only a small fraction of the total variation in egg size and number. Indirect measures of nutrition indicate that food limitation does not cause the seasonal decline in egg size and number. The seasonal change in reproductive traits is well correlated with changes in invertebrate and vertebrate predation. Examination of predator feeding preferences and their impact on Daphnia mortality indicate that variation of reproductive traits is most likely a complex adaptation to changing predation regimes.     

Predators can influence the host‐parasite dynamics of their prey via nonconsumptive effects

Ecological communities are partly structured by indirect interactions, where one species can indirectly affect another by altering its interactions with a third species. In the absence of direct predation, nonconsumptive effects of predators on prey have important implications for subsequent community interactions. To better understand these interactions, we used a Daphnia‐parasite‐predator cue system to evaluate if predation risk affects Daphnia responses to a parasite. We investigated the effects of predator cues on two aspects of host–parasite interactions (susceptibility to infection and infection intensity), and whether or not these effects differed between sexes. Our results show that changes in response to predator cues caused an increase in the prevalence and intensity of parasite infections in female predator‐exposed Daphnia. Importantly, the magnitude of infection risk depended on how long Daphnia were exposed to the cues. Additionally, heavily infected Daphnia that were constantly exposed to cues produced relatively more offspring. While males were ~5× less likely to become infected compared to females, we were unable to detect effects of predator cues on male Daphnia–parasite interactions. In sum, predators, prey, and their parasites can form complex subnetworks in food webs, necessitating a nuanced understanding of how nonconsumptive effects may mediate these interactions.     

Reproductive costs of Chaoborus-induced polymorphism in Daphnia pulex

Although the Chaoborus-induced spined morph of Daphnia pulex survives attacks by Chaoborus over twice as frequently as the typical morph, the spined morph is never found in the absence of Chaoborus. This implies that a disadvantage is associated with the spined morph in the absence of Chaoborus predation. The present study tested the hypothesis that the typical morph has a higher intrinsic rate of increase than the spined morph, by measuring several life history characteristics in controlled laboratory experiments at constant temperature and unlimited food.The results suggest that the spined morph of D. pulex takes longer to reach maturity, is smaller at maturity, but has similar egg number and egg size as the typical morph. These results are consistent with the hypothesis that the Chaoborus-induced spined morph is reproductively inferior to the typical morph.     

Local genetic adaptation generates latitude‐specific effects of warming on predator–prey interactions

Temperature effects on predator–prey interactions are fundamental to better understand the effects of global warming. Previous studies never considered local adaptation of both predators and prey at different latitudes, and ignored the novel population combinations of the same predator–prey species system that may arise because of northward dispersal. We set up a common garden warming experiment to study predator–prey interactions between Ischnura elegans damselfly predators and Daphnia magna zooplankton prey from three source latitudes spanning >1500 km. Damselfly foraging rates showed thermal plasticity and strong latitudinal differences consistent with adaptation to local time constraints. Relative survival was higher at 24 °C than at 20 °C in southern Daphnia and higher at 20 °C than at 24 °C, in northern Daphnia indicating local thermal adaptation of the Daphnia prey. Yet, this thermal advantage disappeared when they were confronted with the damselfly predators of the same latitude, reflecting also a signal of local thermal adaptation in the damselfly predators. Our results further suggest the invasion success of northward moving predators as well as prey to be latitude‐specific. We advocate the novel common garden experimental approach using predators and prey obtained from natural temperature gradients spanning the predicted temperature increase in the northern populations as a powerful approach to gain mechanistic insights into how community modules will be affected by global warming. It can be used as a space‐for‐time substitution to inform how predator–prey interaction may gradually evolve to long‐term warming.     

Two hundred years of a diverse Daphnia community in Lake Naivasha (Kenya): effects of natural and human-induced environmental changes

1. We used fossil diapausing eggs extracted from ²¹⁰Pb‐dated sediment cores to reconstruct historical changes in the Daphnia community of Lake Naivasha, a climate‐sensitive lake in Kenya which over the past 200 years has experienced a series of well‐documented natural and anthropogenic environmental changes. 2. Contiguous sampling and analysis of four cores yielded ephippial capsules of eight Daphnia species. Only two of these had been recorded previously in live collections from Lake Naivasha, and one species is new to science. The four more common species (Daphnia barbata, D. laevis, D. magna, and D. pulex) show striking differences in abundance patterns and population dynamics through time. Four other species (D. lumholtzi, D. curvirostris, D. longispina s.l., and Daphnia sp. nov. type Limuru.) appear to have been present only occasionally. Nevertheless, between 1895 and 1915 seven species of Daphnia inhabited Lake Naivasha simultaneously. 3. Despite considerable natural environmental change associated with climate‐driven lake‐level fluctuations, the Daphnia community of Lake Naivasha has been severely affected by human activities over the past century, especially the introduction of exotic fishes and water‐quality changes because of agricultural soil erosion. The recent reappearance of large‐bodied Daphnia species (D. magna, D. barbata, D. lumholtzi, Daphnia sp. nov. type Limuru) after 20–110 years of absence can be explained by their release from fish predation, following a dramatic increase in turbidity caused by excess clastic sediment input from eroded catchment soils. The small‐bodied species D. laevis has fared less well recently, presumably because the benefit of lowered predation pressure is counteracted by more pronounced negative effects of increased turbidity on this species and loss of submerged macrophyte beds which formerly served as predation refuge. 4. Our results suggest that, despite considerable environmental instability and the absence of specialised zooplanktivores, top‐down control of fish on large zooplankton is important in Lake Naivasha. Predation pressure from fish has led to clear‐cut shifts in local Daphnia species composition, but failed to drive the larger taxa to extinction.     

Predator-mediated genotypic shifts in a prey population: experimental evidence

We demonstrate the effect of fish predation on genotype frequencies in a laboratory population composed of two Daphnia magna clones, with historically contrasting exposures to fish predation. The two clones differed in their responsiveness to predation via differential avoidance/escape behavior. The clone which coexists with fish in nature is more responsive to the presence of a fish predator, while the clone not exposed to fish predation does not exhibit the defensive reaction. Fish caused a rapid (within 18 h) and significant shift in Daphnia clonal composition, from 1:1 to 8:1, in favor of the responsive clone. Genotype-specific defensive abilities (modus defendi) can contribute greatly to the phenomenon of genotype replacement under selective predation.     

Population dynamics of daphnia pulex and utilization by the rainbow trout (salmo gairdneri)

The effect of rainbow trout (Salmo gairdneri) predation on the population dynamics of the water flea,Daphnia pulex, was examined during 1976 and 1977 in Becker Lake, a small, shallow, productive reservoir in northeastern Arizona.Rainbow trout were size-biased feeders, utilizing daphnids which were 1.3 mm in size or larger. Trout predation uponDaphnia pulex occurred mainly during winter and early spring when their numbers were relatively low but their clutch size high, suggesting that trout selectedDaphnia on the basis of brood pigmentation. By far the greatest proportion ofDaphnia mortality was due to nonpredatory sources, since generally less than 6% could be attributed to trout predation.TheD. pulex population exhibited a bimodal seasonal abundance curve which was attributed to ephippial egg production and trout predation during the winter and poor food quality/quantity during the summer.This work was supported in part by the Arizona Department of Game and Fish. The authors wish to thank Jim Novy and Joe Stone of that department for their invaluable assistance in the field collecting segment of this study.     

Experiments on the Selection against Colour Oddity of Water Fleas by Three-Spined Sticklebacks

To determine whether odd individuals were more vulnerable to predation than the majority of a swarm, four types of choice experiments were conducted with three-spined sticklebacks (Gasterosteus aculeatus L.) as predators and water fleas (Daphnia magna) as prey, attempting to separate out the roles of oddity and conspicuousness. Odd-coloured water fleas were attacked more often by the predators than the majority, suggesting that oddity is effective apart from conspicuousness, which, however, also played a role.     

Behavioural responses to co-occurring threats of predation and ultraviolet radiation in Daphnia

1. Organisms in the wild are faced with multiple threats and a common response is a change in behaviour. To disentangle responses to several threats, we exposed two differently sized species of the freshwater invertebrate Daphnia to solar ultraviolet radiation (UVR) and predation from either moving pelagic or benthic ambush predators.
2. Using an advanced nanotechnology-based method, we tracked the three-dimensional movements of those mm-sized animals at the individual level. Each behavioural trial was performed both under conditions resembling night (no UVR) and day (UVR) and we examined patterns of the depth distribution and swimming speed by Daphnia across three treatments: no predator (control); bottom-dwelling damselfly (Calopteryx sp.); and fish (stickleback, Pungitius pungitius) predators. We also quantified the actual predation rate by the two predators on the two Daphnia species, Daphnia manga and Daphnia pulex.
3. We show that individual Daphnia are able to identify predators with different feeding habitats, rank multiple and simultaneously occurring risks and respond in accordance with the actual threat; complex responses that are generally associated with larger animals.
4. In a broader context, our results highlight and quantify how a cocktail of everyday threats is perceived and handled by invertebrates, which advances our understanding of species distribution in space and time, and thereby of population dynamics and ecosystem function in natural ecosystems.