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.     

Vulnerability of Daphnia middendorffiana to Parabroteas sarsi predation: the role of the tail spine

The predaceous calanoid copepod Parabroteas sarsi and Daphniamiddendorffiana co-exist in South Andes ponds. Daphnia middendorffianajuveniles have a tail spine with negative allometric development.A series ofexperiments was carried out with tailed and tail-removedjuveniles of three different instars. In all cases tested, feedingrates were significantly higher on tail-removed prey. Directobservations showed a higher frequency of unsuccessful attackson spined juveniles when compared with tail-removed juveniles.The proportion of dorsal attacks also increased in spined juveniles.Prey total length was a much better predictor of feeding ratethan prey body size. Three groups of juveniles with equal totallength, but differentage, body size and biomass, showed no significantdifferences in their vulnerability to predation.     

Cyclomorphosis in Daphnia: an adaptation to avoid invertebrate predation

Seasonal morphological changes in three Daphnia species were followed over a two-year period in two lakes that differ in invertebrate and fish pressure. Whereas the morphology of D. hyalina, the biggest of the three species, varied little from season to season, D. cucullata, the smallest, exhibited the most pronounced seasonal changes in head height/carapace length ratio. The pattern of seasonal changes of body proportions was similar in all size classes and isometric growth of the head was reported for D. cucullata. Unlike the head, tail spine length/carapace length ratio almost did not vary seasonally. Strong negative allometry of tail spine growth was observed. These results are consistent with the hypothesis that helmets and tail spines provide protection against invertebrates in the two smallest, thus most endangered species.     

Edge effects in patchy seagrass landscapes: The role of predation in determining fish distribution

Predation is often described as an underlying mechanism to explain edge effects. We assessed the importance of predation in determining edge effects in seagrass using two approaches: a video survey to sample predators at small scales across seagrass edges, and a tethering experiment to determine if predation was an underlying mechanism causing edge effects. Underwater videos were placed at four positions: middle of seagrass patches; edge of seagrass; sand immediately adjacent to seagrass and sand distant from seagrass. Fish abundances and the time fish spent in view were measured. The main predatory fish (Australian salmon, Arripis spp.) spent more time over adjacent sand than other positions, while potential prey species (King George whiting, Sillaginodes punctata (Cuvier), recruits) were more common in the middle of seagrass patches. Other species, including the smooth toadfish, Tetractenos glaber (Freminville), and King George whiting adults, spent more time over sand adjacent to seagrass than distant sand, which may be related to feeding opportunities. King George whiting recruits and pipefish (Stigmatopora spp.) were tethered at each of the four positions. More whiting recruits were preyed upon at outer than inner seagrass patches, and survival time was greater in the middle of shallow seagrass patches than other positions. Relatively few pipefish were preyed upon, but of those that were, survival time was lower over sand adjacent to seagrass than at the seagrass edge or middle. Video footage revealed that salmon were the dominant predators of both tethered King George whiting recruits and pipefish. The distribution of predators and associated rate of predation can explain edge effects for some species (King George whiting) but other mechanisms, or combinations of mechanisms, are determining edge effects for other species (pipefish).     

Swimming behaviour of Daphnia clones: differentiation through predator infochemicals

We studied variation in small-scale swimming behavior (SSB)in four clones of Daphnia galeata (water flea) in response topredator infochemicals. The aim of this study was 3-fold. First,we tested for differences in SSB in Daphnia; second, we examinedthe potential of differences in SSB to explain survival probabilityunder predation; and third we tested the effect of differencesin SSB on survival under predation. Four treatments were applied:one kairomone-free control, one Chaoborus- (phantom midge) andone Perca (perch)-conditioned treatment, and a mixed treatmentcontaining both infochemicals. All of the three tested behavioralparameters (swimming speed, trajectory length and vertical distribution)were affected by the presence of the Chaoborus infochemical,and swimming speed and vertical distribution were also affectedby the presence of Perca infochemical. The effect of the treatmentwas interfered with by a clone effect: genetic differences werepronounced in all traits. These results illustrate that clonescan be responsive in only a subset of traits. The general theorythat clones are either responsive or non-responsive is not validfor SSB. The outcome of the predation trial confirms that adecrease in activity is a main factor in lowering Daphnia vulnerabilityto Chaoborus predation.     

Egg predation by copepods in Daphnia brood cavities

Both field observations and enclosure experiments show that juvenile copepods enter Daphnia brood cavities to feed on Daphnia eggs and embryos. The ability to perform such in vivo exploitation is reported for both cyclopoid and calanoid copepods. Copepodites of Acanthocyclops robustus (G.O. Sars) were found to eat eggs in brood cavities of D. magna, D. pulex and D. pulicaria in experimental enclosures rich in algae. Copepodites of Eudiaptomus gracilus (G.O. Sars) were found in brood cavities of D. hyalina in a mesotrophic lake. The copepods' intrusions into brood cavities caused dramatic declines in the clutch size of infested Daphnia, and this predation effect could easily be confused with the effect of severe food limitation.     

The role of pre- and post-dispersal seed predation in determining total seed loss

Most seed predation studies focus on either pre- or post-dispersal predation and may therefore underestimate the role of predation in regulating plant populations. We therefore estimated total seed predation of an invasive tree, mesquite (Leguminoseae: Prosopis spp.), by examining the entire seed pool from tree to seed bank. The spatio-temporal dynamics of total seed predation was examined by sampling across its Australian distribution and through time. The main predator was a host-specialist multivoltine beetle, Algarobius prosopis L. (Bruchidae), previously introduced as a biocontrol agent. Seed predation exceeded 20% in all seed stages (in pods on and off the tree, and seeds within woody endocarps (capsules) and free seeds on and in the ground) but was consistently highest in capsules on the ground (up to 90%). Pre-dispersal predation contributed little. Total seed predation rates were primarily determined by predation rates on the most persistent seed stage, in this case fallen pods if only pods are considered and seeds in capsules for the total seed pool. This pattern was consistent across the surveyed taxa, regions, years and seasonally. Predation rate was relatively unaffected by seed density, potentially because densities were always low (<150 seeds m⁻²). Average total seed predation within a region reached 55%, but we conclude that any population regulation of mesquite by seed predation will principally be through reduced seed bank persistence. Our results highlight the need to consider the entire seed pool, especially the often cryptic and overlooked long-lived stages, when determining seed loss to predation and its likely population consequences.     

Vulnerability of melanic Daphnia to brown trout predation

The co-existence of melanic Daphnia cf longispina and facultativelyplanktivorous brown trout is reported from a clear-water, alpinelake. This cooccurrence is uncommon, presumably due to the vulnerabilityof pigmented Daphnia to fish predation. Lake Bjornesfjorden(Norway) provided an opportunity to test this assumption. About20% of the fish caught in gill nets had fed on Daphnia. Thetrout exerted a marked selection for large-sized Daphnia prey,and a very strong selection for pigmented individuals relativeto transparent ones The persistence of a pigmented Daphnia populationprobably relies on limited recruitment and a low stock of thepredator, and the availability of more favourable benthic preyorganisms. ⁴Present address: Rådgivende Biologer A/S, Bredsgården,Bryggen, N-5003 Bergen, Norway     

Optimising survival under predation: chemical cues modify curvature in Daphnia galeata

Morphological responses to the presence of predator info-chemicals havebeen described for many Daphnia (Cladocera) species, butD. galeata is generally considered to exhibit almost nomorphological changes that could increase its fitness under predation.Therefore, the aim of our study was to examine the nature and magnitude ofmorphological responses of D. galeata to their predatorsindetail and assess their potential role in decreasing the predation threat. Twoclones of Daphnia were exposed to predator info-chemicals(kairomones) from perch, a fish (Perca), and a phantommidge larvae (Chaoborus) an invertebrate, and a kairomone mixture fromboththese organisms. Laboratory life-table experiments were carried out and fiveparameters characterising the body shape of the daphnids were measured: helmetlength, head- and carapace width, eye diameter and body size. The last-namedthree parameters did not differ significantly between the clones or thetreatments. The differences found between the clones were significant for headwidth and helmet length, but only in combination with the treatment effects.Ourresults on genotype-dependent phenotypic plasticity indicated that, althoughphenotypic plasticity is present, the clonal composition of aDaphnia population can be altered by selection on themorphotype. This potential for a change in clonal frequencies is given by thedifferences measured between the two clones in head width and helmet length,altering the curvature of the Daphina body in response tokairomone presence.     

Chaoborus predation and delayed reproduction in Daphnia: a demographic modeling approach

I develop a demographic model that examines the impact of Chaoborus predation on the population dynamics and life history of Daphnia. Predation effects are determined through analysis of the various components of the predator-prey interaction (encounter, attack, strike efficiency), and are integrated into a stage-classified matrix population model. The model is parameterized with data from interactions between D. pulex and fourth-instar C. americanus. I test this model with two laboratory experiments that examine population growth of D. pulex under the influence of five different levels of Chaoborus predation. With the exception of a single predation treatment in each experiment, the model accurately predicted the observed reduction in Daphnia numbers with increasing Chaoborus predation. I then use this model to investigate the evolution of delayed reproduction in D. pulex that are exposed to Chaoborus. I ask whether delayed reproduction may evolve in Daphnia that are subjected to Chaoborus predation as a trade-off for the benefits of larger body size. The model predicts that the effectiveness of such a life history trade-off depends on the relative sizes of predator and prey. In some interactions between Chaoborus and Daphnia, increasing Daphnia body length by as little as 5% from base growth trajectories sufficiently increases fitness (by reducing vulnerability to Chaoborus predation) to compensate for the cost of delayed reproduction. In other interactions, however, increased body length provides no benefit to Daphnia (and may even reduce fitness), and selection would act against the evolution of delayed reproduction. This revised version was published online in July 2006 with corrections to the Cover Date.     

Chaoborus predation and the function of phenotypic variation in Daphnia

To investigate the role of helmet formation in defense against predation, laboratory experiments were used to analyze the effects of morphological changes in Daphnia on susceptibility to Chaoborus predation. Behavioral observations of Chaoborus preying on helmeted and non-helmeted Daphnia suggest pre-contact advantages for helmeted prey but post-contact advantages for non-helmeted prey. Helmeted Daphnia are better at evading capture by Chaoborus but may also be more easily handled by the predator. Swimming behavior of the prey, which is influenced by the presence of a tailspine, may affect Chaoborus strike distance. These results re-emphasize the potential hydromechanical importance of body shape changes in defense against predation.     

Impact of larval Dorosoma predation on Daphnia parvula dynamics

• 1 In reservoirs of the south-eastern United States, Daphnia typically decline in abundance during the summer months. Current understanding suggests that adult shad (Dorosoma spp.), the dominant planktivore in these reservoirs, plays a limited role in Daphnia midsummer declines. The impact of shad larvae is beginning to be documented.
• 2 To examine the impact of larval shad on Daphnia seasonal dynamics, larval dynamics were studied over two seasons and stomach contents were analysed during one season.
• 3 Initial Daphnia panwla declines correlated with abundance of shad larvae. During the decline, shad stomach contents consisted of 97—100%D. parvula. After D. parvula declined, Bosmina and Diaphanosoma accounted for up to 97% of larval diets. When shad diets included other cladocerans, D. parvula abundance increased in the reservoir.
• 4 When this predation pattern was incorporated into a predictive model, larval shad predation accounted for an initial, temporary decline in D. parvula abundance. However, when other crustaceans were incorporated into the shad diet, D. parvula populations increased.
• 5 These results suggest that larval shad have a temporary, strong influence on the structure and abundance of D. parvula populations in south-eastern reservoirs of the United States.

     

捕食风险对高原鼠兔行为的影响

我们在野外条件下利用赤狐的粪尿增加高原鼠兔的捕食风险 ,采用目标动物抽样法对高原鼠兔的 5种行为进行观察 ,分析天敌动物气味增加与天敌动物数量增加对高原鼠兔行为影响的差异 ,探讨高原鼠兔对捕食风险的权衡能力及面对不同风险的行为决策。 2 0 0 1年 ,在捕食风险处理样方中观察雄性成体 1 8只 ,雌性成体2 5只 ,雄性幼体 35只和雌性幼体 4 2只 ,在对照样方中观察雄性成体 1 4只 ,雌性成体 1 5只 ,雄性幼体 2 2只和雌性幼体 1 1只 ;2 0 0 2年 ,在捕食风险处理样方中观察雄性成体 7只 ,雌性成体 1 2只 ,在对照样方中观察雄性成体 8只 ,雌性成体 1 5只。研究结果表明 :增加赤狐的气味后 ,高原鼠兔通过改变行为策略以适应捕食风险的增加。当气味源刚放入后 ,与对照样方比较 ,高原鼠兔明显增加了观察和鸣叫的频次 ,相应减少了取食的频次 ,且随着时间的推移 ,高原鼠兔并未对气味产生适应性。当天敌动物的数量增加后 ,赤狐气味的增加对高原鼠兔行为的影响较小 ,在捕食风险超出高原鼠兔的耐受范围时 ,高原鼠兔扩散。同时 ,高原鼠兔的行为在雌雄之间、成体与幼体之间没有明显的不同 ,雌雄个体、成幼体均采用相同的行为策略减小捕食风险。以上结果表明 :捕食风险明显影响高原鼠兔的行为 ,高原鼠兔能够权衡捕食     

Effects of predation risk on mating behavior of the Kanzawa spider mite

In the Kanzawa spider mite, Tetranychus kanzawai, adult males locate pre-reproductive quiescent females and engage in precopulatory mate guarding. We found that re-reproductive quiescent females preferred to be near veins, rather than other leaf parts, and moreover, adult males spent more time along the vein than on other parts. Consequently, T. kanzawai males found more quiescent females along veins than those on other parts. However, the predatory mite Neoseiulus womersleyi also found more quiescent T. kanzawai females along veins than those on other parts. Moreover, N. womersleyi found more guarding males than solitary males of T. kanzawai. Thus, we experimentally examined the effects of predation risk on the mating behavior of T. kanzawai. The presence of N. womersleyi reduced T. kanzawai female preference for vein vicinity as a quiescent site. Although the predation risk of guarding T. kanzawai males was lower than that of solitary males after detection by predators, the presence of N. womersleyi also reduced the proportion of guarding T. kanzawai males. These results suggest that the possible benefits of preferring vein vicinity as quiescent sites by T. kanzawai females is outweighed by predation risk in the presence of predators, and that the risk of detection by predators would be more important for T. kanzawai males than the risk of being preyed upon.     

Predicting extinctions: interspecific competition,predation and population variability in experimental Daphnia populations

We examine how interspecific competition and two types of size-selective predation affect population density, variability and persistence in laboratory cultures of two species of Daphnia, D. magna and D. longispina. When both species were analysed together, and for D. longispina alone, there were weak negative relationships between mean population density and population variability. Interspecific competition resulted in lower population densities and higher population variability. Extinct populations had lower densities and were also more variable than persisting ones. There was still an effect of population variability on extinction probability after the effect of density on population variability had been accounted for. Hence, the effects of population density and variability on population persistence were partly independent of each other. The effects of size-selective predation on population persistence were more species-specific and not directly related to density or variability. Since the effects of species interactions on persistence were large, we suggest that it is likely that population vulnerability analyses not incorporating effects of interspecific interactions are often misleading.     

The effects of predation risk from crab spiders on bee foraging behavior

Recent studies have suggested that top–down effects ofpredation on plant–pollinator interactions may not be,as previously thought, rare and/or weak. In this paper, we explorethe effects of crab spiders (Araneae: Thomisidae) on the behaviorof 2 species of bee (Hymenoptera: Apidae) foraging for nectarand pollen on 3 different plant species in central Portugal.In 2 experiments, we found that the eusocial bee Apis melliferawas significantly less likely to inspect and accept a floweror inflorescence if it harbored a spider. In contrast, we foundno such effects of spiders on the behavior of the solitary beeEucera notata. Further experiments showed that the effects ofenvironmental cues associated with predators on flower visitationby A. mellifera were detectable even when no spider was presentat the moment a flower was encountered. Such indirect effectswere only identified, however, in bees foraging on 1 of 2 plantspecies studied. In a final experiment, A. mellifera was shownto respond negatively to the presence of the corpses of conspecificsglued to flowers. This suggests that prey corpses left exposedon petals or bracts by spiders provide an obvious cue that beescan use to avoid predators. These results add to a growing bodyof evidence that plant–pollinator interactions are notimmune to the effects of predation and suggest that the strengthof such effects vary both between and within species.     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

Assessment of local predation risk: the role of subthreshold concentrations of chemical alarm cues

The ability to accurately assess local predation risk is criticalto prey individuals, as it allows them to maximize threat-sensitivetrade-offs between predator avoidance and other fitness relatedactivities. A wide range of taxonomically diverse prey (includingmany freshwater fishes) relies on chemical alarm cues (alarmpheromones) as their primary information source for local riskassessment. However, the value of chemical alarm cues has beenquestioned due to the availability of additional sensory inputs(i.e., visual cues) and the lack of an overt antipredator responseunder conditions of low perceived risk. In this paper, we testthe hypothesis that chemical alarm cues at concentrations belowthe point at which they elicit an overt behavioral responsefunction to increase vigilance towards other sensory modalities(i.e., visual alarm cues). Shoals of glowlight tetras (Hemigrammuserythrozonus) exposed to the subthreshold concentration of hypoxanthine-3-N-oxide(the putative Ostariophysan alarm pheromone) did not exhibitan overt antipredator response in the absence of secondary visualcues (not different than the distilled water control). However,when exposed to the sight of a visually alarmed conspecific,they significantly increased the intensity of their antipredatorresponse (not different from shoals exposed to the suprathresholdalarm cue). This study demonstrates that prey may benefit fromresponding to low concentration alarm cues by increasing vigilancetowards secondary cues during local risk assessment, even inthe absence of an overt behavioral response. By increasing vigilancetowards secondary risk assessment cues in the presence of alow risk chemical cue, individuals are likely able to maximizethe threat-sensitive trade-offs between predator avoidance andother fitness related activities.     

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.