Effect of the cyclopoid copepod Mesocyclops thermocyclopoides on the interactions between the predatory rotifer Asplanchna intermedia and its prey Brachionus calyciflorus and B. angularis

In many shallow, eutrophic subtropical ponds, brachionid rotifers are common prey of the predatory copepod Mesocyclops thermocyclopoides. The predatory rotifer Asplanchna intermedia, which is itself a potential prey of the copepod, also feeds preferentially on brachionids. We studied in the laboratory the population dynamics of two mutually competing prey species, Brachionus angularis and B. calyciflorus, in the presence of the two predators A. intermedia and M. thermocyclopoides. The experimental design included separate population dynamics studies with one prey–one predator, two prey–one predator, one prey–two predator, and two prey–two predator systems. These combinations were compared with controls, in which both the prey species (B. angularis and B. calyciflorus) were grown separately and in combination with each other. In the absence of any predator, B. angularis generally eliminated the larger B. calyciflorus. Selective predation by the copepod allowed B. calyciflorus to persist longer in competition with B. angularis. Feeding by M. thermocyclopoides on A. intermedia reduced the predation pressure on B. calyciflorus. However, given enough time, the cyclopoid copepod was able to eliminate both the brachionids as well as the predatory Asplanchna.     

Diet‐dependent intraguild predation between the predatory mites Neoseiulus californicus and Neoseiulus cucumeris

Based on the hypothesis that matching diets of intraguild (IG) predator and prey indicate strong food competition and thus intensify intraguild predation (IGP) as compared to non‐matching diets, we scrutinized diet‐dependent mutual IGP between the predatory mites Neoseiulus cucumeris and N. californicus. Both are natural enemies of herbivorous mites and insects and used in biological control of spider mites and thrips in various agricultural crops. Both are generalist predators that may also feed on plant‐derived substances such as pollen. Irrespective of diet (pollen or spider mites), N. cucumeris females had higher predation and oviposition rates and shorter attack latencies on IG prey than N. californicus. Predation rates on larvae were unaffected by diet but larvae from pollen‐fed mothers were a more profitable prey than those from spider‐mite fed mothers resulting in higher oviposition rates of IG predator females. Pollen‐fed protonymphs were earlier attacked by IG predator females than spider‐mite fed protonymphs. Spider mite‐fed N. californicus females attacked protonymphs earlier than did pollen‐fed N. californicus females. Overall, our study suggests that predator and prey diet may exert subtle influences on mutual IGP between bio‐control agents. Matching diets did not intensify IGP between N. californicus and N. cucumeris but predator and prey diets proximately influenced IGP through changes in behaviour and/or stoichiometry.     

Estimating predation risk in zooplankton communities: the importance of vertical overlap

In order to estimate predation risk in nature, two basic components of predation need to be quantified: prey vulnerability, and density risk. Prey vulnerability can be estimated from clearance rates obtained from enclosure experiments with and without predators. Density risk is a function of predator density, and the spatial and temporal overlap of the predator and prey populations. In the current study we examine the importance of the vertical component of overlap in making accurate estimates of predation risk from the invertebrate predator Mesocyclops edax on rotifer versus crustacean prey. The results indicate that assumptions of uniform predator and prey densities cause a significant underestimation of predation risk for many crustacean prey due to the coincident vertical migration of these prey with the predator. The assumption of uniformity is more reasonable for estimating predation risk for most rotifer prey.     

Influence of Predator Presence and Prey Density on Behavior and Growth of Damselfly Larvae (Ischnura elegans) (Odonata: Zygoptera)

Foraging behavior is often determined by the conflicting benefits of energy gain and the risk of mortality from predation or other causes. Theory predicts that animals should have lower activity levels when either the risk of predation or the availability of resources in the environment is high. We investigated the adjustment of the behavior of I. elegans larvae to predator presence (Anax imperator) and prey density (Daphnia sp.) and their interaction in a completely crossed factorial experiment in the lab and the effect of behavior on growth. The foraging activity of the I. elegans larvae was significantly reduced in the presence of a free-swimming predator but not a caged predator. Abdominal movements were significantly reduced at a low prey density. Growth was significantly reduced by the presence of a free swimming predator and low prey densities. These results provide evidence that these damselfly larvae adjust their behavior to the presence of predators to increase their survival at the expense of reduced growth and development.     

Further observations on developmental polymorphism and its evolution in the rotifer Brachionus calyciflorus

SUMMARY. In laboratory experiments, long, Asplanchna-induced posterolateral spines of Brachionus calyciflorus were very effective in preventing their capture and subsequent ingestion by the predator Asplanchna sieboldi but provided no protection against predation by Mesocyclops edax. Young, short-spined B. calyciflorus were always captured after attack by adult A. sieboldi and were ingested in about 12 seconds. Adult, short-spined forms were captured on c. 35% of occasions when attacked by this predator and were ingested in about 50 s. Young and adult long-spined forms were captured by this predator on c. 60% of occasions when attacked, but they both almost invariably escaped or were rejected 20–35 s after capture. Short- and long-spined B. calyciflorus adults were always captured when attacked by adult, female M. edax and were completely ingested in about 20 s and 30 s, respectively. Life-table experiments conducted with B. calyciflorus at several levels of the food organism, Aerobacter aerogenes, showed that neither the possession of long posterolateral spines nor the production of offspring with long posterolateral spines interfered with survivorship, fecundity, or reproductive potential. In the laboratory, the volumes of the amictic parthenogenetic eggs of B. calyciflorus cultured on Euglena or Aerobacter were significantly greater in individuals from populations maintaining long posterolateral spines than in comparable-sized individuals from populations maintaining short spines. Egg volume was generally independent of adult body length, but it was significantly greater in Brachionus fed on Euglena compared with Aerobacter. Possible reasons why B. calyciflorus does not produce long posterolateral spines in the absence of Asplanchna are discussed. Few organisms other than B. calyciflorus are known to develop novel defensive phenotypes in direct response to the presence of a predator. It is suggested that such developmental responses evolve only when two conditions apply: (1) the defensive structure is primarily effective against a single type of predator, and (2) the prey organism exhibiting the response has a short generation time.     

Population growth of Asplanchna sieboldi fed two Brachionus spp. (Rotifera) raised on green alga and baker's yeast

We conducted population growth experiments of A. sieboldi using Brachionus calyciflorus and Brachionus patulus as prey. The prey rotifers were mass cultured separately on Chlorella vulgaris, Saccharomyces cerevisiae or on their mixture. Data on population growth of A. sieboldi showed prey type and food density-related differences. At any given prey concentration, both B. calyciflorus and B. patulus raised on a mixture of alga and yeast, resulted in higher abundance of the predator than those raised solely on alga or yeast. The rate of population increase per day (r) of A. sieboldi increased with increasing prey density for both prey species. However, predators grown on B. patulus showed higher r values compared to those grown on B. calyciflorus.     

Predator‐Naïve Brown Trout (Salmo trutta) Show Antipredator Behaviours to Scent from an Introduced Piscivorous Mammalian Predator Fed Conspecifics

Introduced mammalian predators may pose a high risk for native and naïve prey populations, but little is known about how native fish species may recognize and respond to scents from introduced mammalian predators. We investigated the role of diet‐released chemical cues in facilitating predator recognition, hypothesizing that native brown trout (Salmo trutta) would exhibit antipredator behaviours to faeces scents from the introduced American mink (Neovision vison) fed conspecifics, but not to non‐trout diets. In treatments‐control and replicate stream tank experiments, brown trout showed significant antipredator responses to faeces scent from mink fed conspecifics, but not to faeces scent from mink fed a non‐trout diet (chicken), or the non‐predator food control, Eurasian beaver (Castor fiber). We conclude that native and naïve brown trout show relevant antipredator behaviours to an introduced mammalian predator, presumably based on diet‐released conspecific alarm cues and thereby estimate the predation risk.     

Predatory behaviour ofScatophaga stercoraria under laboratory conditions

Laboratory experiments tested the effectiveness ofScatophaga stercoraria (L.) as a predator of various insect species. Photophases, ages and gender of predators and effects of dung presence on predation rates were compared for either sex using counts of prey cadavers, behavioral observations and oviposition rates. Males consistently killed more prey than females, but longer photophases did not significantly increase daily predation rates. Nine to 10-day-old flies approaching sexual maturity achieved highest predation rates. Dung was found to affect predatory behaviour but not cumulative predation. Predatory behaviour of sexually mature males was more affected than female behaviour by the presence of dung, whereas sexually immatures flies showed no response to dung presence. Preconditioning ofS. stercoraria adults using 3 prey species of different sizes did not affect the prey chosen in subsequent random choice experiments. Larger prey species such asMusca domestica (L.) orDelia antiqua (Meigen) were preferred toDrosophila sp. by both sexes ofS. stercoraria. This species may be a useful predator for future integrated pest management techniques in vegetable crops or control of house flies in barns.      

Stylochus tauricus,a predator of the barnacle Balanus improvisus in the Black Sea

The flatworm Stylochus tauricus Jacubova has been found associated with the barnacle Balanus improvisus Darwin, on which it feeds. The predation rate (the number of barnacles eaten by one polyclad in a month) ranges between 5–10. Inside the empty shells of B. improvisus some egg-plates of S. tauricus were observed. Pelagic Götte's larvae aged 2–3 days possess 4 lobes while those aged 7–8 days have 5 lobes. Flatworms can prey on the young of another species Balanus eburneus Gould, whereas predation on the mussels Mytilus galloprovincialis Lam. is rare. There is a direct correlation between predator abundance and prey ingested.     

Density of an intraguild predator mediates feeding group size, intraguild egg predation, and intra- and interspecific competition

Intraguild predation (IGP) is common in most communities, but many aspects of density-dependent interactions of IG predators with IG prey are poorly resolved. Here, we examine how the density of an IG predator can affect feeding group size, IG egg predation, and the growth responses of IG prey. We used laboratory feeding trials and outdoor mesocosm experiments to study interactions between a social intraguild predator (larvae of the wood frog; Rana sylvatica) and its prey (spotted salamander; Ambystoma maculatum). Larvae of R. sylvatica could potentially affect A. maculatum by consuming shared larval food resources or by consuming eggs and hatchlings. However, successful egg predation requires group feeding by schooling tadpoles. We established from five to 1,190 hatchlings of R. sylvatica in mesocosms, then added either 20 A. maculatum hatchlings to study interspecific competition, or a single egg mass to examine IGP. Crowding strongly suppressed the growth of R. sylvatica, and IGP was restricted to the egg stage. In the larval competition experiment, growth of A. maculatum was inversely proportional to R. sylvatica density. In the predation experiment, embryonic mortality of A. maculatum was directly proportional to the initial density of R. sylvatica and the mean number of tadpoles foraging on egg masses. IGP on eggs reduced A. maculatum hatchling density, which accelerated larval growth. Surprisingly, the density of R. sylvatica had no overall effect on A. maculatum growth because release from intraspecific competition via egg predation was balanced by increased interspecific competition. Our results demonstrate that the density of a social IG predator can strongly influence the nature and intensity of interactions with a second guild member by simultaneously altering the intensity of IGP and intra- and interspecific competition.L . A. Burley and A. T. Moyer contributed equally to this work.     

Predation in a temporary pond with special attention to the trophic role of Triops cancriformis (Crustacea: Branchiopoda: Notostraca)

Temporary ponds are described as environments with a low predation pressure. Notostracans inhabit these types of ponds, and some populations acquire a high proportion of larger individuals, whose feeding behaviour is mainly predatory. The predation nature of Triops cancriformis is not widely accepted, because its diet is still partly controversial. We analysed the diet of one population of T. cancriformis in a Mediterranean temporary pond (Espolla pond, NE Iberian peninsula) to evaluate its predation behaviour. The gut content of the individuals bigger than 10 mm is mainly composed of detritus, plant fibres and microcrustaceans (cladocerans, ostracods and copepods). The prey number increases with the body size of individual T. cancriformis as previously described. Sex-biased predation was observed for one copepod prey, but not for the other. Predation pressure in that community was monitored along six hydroperiods as the percentage of predator biomass (not only T. cancriformis) in relation to the non-predator biomass. The proportion of predator biomass is high, and this contrasts with the low predation pressure expected for a temporary system with short hydroperiods.     

Differential vulnerability to predation and refuge use in competing larval salamanders

The aquatic larvae of two species of salamanders coexist as a result of differences in their competitive abilities: Ambystoma talpoideum is a superior aggressor, whereas A. maculatum is a superior forager. I examined the behavioral mechanisms that permit these species to coexist with their predatory congener, A. opacum. I asked whether the two prey species differ in their vulnerability to predation and in their use of structural and spatial refugia when under the risk of predation; such inter-specific variation may allow predation to contribute indirectly to prey coexistence. Larval A. maculatum (the superior forager) was more vulnerable to predation by A. opacum than was A. talpoideum, and only the latter species significantly increased its use of structural refugia (leaf litter) in the presence of the predator. In pond enclosures, both species of prey exhibited diel patterns of microhabitat use; significantly more larvae occupied shallow regions of enclosures during the day and migrated to deeper water (a spatial refugium) at night. However, when considered separately, neither (1) the presence of a predatory larval A. opacum nor (2) an increased density of intra- and interspecific competitors significantly altered this habitat shift for either prey species. Rather, diel microhabitat usage in A. talpoideum was significantly affected by an interaction between predator presence and competitor density. My results demonstrate the importance of refugia to coexistence in this predator-prey assemblage. Furthermore, predation by A. opacum may mediate prey competition; that is, preferential consumption of A. maculatum may reduce the competitive impact of this superior forager on A. talpoideum, thus enhancing their coexistence.     

Differential prey utilization by the generalist predator Coleomegilla maculata lengi according to prey size and species

Prey utilization by the generalist predator Coleomegilla maculata lengi Timb. (Coleoptera: Coccinellidae) of three sympatric lepidopterous species was quantified in relation with prey size (age) and prey species. Based on optimal foraging theory, we argued that costs associated with the utilization of small and large prey are higher than those of intermediate prey size. As a result, we expected a higher prey consumption rate on intermediate prey size leading to a convex prey consumption curve. Laboratory experiments showed that, within a given prey instar, Coleomegilla maculata lengi preyed more on Plutella xylostella (L.) compared to Artogeia rapae (L.) and Trichoplusia ni (Hübner). Generally, prey consumption rate by Coleomegilla maculata lengi on the three prey species decreased with increasing immature prey size (age). The predation efficacy of Coleomegilla maculata lengi adults and fourth instar larvae was higher compared to younger coccinellids (L2). Although, Coleomegilla maculata lengi showed a higher level of predation on smaller immature prey, we demonstrated that it is not the optimal size range for this predator. As predicted, prey weight consumption rate by Coleomegilla maculata lengi was higher at intermediate prey size leading to a convex prey utilization curve. The beneficial impact of Coleomegilla maculata lengi predation on the host plant was also estimated by using a Protection Index that considers the differential predation caused by the coccinellids and the relative importance of each pest species in terms of plant injury. Coleomegilla maculata lengi has a more significant beneficial impact when it preys on T. ni immatures.     

Pre-adult development of Phytoseiulus persimilis on diets of Tetranychus urticae and Tetranychus lintearius: implications for the biological control of Ulex europaeus

Predation by the phytoseiid mite, Phytoseiulus persimilis, is considered a major threat to the effectiveness of biological control of gorse, Ulex europaeus, using Tetranychus lintearius. To assess this threat and to determine if the impact of P. persimilis on T. lintearius populations is comparable to its impact on T. urticae populations, its development and predator : prey generation time ratios were assessed. The pre-adult mortality and development time of two populations of P. persimilis fed on two diets, T. urticae and T. lintearius, were determined at two temperatures, 14 and 24°C. There were no significant differences in either mortality or development time between the two populations of P. persimilis at these temperatures. There is therefore no evidence that the two tested populations of P. persimilis are behaving as different strains. Similarly, diet had no significant effect on either mortality or development time at these temperatures. At 14°C the mortality of P. persimilis was significantly higher and development was significantly longer than at 24°C. Using pre-adult development as a surrogate for generation times, predator : prey generation time ratios were calculated between P. persimilis and both T. urticae and T. lintearius using data from this and other studies. The predator : prey generation time ratios between P. persimilis and T. lintearius were lower than those between P. persimilis and T. urticae. These results indicate that the impact of P. persimilis on T. lintearius populations is likely to be comparable to its impact on T. urticae populations. This provides further evidence that predation by P. persimilis is having a deleterious effect on T. lintearius populations and therefore reducing its effectiveness as a biological control agent for gorse.     

Effects of chlorogenic acid-and tomatine-fed caterpillars on the behavior of an insect predator

If generalist insect predators are a selective force contributing to patterns of feeding specialization by insect herbivores, then predators should be deterred from eating allelochemical-fed prey. The attack and feeding behaviors of naive predators (Podisus maculiventris stinkbugs) reared on control caterpillars (Manduca sexta) fed plain diet were compared to experienced predators reared on caterpillars fed tomato allelochemicals. Tomatine-fed prey were found more quickly by both naive and tomatine-experienced predators, and chlorogenic acid-experienced predators were more stimulated to begin searching for prey. However, experienced predators were less likely to attack both chlorogenic acidfed and tomatine-fed caterpillars than were naive predators. These results indicate that allelochemical-fed prey were easier for predators to locate, but allelochemical-containing prey often deterred predation by experienced predtors.     

Epizoic mode of life in Brachionus rubens Ehrenberg as a deterrent against predation by Asplanchna intermedia Hudson

The planktonic rotifer Brachionus rubens has a propensity for an epizoic mode of life, and in nature is often found attached to cladocerans. In this way the rotifer avoids to a certain extent the adverse effects of interference competition with cladocerans. We test the hypothesis that the epizoic habit of B. rubens acts also as a deterrent against invertebrate predation. Using Asplanchna intermedia as predator, we followed the population growth patterns of B. rubens alone and in the presence of the host species Daphnia carinata and Ceriodaphnia rigaudi. In the absence of cladocerans, the prey was eliminated within three days, followed by extinction of the predator due to starvation. With D. carinata in the medium, the prey-predator system persisted much longer, with B. rubens reaching high population densities. With the smaller-sized C. rigaudi, allowing a significantly smaller fraction of B. rubens population to be epizoic, the system persisted longer than in the controls, but both the prey and predator eventually became extinct. We conclude that the epizoic habit of B. rubens, by acting as a prey refugium, helps a portion of the population to escape from predation, and facilitates its coexistence with Asplanchna intermedia.     

Influence of predator density on nonindependent effects of multiple predator species

Interactions between multiple predator species are frequent in natural communities and can have important implications for shared prey survival. Predator density may be an important component of these interactions between predator species, as the frequency of interactions between species is largely determined by species density. Here we experimentally examine the importance of predator density for interactions between predator species and subsequent impacts on prey. We show that aggressive interactions between the predatory shore crabs Carcinus maenas and Hemigrapsus sanguineus increased with predator density, yet did not increase as fast as negative interactions between conspecifics. At low density, interactions between conspecific and heterospecific predators had similar inhibitory impacts on predator function, whereas conspecific interference was greater than interference from heterospecifics at high predator density. Thus the impact of conspecific interference at high predator density was sufficient in itself that interactions with a second predator species had no additional impact on per capita predation. Spatial and temporal variability in predator density is a ubiquitous characteristic of natural systems that should be considered in studies of multiple predator species.     

The Effect of Refuge on Dermestes ater (Coleoptera: Dermestidae) Predation on Musca domestica (Diptera: Muscidae): Refuge for Prey or the Predator?

It is believed that habitat heterogeneity can change the extent of predator-prey interactions. Therefore, in this study we examined the effect of habitat heterogeneity (characterized here as an addition of refuge) on D. ater predation on M. domestica. Predation of D. ater on M. domestica larvae was carried out in experimental habitats with and without refuge, and examined at different prey densities. The number of prey eaten by beetles over 24 h of predator-prey interaction was recorded, and we investigated the strength of interaction between prey and predator in both experimental habitats by determining predator functional response. The mean number of prey eaten by beetles in the presence of refuge was significantly higher than in the absence of refuge. Females had greater weight gains than males. Logistic regression analyses revealed the type II functional response for both experimental habitats, even though data did not fit well into the random predator model. Results suggest that the addition of refuge in fact enhanced predation, as prey consumption increased in the presence of refuge. Predators kept in the presence of refuge also consumed more prey at high prey densities. Thus, we concluded that the addition of refuge was an important component mediating D. ater-M. domestica population interactions. Refuge actually acted as a refuge for predators from prey, since prey behaviors detrimental to predators were reduced in this case.     

Morphometric and demographic responses of brachionid prey (Brachionus calyciflorus Pallas and Plationus macracanthus (Daday)) in the presence of different densities of the predator Asplanchna brightwellii (Rotifera: Asplanchnidae)

We quantified the indirect effects of different densities (1, 4, 16 individuals per jar) of the predator Asplanchna brightwellii on the morphometry and demography of their prey Brachionus calyciflorus and Plationus macracanthus. Population growth and life table demography experiments were separately conducted for each of the two prey rotifer species while keeping A. brightwellii in indirect contact. The brachionids were fed the green alga Chlorella vulgaris at a density of 1 × 10⁶ cells ml⁻¹. As compared to those cultured in the absence of the predator, in the presence of A. brightwellii the postero-lateral spines of P. macracanthus increased by about 15 μm, while in B. calyciflorus the increase was more (>80 μm). For both the brachionid species, the peak population density significantly decreased in the presence of A. brightwellii. The reproductive variables viz., net reproductive rate, generation time, and the rate of population increase of B. calyciflorus and P. macracanthus were negatively affected in the presence of kairomones from A. brightwellii.     

Predation pressure on an imperfect Batesian micicry complex in the presence of alternative prey

Summary Differential predation pressure and the probability of predation on a Batesian mimicry complex and on alternative prey were estimatedin a field experiment. The mimicry complex was composed of a noxious model (Eleodes obscura (Say)) and a palatable mimic (Stenomorpha marginata (LeConte)). House crickets (Acheta domesticus) (Linn.) were used as alternative prey. The experiment was conducted for 23 nights in August and September to approximate the peak seasonal activity time period during which both models and mimics normally are exposed to predation while foraging and depositing eggs. Each night thirty prey in ratios of 16 models: 7 mimics: 7 crickets were exposed for 2.5 h to a suite of predators consisting of pallid bats (Antrozous pallidus), striped skunks (Mephitis mephitis) and ringtails (Bassariscus astutus) that had free access to the prey. The model-mimic ratio was similar to that found in nature. Predators obtained prey on 11 of the 23 nights and preferred the alternative prey (crickets) in proportions higher than was expected from a predation rate that was equal on all species of prey. Mimics were taken by predators at a rate proportional to their abundance, while models were taken at a rate considerably lower than their relative abundance. This suggests that at least some of the predators could distinguish between models and mimics and were willing to eat the mimics at higher frequencies than they were willing to eat the models. However, although the mimicry is not perfect with respect to the entire predator suite, the mimics still gain an advantage by resembling the models, compared to the predation levels on the alternate prey.