Walking,feeding and intraspecific interaction of larvae of Metaseiulus occidentalis,Typhlodromus pyri,Neoseiulus fallacis and Amblyseius andersoni held with and without eggs of Tetranychus urticae

Larvae of Metaseiulus occidentalis (Nesbitt), Typhlodromus pyri Scheuten, Neoseiulus fallacis (Garman) and Amblyseius andersoni Chant exhibited different activity levels when held on apple leaf or on tile arenas and given or not given eggs of Tetranychus urticae Koch and water (tiles only). M. occidentalis larvae held without prey exhibited high levels of walking (includes searching) during 24 hours of evaluation, whereas M. occidentalis larvae held with prey fed quickly and then became less active. Fed larvae of M. occidentalis were less active on leaves than tile. Larvae of T. pyri on leaves had a very low frequency of walking, almost never fed and quickly assumed a resting position during development. While much less active than M. occidentalis, fed and unfed T. pyri larvae walked more on tiles than leaves before resting. Larvae of N. fallacis and A. andersoni fed at low rates and were similarly active on tiles and leaves. Free water increased walking by M. occidentalis, A. andersoni and T. pyri on tile but not N. fallacis. M. occidentalis larvae interacted 5–7 times more often than larvae of the other three species. Cannibalism or scavenging was rarely seen and then only for M. occidentalis larvae. Larvae of all four mites walked, fed and interacted much more in the first 12 hours than the second 12 hours of tests, except unfed M. occidentalis. Unfed M. occidentalis larvae did not molt to protonymphs but unfed larvae of the other three species did. Unfed and fed protonymphs of all four species walked more at 4 hours after molting than larvae at 12–24 hours. Unfed and fed protonymphs of T. pyri or A. andersoni had similar walking frequencies, but unfed protonymphs of N. fallacis were more active than fed ones. Trends in larval activities are discussed relative to the life history of each species.     

Geographic variation of diapause intensity in the spider mite Tetranychus urticae

Abstract. Eight strains of the spider mite Tetranychus urticae, originating from different localities in western and central Europe, with latitudes ranging from 40.5 to 60^oN, displayed marked differences in the period of chilling at 4^oC required for diapause termination under a diapause-maintaining short-day photoperiodic regime at 19^oC, to which the mites were transferred after the cold period. The higher the latitude from which the strains originated the longer was the period of chilling required for diapause termination, suggesting the presence of a gradient in diapause intensity, diapause being deeper the more northern the origin of the strains. Two strains originating from higher altitudes appeared to have a much deeper diapause than expected from their latitudinal origin. In addition, these two mountain strains showed mutual differences in diapause intensity, notwithstanding the fact that they originated from similar latitudes and altitudes; local climatic conditions probably act as strong selective forces with regard to diapause depth. All strains appeared to be sensitive to photoperiod during the period of diapause development. Diapause was quickly completed by a long-day photoperiod (LD 17:7 h), but was maintained by a short-day photoperiod (LD 10:14h). However, even under the latter regime sensitivity to photoperiod gradually diminished and eventually disappeared, thus leading to ‘spontaneous’ termination of diapause. The length of the period of diapause development, as measured by the sensitivity to photoperiod of diapausing mites, varied between strains; it was shorter in the southern strains and longer in the northern strains. The results indicate great variation in diapause intensity between strains, which is probably genetically determined and may have adaptive significance for this widespread species. When young females which had just entered diapause were kept for ever longer periods of time under the diapause inducing short-day regime at which they had been reared, before being transferred to the cold room, the duration of the period of chilling required for diapause termination was found to decrease proportionally in all three strains tested. These results suggest that intensification of diapause does not occur in T. urticae; diapause intensity seems to be highest at the beginning of diapause and to diminish gradually during diapause development.     

Acoustic stimuli from predators trigger behavioural responses in aggregate caterpillars

Prey detect their predators through predator signals and cues and, consequently, respond with anti‐predatory behaviours to inhibit the action of their aggressors. Lepidopterans can intercept signals emitted by predators and may defend themselves through chemical, morphological or behavioural responses. In this study, we investigated the effect of acoustic stimuli of different predators on defensive behaviour of gregarious caterpillars. Our results demonstrated that Hylesia nigricans (Lepidoptera, Saturniidae) caterpillars alter their behaviour (i.e. abruptly raising the head) in response to the acoustic stimulus of the predators (i.e. predation risk signals from birds and wasps). The magnitude of this response depended on predator identity and caterpillar body size. Larger caterpillars responded more strongly to predatory stimuli than smaller caterpillars. However, regardless of the size of the caterpillars, they responded more strongly to the stimuli of wasps. In addition, we identified that H. nigricans caterpillars emit ultrasonic noise after detecting the stimuli of the predators – this noise seems to function as an alert about the risk of predation during the early stages of development (second and fifth instars). The duration of ultrasonic emission (i.e. milliseconds) increases with the number of repetitions of the stimuli (i.e. wing‐beat sounds of the wasps and insectivorous birds). These results provide novel information about predation risk in interactions among caterpillars and their predators, and indicate possible communication among invertebrates mediated by the risk of predation.     

Nonconsumptive effects in a multiple predator system reduce the foraging efficiency of a keystone predator

Many studies have demonstrated that the nonconsumptive effect (NCE) of predators on prey traits can alter prey demographics in ways that are just as strong as the consumptive effect (CE) of predators. Less well studied, however, is how the CE and NCE of multiple predator species can interact to influence the combined effect of multiple predators on prey mortality. We examined the extent to which the NCE of one predator altered the CE of another predator on a shared prey and evaluated whether we can better predict the combined impact of multiple predators on prey when accounting for this influence. We conducted a set of experiments with larval dragonflies, adult newts (a known keystone predator), and their tadpole prey. We quantified the CE and NCE of each predator, the extent to which NCEs from one predator alters the CE of the second predator, and the combined effect of both predators on prey mortality. We then compared the combined effect of both predators on prey mortality to four predictive models. Dragonflies caused more tadpoles to hide under leaf litter (a NCE), where newts spend less time foraging, which reduced the foraging success (CE) of newts. Newts altered tadpole behavior but not in a way that altered the foraging success of dragonflies. Our study suggests that we can better predict the combined effect of multiple predators on prey when we incorporate the influence of interactions between the CE and NCE of multiple predators into a predictive model. In our case, the threat of predation to prey by one predator reduced the foraging efficiency of a keystone predator. Consequently, the ability of a predator to fill a keystone role could be compromised by the presence of other predators.     

Effects of mating status and age on dispersal behavior in the twospotted spider mite, Tetranychus urticae in response to fenvalerate-treated leaf surfaces

Effects of variation in age and mating status on dispersal of the twospotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae), from fenvalerate-treated leaf surfaces were investigated. Unmated and mated female mites, 1 to 6 days in age, were scored for walkoff and spindown dispersal response on bean leaves treated with a sub-lethal, residual dose of fenvalerate. Variation in mite age did not have a pronounced effect on dispersal response. Significant differences in dispersal response due to mating type were observed, yet the mean differences in dispersal response between unmated and mated females was generally small. Our data suggest that the current practice of using mated female mites of unknown age when testing for among-population differences in dispersal response, in most cases, provides an adequate estimate of population response when differences among treatments in dispersal response are large. However, when testing for within-population differences in dispersal response, small treatment effects may be more easily measured using a single age class of mites, because this could decrease within-treatment variance.     

Linking anti‐predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore

Ecological theory predicts that the diffuse risk cues generated by wide‐ranging, active predators should induce prey behavioural responses but not major, population‐ or community‐level consequences. We evaluated the non‐consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high‐risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20‐fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.     

Diet and feeding selectivity of a benthivorous fish in streams: responses to the effects of urbanization

This study evaluated the diet and feeding selectivity of the catfish Imparfinis mirini in streams with different degrees of urbanization and the effect of rainfall on the availability of prey. The diet was based especially on Chironomidae and Trichoptera. Significant spatial differences in diet were found between the streams; the diet of the fish was similar in the rural and peri‐urban streams, and differed from that in the urban stream. Seasonality was an unimportant factor affecting the species' diet, which did not differ significantly between the rainy and dry periods in any of the streams. Fish from the urban stream fed more according to what was available in the environment, while fish from the peri‐urban and rural streams showed higher degrees of selectivity. The results indicate that environmental conditions influenced the diet and prey selection of this species in response to the differences in diversity and abundance of the food organisms. They indicate that trophic studies of benthivorous fishes, such as I. mirini, may represent an alternative way to assess human effects on streams.     

Acaricidal and stimulatory effects of insecticides on Tetranychus urticae Koch (Acari: Tetranychidae) in cotton

Possible stimulatory or acaricidal effects of thiodicarb, dimethoate and endosulfan on the two-spotted spider mite, Tetranychus urticae Koch, were tested in a glasshouse. Population development of T . urticae was monitored, with the expectation that any stimulatory or acaricidal effects would be reflected in different population growth relative to untreated plants. Acaricidal activity of the three insecticides was also investigated using a standard bioassay technique. Population growth on thiodicarb- or dimethoate-treated plants was no different to that on untreated plants but was slower on plants treated with endosulfan. Bioassays showed that endosulfan was moderately acaricidal, that T . urticae was highly resistant to dimethoate and that thiodicarb was of very low acaricidal activity. No evidence of stimulatory effects of dimethoate or thiodicarb was found. The data suggest that stimulation is an unlikely explanation for outbreaks of T . urticae in cotton and that endosulfan sprayed for control of Helicoverpa spp. may delay the development of mite outbreaks.     

Prey stage preference and feeding behaviour of Oligota kashmirica benefica (Coleoptera: Staphylinidae), an insect predator of the spider mite Tetranychus urticae (Acari: Tetranychidae)

We studied the prey stage preference and feeding behaviour of the first to third instar larvae and adult females of Oligota kashmirica benefica Naomi (Coleoptera: Staphylinidae), a predator of the spider mite Tetranychus urticae Koch (red form) (Acari: Tetranychidae), on leaves of the kudzu vine (Pueraria lobata (Wild.) Ohwi (Leguminosae)) under laboratory conditions. The number of mites eaten increased with the growth of predator larvae. Third instar larvae preyed on all stages of spider mite, whereas first instar larvae preyed mainly on immobile stages (eggs and quiescent stages). The predator larvae showed two types of foraging behaviour (active searching and ambush behaviour) when targeting the mobile stages (larval, nymph and adult stages of prey). Although no difference was found in the number of prey consumed by adult females and third instar larvae of the predator, the adult females mainly attacked and consumed the immobile stages.     

Early exposure to predation risk carries over metamorphosis in two distantly related freshwater insects

1. Predation and competition play a central role in ecological communities, and it is increasingly recognised that animals use early warning cues to reduce the impact of these antagonistic interactions. 2. Strategies to avoid risk can occur during embryo development through plasticity in egg hatching time. This strategy, and its associated costs and carryover effects on adults are little understood in insects. In this study, these are explored in two distantly related freshwater insects: the damselfly Ischnura elegans and the mosquito Aedes albopictus. 3. As predicted, damselfly eggs hatched earlier in response to larval predators cues, a treatment that also affected adult size. Risk cues did not affect mosquito egg hatching time, but they did affect larval development time in a sex‐dependent manner. 4. The results suggest that responses aimed at avoiding risks can be triggered during the egg stage, and although they can vary dramatically among species, they are likely to be widespread in insects. Early warning responses can be particularly important to understand the ecology of aquatic insects, some of them global vectors of human diseases.     

Temporal variation in behavioral responses to dietary cues from a gape‐limited predator in tadpole prey: A test of the phylogenetic relatedness hypothesis

The ability of prey to recognize and adequately respond to predators determines their survival. Predator‐borne, post‐digestion dietary cues represent essential information for prey about the identity and the level of risk posed by predators. The phylogenetic relatedness hypothesis posits that prey should respond strongly to dietary cues from closely related heterospecifics but respond weakly to such cues from distantly related prey, following a hierarchical pattern. While such responses have mostly been observed in prey at their first encounter with predators, whether prey maintain such hierarchical levels of investment through time remains unclear. We investigated this question by exposing Rhacophorus arboreus tadpoles to the non‐consumptive effect of gape‐limited newt predators Cynops pyrrhogaster that were fed one of five prey diets across a gradient of phylogenetic relatedness: frog tadpoles (Rhacophorus arboreus, Rhacophorus schlegelii, Pelophylax nigromaculatus, and Hyla japonica) and medaka fish (Oryzias latipes). Predators’ diet, time, and their interaction significantly influenced tadpole activity level. We found support for the phylogenetic relatedness hypothesis: Investments in defense were stronger to cues from tadpole diets than to cues from fish diet. However, such a hierarchical response was recorded only in the first four days following predator exposure, then gradually disappear by day 8 on which the tadpoles exhibited similar activity level across all predator treatments. The findings suggest that, at least under the threat of gape‐limited predators, prey use phylogenetic information to evaluate risk and appropriately invest in defense during early encounters with predators; however, energy requirements may prevent prey from maintaining a high level of defense over long exposure to predation risk.     

Overwintering and effects of autumn habitat manipulation and carbofuran on Neoseiulus fallacis and Tetranychus urticae in peppermint

The percentages of Neoseiulus fallacis (Garman) in four subhabitats of peppermint fields in November were 37% on dead leaves, 45% in debris below dead leaves, 3% in hollow stems and 15% on live foliage. In December, the percentages were 36, 39, 7 and 18%, respectively, for these same sites. The values were 40, 37, 18 and 5% and 6, 80, 14 and 0% for January and February, respectively. By April, 95% of all predators were on new foliage feeding on Tetranychus urticae Koch, while the rest were still in debris. Augmenting plots with debris increased overwintering survival of N. fallacis compared to the natural condition while removal of debris decreased it. An autumn carbofuran (Furadan 4F) treatment completely eliminated overwintering N. fallacis, thus leading to outbreaks of T. urticae the following spring. The results of these overwintering studies are discussed in relation to improved biological control and spider mite management.     

Wise flies: a pre‐dispersal seed predator prefers hermaphrodites over females in the gynodioecious Polemonium foliosissimum

1. Oviposition choices can profoundly affect offspring performance. Oviposition decisions of the dipteran pre‐dispersal seed predator, Hylemya sp. (Diptera: Anthomyiidae), when choosing among sex morphs of their host‐plant—Polemonium foliosissimum Hook—were evaluated. Polemonium foliosissimum is gynodioecious, with female and hermaphrodite sex morphs that differ in flower size. 2. It was asked: Do female flies preferentially oviposit on hermaphrodite plants and, if so, are oviposition choices correlated with flower size? Is larval survivorship higher on hermaphrodite plants and, if so, is larval success correlated with flower size? Can the differences in oviposition and/or larval success be explained by slight differences in flowering phenology between the sexes? 3. Hermaphrodite flowers received 45% more Hylemya eggs than females. Although hermaphrodites had larger petals and sepals than females, egg loads were not correlated with petal or sepal size. Larval survival was 30% greater on hermaphrodites than females and higher on plants with larger sepals. However, the difference in sepal area between genders did not fully explain larval survival differences. Egg numbers declined over weeks, but differences in egg loads between the sex morphs were not attributable to a slight phenological delay of females. Larval survival declined over the season; however, larval survival differences between sex morphs were consistent. 4. Hylemya preferentially oviposited on hermaphrodites where their larvae had a significantly greater survival rate compared with females. The present results add to the knowledge that intra‐specific choices have consequences for phytophagous insects and that the relationship between antagonists and their gynodioecious host plants is complex.     

Development of species‐specific primers for rapid diagnosis of Tetranychus urticae,T. kanzawai,T. phaselus and T. truncatus (Acari: Tetranychidae)

Species diagnosis is of the utmost importance to both pest management and plant quarantine services. Because of difficulties in the morphological diagnosis of spider mites, molecular techniques are of great value to rapidly and accurately diagnose closely related species. We examined four species of genus Tetranychus (the green and red forms of T. urticae, and T. kanzawai, T. phaselus and T. truncatus), which are found in Korea and are of significance to plant quarantine services. DNA samples isolated from a single egg, larva or adult weighed 64–188 ng. We designed species‐specific primers by performing sequence alignment for 107 sequences of the ribosomal internal transcribed spacer 2 (ITS2) region, which we obtained from GenBank, and sequences generated in this study. Specific nucleotides of each species were selected for designing primers specific for each species. Each species‐specific primer pair, when used to perform PCR analyses, detected only the species from which it originated. However, a T. urticae‐specific primer pair did not discriminate between the green and red forms of this species. These species‐specific primers can be applied in practice for the rapid and accurate diagnosis of spider mite species in plant quarantine and in agricultural fields.     

Artificial selection for timing of dispersal in predatory mites yields lines that differ in prey exploitation strategies

Dispersal is the main determinant of the dynamics and persistence of predator–prey metapopulations. When defining dispersal as a predator exploitation strategy, theory predicts the existence of a continuum of strategies: from some dispersal throughout the predator–prey interaction (the Milker strategy) to dispersal only after the prey had been exterminated (the Killer strategy). These dispersal strategies relate to differences in prey exploitation at the population level, with more dispersal leading to longer predator–prey interaction times and higher cumulative numbers of dispersing predators. In the predatory mite Phytoseiulus persimilis, empirical studies have shown genetic variation for prey exploitation as well as for the timing of aerial dispersal in the presence of prey. Here, we test whether artificial selection for lines that differ in timing of dispersal also results in these lines differing in prey exploitation. Six rounds of selection for early or late dispersal resulted in predator lines displaying earlier or later dispersal. Moreover, it resulted—at the population level—in predicted differences in the local predator–prey interaction time and in the cumulative numbers of dispersers in a population dynamics experiment. We pose that timing of dispersal is a heritable trait that can be selected in P. persimilis, which results in lines that show quantitative differences in local predator–prey dynamics. This opens ways to experimentally investigate the evolution of alternative prey exploitation strategies and to select for predator strains with prey exploitation strategies resulting in better biological control.     

Predator control of ecosystem nutrient dynamics

Predators are predominantly valued for their ability to control prey, as indicators of high levels of biodiversity and as tourism attractions. This view, however, is incomplete because it does not acknowledge that predators may play a significant role in the delivery of critical life‐support services such as ecosystem nutrient cycling. New research is beginning to show that predator effects on nutrient cycling are ubiquitous. These effects emerge from direct nutrient excretion, egestion or translocation within and across ecosystem boundaries after prey consumption, and from indirect effects mediated by predator interactions with prey. Depending on their behavioural ecology, predators can create heterogeneous or homogeneous nutrient distributions across natural landscapes. Because predator species are disproportionately vulnerable to elimination from ecosystems, we stand to lose much more from their disappearance than their simple charismatic attractiveness.     

Experimentally induced life-history evolution in a killifish in response to the introduction of guppies

Life-history theory predicts that increased predation on juvenile age/size-classes favors delayed maturation and decreased reproductive investment. Although this theory has received correlative support, experimental tests in nature are rare. In 1976 and 1981, guppies (Poecilia reticulata) were transplanted into localities that previously only contained a killifish, Rivulus hartii. This situation presents an opportunity to experimentally test this life-history prediction because guppies prey upon young Rivulus. We evaluated the response to selection in Rivulus by measuring phenotypic and genotypic divergence between introduction and upstream "control" localities that lack guppies. Contrary to expectations, Rivulus from the introduction sites evolved earlier maturation and increased reproductive investment within 25 years. Such evolutionary changes parallel previous investigations on natural communities of Rivulus, but do not comply with predictions of age/size-specific theory. Guppies also caused reduced densities and increased growth rates of Rivulus, which are hypothesized indirect effects of predation. Additional life-history theories show that changes in density and growth can interact with predator-induced mortality to alter the predicted trajectory of evolution. We discuss how these latter frameworks improve the fit between theory and evolution in Rivulus.