Intraguild predation of the aphid pathogenic fungus Pandora neoaphidis by the invasive coccinellid Harmonia axyridis

Abstract.  1. The ladybird Harmonia axyridis is an invasive alien species in many countries and is predicted to have a negative impact on native biodiversity. However, little is known on the status of this aphidophage as an intraguild predator of natural enemies of aphids such as insect-pathogenic fungi.
2. The study assessed the predation of the aphid-specific pathogenic fungus Pandora neoaphidis by adult and larval H. axyridis collected from the U.K. (an invasive population) and Japan (a native population) relative to that of the ladybird Coccinella septempunctata (native to the U.K.) and the non-U.K. C. septempunctata subspecies brucki that were either starved or unstarved.
3. Overall, predation of uninfected aphids was greater than infected aphids and, when given a choice, a preference for aphids was shown. However, the extent of this preference was dependent on the species and origin of the coccinellid. Harmonia axyridis (U.K.) consumed the greatest quantity of fungal cadavers and showed little preference for uninfected aphids over infected aphids. In contrast, C. septempunctata rarely consumed infected aphids. Life stage had no direct effects on predation but starved coccinellids consumed more uninfected aphids than infected aphids.
4.  Harmonia axyridis (U.K.) is a stronger intraguild predator of P. neoaphidis cadavers than the native species C. septempunctata and, therefore, may have an impact on the occurrence and persistence of P. neoaphidis . The differences in intraguild predation by H. axyridis collected in the U.K. and those from Japan suggests that individuals that invaded the U.K. could have a different genetic profile to those in its native range.     

东亚小花蝽对西方花蓟马和二斑叶螨的捕食选择性

多食性天敌对猎物的捕食选择性及猎物密度对天敌捕食的干扰作用直接影响到天敌对不同猎物的控制作用.在实验室条件下研究了东亚小花蝽对西方花蓟马和/或二斑叶螨不同虫态的捕食选择性,以及两猎物中一种猎物的密度变化对小花蝽取食另一种猎物的影响.结果如下:东亚小花蝽5龄若虫和成虫对西方花蓟马2龄若虫的捕食选择性均强于对其成虫,对二斑叶螨的选择性为雌成螨>若螨>幼螨.实验中西方花蓟马2龄若虫是东亚小花蝽最喜好的虫态.二斑叶螨雌成螨密度固定为60头/19.63cm2,西方花蓟马若虫密度从10增加到60时显著地减少了东亚小花蝽对二斑叶螨的取食.反之,固定西方花蓟马同样密度,增加二斑叶螨密度却没有显著改变小花蝽对西方花蓟马的取食.此结果进一步表明,西方花蓟马是东亚小花蝽更喜好的猎物.     

The role of vision and color in the close proximity foraging behavior of four coccinellid species

The role of vision and color in close-proximity foraging behavior was investigated for four species of lady beetles: Coccinella septempunctata, Hippodamia convergens, Harmonia axyridis, and Coleomegilla maculata. The effect of light level and color cues on consumption rates varied among the four predator species. The consumption rates of these predators on the pea aphid Acyrthosiphon pisum (Harris) was measured under light and dark conditions. C. septempunctata,H. convergens, and Ha. axyridis consumed significantly more aphids in the light than in the dark, while the consumption rate of Col. maculata was not affected by light level. Foraging ability was also measured on red and green color morphs of the pea aphid on red, green, and white backgrounds. C. septempunctata consumed significantly more of the aphid morph that contrasted with the background color, and showed no difference between morphs on the white background. H. axyridis consumed significantly more red morph aphids regardless of background. The remaining two species showed no difference in consumption rates on the two color morphs. The variation in the use of visual cues demonstrates how different species of predators can exhibit different foraging behaviors when searching for the same prey. Received: 4 August 1997 / Accepted: 3 February 1998     

Predation preferences and discrimination between con- and heterospecific prey by the phytoseiid mites Phytoseiulus persimilis and Neoseiulus californicus

Discrimination between and predation preference for con- or heterospecific larvae was examined for adult females of P. persimilis and N. californicus in plexiglass cages with and without their primary prey T. urticae. Rates of intra- and interspecific predation on larvae were measured for females held on leaves and provided with excess amounts of spider mites. Females of the generalist N. californicus distinguished con- and heterospecific larvae and preferred to prey upon the latter. Females of the specialist P. persimilis appeared to lack discrimination ability and fed equally on con- and heterospecifics. When spider mites and phytoseiids were offered simultaneously, all P. persimilis females chose to first attack T. urticae, whereas N. californicus females attacked both tetranychids and heterospecific phytoseiids. Females of both predators preyed upon phytoseiid larvae when held on leaves with surplus T. urticae: while P. persimilis fed on both con- and heterospecifics, N. californicus attacked larvae of P. persimilis but avoided cannibalizing larvae. The different behaviors of P. persimilis and N. californicus are discussed with regard to different predation types (generalists vs. specialists) and the possible consequences of mixed release for biological control of spider mites in greenhouses.     

Euseiusfinlandicus (Acari: Phytoseiidae) as a potential biocontrol agent against Tetranychus urticae (Acari: Tetranychidae): life history and feeding habits on three different types of food

Life history and reproductive parameters of the generalist predatory mite Euseius (Amblyseius) finlandicus (Oudemans) were studied in the laboratory at 25 +/- 1 degrees C, with a 16L:8D photoperiod and 60 +/- 15% RH, to investigate its response to different food sources: an eriophyid mite Aceria sp., tulip pollen Tulipa gesnerana L., and two-spotted spider mite Tetranychus urticae Koch. Total developmental time of the immature stages was the shortest on eriophyid mites, followed by pollen, and then spider mites. Fecundity was highest on pollen (43.69 eggs; 1.63 eggs/female/day), then eriophyid mites (39.73 eggs; 1.37 eggs/female/day) and lowest on spider mites (18.16 eggs; 0.80 eggs/female/day). Intrinsic rate of increase (Rm), net reproductive rate (Ro) and finite rate of increase (lambda) followed the same pattern [pollen (0.168, 27.96 and 1.183, respectively), eriophyid mites (0.153, 20.81 and 1.167), spider mites (0.110, 9.44 and 1.119)]. Mean generation time (days) was the shortest on pollen (19.90), followed by eriophyid mites (20.02), and then spider mites (20.59). Average spider mite larvae consumed by E. finlandicus during immature stages were 9.18 for males and 11.85 for females. Adult E. finlandicus females consumed an average of 166.38 spider mite protonymphs during adult stage compared to an average of 66.55 by males. The number of prey protonymphs consumed per day by females was highest in the oviposition period, lower in the pre-oviposition period and the lowest in the post-oviposition period. The eriophyid mite as a prey recorded the shortest developmental time, while pollen as food recorded the highest oviposition rate in E. finlandicus. The potential of this predator as a biocontrol agent against T. urticae is discussed.     

Prey preference of Orius sauteri between Western Flower Thrips and spider mites

Prey preference of polyphagous predators plays an important role in the suppression of various species of pest insects. In this study, the prey preference of Orius sauteri (Poppius) (Heteroptera: Anthocoridae) between deutonymphs of Tetranychus urticae Koch (Acari: Tetranychidae) and second instars of Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), and the influence of spider mite density on the preference were examined in the laboratory in three-dimensional set-ups at 25 ± 1 °C and 70 ± 5% r.h. The results showed that the predatory bug in the presence of equal densities of the two prey species had a clear preference for thrips (preference index β: 0.86 ± 0.02), consuming 7.2 thrips larvae and 1.5 spider mite deutonymphs during the experimental periods of 6 h. The number of thrips consumed by O. sauteri did not change when the density ratio of thrips to spider mites was decreased from 1:1 to 1:5. Predation on spider mites increased when their ratio to thrips increased, but the disproportionate predation (i.e., the functional response) of O. sauteri towards spider mites resulted in a linear increase in the preference for F. occidentalis . The possible implications of these findings for the suppression of spider mites by O. sauteri in relation to the recent invasion of F. occidentalis into China are discussed.     

Effects of nutrients and predators on an old-field food chain: interactions of top-down and bottom-up processes

In theory, selection favours predators that select prey in order to maximise reproductive success. We studied the association between preference and performance of the generalist predator Orius laevigatus with respect to two prey species: spider mites ( Tetranychus urticae ) and western flower thrips ( Frankliniella occidentalis ). Under ample prey supply, the predators had higher maximum reproductive success (measured as intrinsic population growth rate r ) on thrips than on spider mites; hence thrips represent a higher prey quality to the bugs. This was at odds with the observed preference of the predatory bug for plants (patches) with high densities of spider mites to plants with moderate densities of thrips in release-recapture experiments. Thus, prey quality does not suffice to explain the preference of predators for plants with prey. The quality of a patch as an oviposition site (i.e. the number of eggs produced on a patch per bug per day) also did not match preference patterns. Hence, patch preference was not correlated to prey quality or oviposition rate on prey patches. However, patch productivity, i.e. the total number of offspring surviving until adulthood that can be produced by one female on a patch, was correlated with preference. This was further tested by offering the predators a choice between plants with high densities of spider mites and plants with high densities of thrips in an independent set of release-recapture experiments. These two types of prey patches were found equivalent in terms of patch productivity. Indeed, the predators showed no preference for either of the two types of patches, which is in agreement with our predictions. This suggests that the predatory bugs select patches based on patch productivity rather than on prey quality or oviposition rate on a patch.     

Effect of cannibalism and predation on the larval performance of two ladybird beetles

The relative impact of cannibalism and predation on the development and survival of fourth instar larvae was assessed in two species of aphidophagous ladybird beetles, Coccinella septempunctata and Harmonia axyridis. The effect of eating aphids, conspecific larvae or heterospecific larvae on larval performance differed in the two species: aphids were the best food for C. septempunctata and survival of C. septempunctata larvae was significantly lower when offered heterospecific larvae rather than conspecific larvae or aphids as food, indicating that H. axyridis larvae were not suitable food for C. septempunctata. However, as the different foods did not affect the larval performance of H. axyridis, this species appears to be more polyphagous. Both intraguild predation by the aggressive larvae of H. axyridis and the polyphagous food habit of this species may account for its dominance in ladybird assemblages and its displacing other ladybird beetles in several places in the world.     

Altered host plant preference of Tetranychus urticae and prey preference of its predator Phytoseiulus persimilis (Acari: Tetranychidae, Phytoseiidae) on transgenic Cry3Bb-eggplants

The behavior of the two-spotted spider mite, Tetranychus urticae Koch and the predatory mite Phytoseiulus persimilis A.-H. was investigated in laboratory experiments with transgenic Bt-eggplants, Solanum melongena L., producing the Cry3Bb toxin and corresponding isogenic, non-transformed eggplants. In bitrophic experiments, dual-choice disc tests were conducted to reveal the effects of transgenic eggplants on host plant preference of T. urticae. Adult spider mite females were individually placed on leaf discs (2 cm diameter) and were observed during five days. Females occurred significantly more frequently on transgenic halves on which also significantly more T. urticae eggs were found. The effects of a Cry3Bb-eggplant fed prey on the feeding preference of P. persimilis were investigated in tritrophic experiments. Sixteen spider mite females, eight of which had been taken from transgenic and eight from isogenic eggplants, were offered to well-fed females of P. persimilis and numbers of respective spider mites consumed were registered 12 h later when the predators were offered new spider mites again. This procedure was repeated six times. The results revealed that predatory mites consumed significantly less Bt-fed spider mites than prey that had been raised on control eggplants. These results indicate that eggplants expressing the Cry3Bb toxin for resistance against the Colorado potato beetle are more preferred by spider mites but are less preferred by their predator P. persimilis. Possible consequences of these findings for biological control of spider mites on eggplants are discussed.     

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.     

Prey preference,intraguild predation and population dynamics of an arthropod food web on plants

The theory of intraguild predation (IGP) largely studies effects on equilibrium densities of predators and prey, while experiments mostly concern transient dynamics. We studied the effects of an intraguild (IG) predator, the bug Orius laevigatus, on the population dynamics of IG-prey, the predatory mite Phytoseiulus persimilis, and a shared prey, the phytophagous two-spotted spider mite Tetranychus urticae, as well as on the performance of cucumber plants in a greenhouse. The interaction of the predatory mite and the spider mite is highly unstable, and ends either by herbivores overexploiting the plant or predators exterminating the herbivores. We studied the effect of IGP on the transient dynamics of this system, and compared the dynamics with that predicted by a simple population-dynamical model with IGP added. Behavioural studies showed that the predatory bug and the predatory mite were both attracted to plants infested by spider mites and that the two predators did not avoid plants occupied by the other predator. Observations on foraging behaviour of the predatory bug showed that it attacks and kills large numbers of predatory mites and spider mites. The model predicts strong effects of predation and prey preference by the predatory bugs on the dynamics of predatory mites and spider mites. However, experiments in which the predatory bug was added to populations of predatory mites and spider mites had little or no effect on numbers of both mite species, and cucumber plant and fruit weight.     

Feeding and egg distribution studies of Heringia calcarata (Diptera: Syrphidae), a specialized predator of woolly apple aphid (Homoptera: Eriosomatidae) in Virginia apple orchards

Predation by the aphidophagous syrphid fly Heringia calcarata (Loew) on woolly apple aphid, Eriosoma lanigerum (Hausmann), was studied in the laboratory and in Virginia apple orchards. Feeding studies compared the prey suitability of three temporally sympatric aphid pests of apple: spirea aphid, Aphis spiraecola Patch; rosy apple aphid, Dysaphis plantaginea (Passerini); and woolly apple aphid. Significantly more H. calcarata larvae survived and completed development on a pure diet of woolly apple aphid than on rosy apple aphid, and none survived on spirea aphid. Final larval weights were significantly greater, and the larval developmental period was significantly shorter on woolly apple aphid than on rosy apple aphid, but neither the duration of pupal development nor adult weight differed between diets. H. calcarata larvae consumed an average of 105 woolly apple aphids during their development. Na?ve, neonate larvae given access to all possible pair combinations of woolly apple aphid, rosy apple aphid, and spirea aphid consumed significantly more woolly apple aphids in all pairings that included woolly apple aphid. When given a choice of rosy apple aphid and spirea aphid, significantly more rosy apple aphids were consumed. Weekly counts of syrphid eggs found in woolly apple aphid, rosy apple aphid, and spirea aphid colonies collected from apple trees showed that two generalist hover fly predators, Eupeodes americanus (Wiedemann) and Syrphus rectus Osten Sacken, were present in colonies of all three aphid species and that E. americanus was the most abundant syrphid predator in A. spiraecola and D. plantaginea colonies. H. calcarata eggs were found only in woolly apple aphid colonies and were more abundant there than E. americanus and S. rectus. These data suggest that H. calcarata is a specialized predator of woolly apple aphid in the apple ecosystem in Virginia.     

The effect of powdery mildew on the number of prey consumed by Typhlodromus pyri (Acari: Phytoseiidae)

Abstract:  Prey consumption by Typhlodromus pyri Scheuten was studied in the presence and absence of apple powdery mildew, Podosphaera leucotricha (Ell. and Everh.) under constant laboratory conditions. Eggs of Tetranychus urticae Koch were offered to predatory mites as a prey. Seven densities ranging from five to 100 T. urticae eggs per arena were used. Mildew conidia (approximately 0.5 mg) were added to half of the arenas by brushing them from infected apple leaves. A single adult female of T. pyri was introduced onto each arena and number of prey eggs consumed was counted 12 h later when the predator was offered new T. urticae eggs again and the procedure was repeated once. Data showed that predators consumed in both experimental periods nearly all prey in experiments with densities up to 40 eggs per arena and no mildew. However, the number of eggs consumed decreased more than twofold when mildew conidia were supplied, even at high prey densities. Differences in predation rate between treatments with and without mildew were highly significant. The results thus indicate that availability of mildew as an alternative food can reduce prey suppression by T. pyri . Possible implications of these findings in biological control of spider mites by generalist predatory mites are discussed.     

Intra- and interspecific predation by adult female Metaseiulus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae) when provisioned with varying densities and ratios of Tetranychus urticae (Acari: Tetranychidae) and phytoseiid larvae

The phytoseiid mites Metaseiulus occidentalis (Nesbitt) and Typhlodromus pyri Schueten are used together and alone as biological control agents against tetranychid pest mites of apple. Their effectiveness as control agents may be impacted by intraguild predation. The effects of prey species and prey density on the rates of inter- and intraspecific predation and oviposition by these two predators were investigated through a series of experiments. Adult female predators were given prey as mixed populations of phytoseiid larvae and larvae of a more preferred species, the spider mite, Tetranychus urticae Koch, at different densities and ratios. Typhlodromus pyri, more of a generalist predator, showed higher rates of predation and cannibalism on phytoseiid immatures at most prey densities and ratios. Manly preference indices indicated that T. pyri switched to feed on phytoseiid larvae at higher prey levels and ratios of T. urticae than M. occidentalis. This greater ability to use phytoseiid larvae as prey may help stabilize T. pyri populations when more preferred prey is unavailable. This may, in part, explain the observed persistence of T. pyri populations when M. occidentalis populations were decreasing in orchard test plots.     

Prey preferences of six species of Carabidae (Coleoptera) and one Lycosidae (Araneae) commonly found in UK arable crop fields

Abstract: Laboratory experiments were used to determine the feeding preferences of six carabid beetles and one lycosid spider on aphid and collembolan prey. The first investigation used only five species of carabid Pterostichus melanarius Illiger, Pterostichus madidus F., Harpalus rufipes DeGeer, Nebria brevicollis F. and Carabus violaceus L., which had been caught most commonly in pitfall traps in the headland region of arable crop fields. When offered Brevicoryne brassicae L., Sitobion avenae F., Metopolophium dirhodum Walker and Rhopalosiphum padi L. as prey items, the species consistently consumed in high numbers was M. dirhodum . In subsequent experiments four carabids Pterostichus cupreus L., P. melanarius, P. madidus. H. rufipes and a lycosid spider Trochosa ruricola DeGeer whose distribution was shown by pitfall trapping to extend throughout the arable crop, were the chosen predators. These predators were offered a choice between M. dirhodum and entomobryid collembolans (a recognised alternative prey item) to gauge their preference between the two prey types. Both male and female P. cupreus and P. melanarius showed a significant preference for the aphid prey, while there was no significant preference displayed by the other species. The effect of temperature on the voracity of these five predators fed on M. dirhodum was investigated. There were significant differences in the number of aphids consumed by the species and sexes at the different temperatures. Regression analysis on the mean numbers of aphids eaten by each sex of the five predators, showed that in the majority of cases there was a significant increase in predation with increasing temperature. In considering the dietary preferences illustrated by these experiments, it appeared that P. cupreus and P. melanarius offered the greatest potential in controlling aphids on arable crops.     

Interactions between introduced and native predatory ladybirds (Coleoptera, Coccinellidae): factors influencing the success of species introductions

Abstract.  1. Why some exotic species invade successfully while others do not is poorly understood. This study focuses on the inability of the exotic predatory ladybird Adalia bipunctata (L.) (Coleoptera, Coccinellidae) to establish and spread in Japan. In particular, this study considers the interactions between A. bipunctata and three native Japanese ladybird species, Harmonia axyridis Pallas, Coccinella septempunctata Linn., and Propylea japonica Mulsant.
2. The number of aphids occurring on the host plants declined dramatically over the course of larval development in the presence of H. axyridis or C. septempunctata . In contrast, many aphids remained at the end of the experiment when only A. bipunctata or P. japonica was present.
3. The survivorship of A. bipunctata from second instar to adult was significantly reduced in the presence of H. axyridis , but not in the presence of C. septempunctata or P. japonica . Most deaths were the result of intra-guild predation when H. axyridis or C. septempunctata larvae were present. Larvae of H. axyridis and C. septempunctata suffered very high mortality in this experiment, but survived better when co-occurring with A. bipunctata than when occurring only with conspecifics. In contrast, P. japonica survived better when occurring with conspecifics than with A. bipunctata .
4. The finding that larvae of the exotic species A. bipunctata died at high rates from intra-guild predation by the native Japanese ladybirds H. axyridis and C. septempunctata may bear on the question of why A. bipunctata has not established more successfully in Japan whereas H. axyridis and C. septempunctata have been very successful in establishing themselves in North America.     

Predation and oviposition by predatory Stethorus japonicus, Oligota kashmirica benefica, and Scolothrips takahashii in egg patches of various spider mite species

Predation and oviposition by three predacious insects [ Scolothrips takahashii Priesner (Thysanoptera: Thripidae), Stethorus japonicus H. Kamiya (Coleoptera: Coccinellidae), and Oligota kashmirica benefica Naomi (Coleoptera: Staphylinidae)] were examined using egg patches of three spider mite species [ Amphitetranychus viennensis (Zacher), Tetranychus urticae Koch, and Panonychus mori Yokoyama (all Acari: Tetranychidae)] that are pest species on Japanese pear [ Pyrus serotina Rehder (Rosaceae)]. Scolothrips takahashii females consumed more prey and laid more eggs in A. viennensis and T. urticae egg patches than in P. mori egg patches. Females also left P. mori egg patches sooner than they left other egg patches even at high prey density. Predation and oviposition of S. takahashii in a prey patch were greatly enhanced by the complicated webs produced by A. viennensis . Stethorus japonicus females consumed significantly more prey and laid more eggs in A. viennensis egg patches than in P. mori egg patches, with intermediate values in T. urticae egg patches, reflecting the quality of each spider mite species as prey. Oligota kashmirica benefica females consumed more prey and laid more eggs in A. viennensis and T. urticae egg patches than in P. mori egg patches, particularly at high prey density. However, predation and oviposition by O. kashmirica benefica increased greatly with increasing prey density, even in P. mori egg patches, indicating that prey density was the most important factor in predation and oviposition by this species. These results are discussed in relation to the potential effect of each predator on the suppression of different spider mite species on Japanese pear.     

A comparative study on effects of normal versus elevated temperatures during preimaginal and young adult period on body weight and fat body content of mature Coccinella septempunctata and Harmonia axyridis (Coleoptera: Coccinellidae)

Two climate chamber experiments were performed to simulate the effects of global warming on life table parameters of coccinellids. We investigated the effects of two daily temperature profiles during preimaginal development (larval and pupal) and the young adult period (first 10 d) on body weight and fat body content of adult Coccinella septempunctata L. and Harmonia axyridis (Pallas) fed English grain aphids [Sitobion avenae (F.)] ad libitum: 1) normal, i.e., current daily temperatures in central Europe (T0: mean, 17.8°C; maximum, 21.8°C; minimum, 13.4°C) and 2) increased by 3K (T3: mean, 20.8°C; maximum, 25.5°C; minimum, 15.7°C). The first experiment was performed at the same temperatures (T0 or T3) during both periods to establish the responses of the two species to temperature. The second was conducted to identify the period (preimaginal or adult) in which the responses occurred and to confirm the results of the first experiment. Compared with normal temperatures (T0), elevated temperatures (T3) resulted in significant decreases in preimaginal development time and increases in aphid consumption rates in both species. C. septempunctata (10-d-old adults) had the highest weights when reared at T3, H. axyridis at T0. C. septempunctata was significantly heavier than H. axyridis in most cases, particularly in females. The body fat content of C. septempunctata was higher than that of H. axyridis at T0 and T3 temperatures. At T3 temperatures, fat accumulation in C. septempunctata increased, whereas that in H. axyridis remained relatively low. Body weight and fat body content of 10-d-old adults of both species seemed to be determined by temperature conditions during preimaginal development.     

Spider Mites Avoid Plants with Predators

While searching for food, prey can use cues associated with their predators to select patches with a reduced predation risk. In many cases, odours indicate the presence of both food and predators. Spider mites are known to use odours to locate food and mates, but also to avoid interspecific competitors. We studied the response of the two-spotted spider mite, Tetranychus urticae, to cues associated with the presence of their predators, the phytoseiid Phytoseiulus persimilis. We found that the spider mites strongly avoid plants defended by this predator, but do not avoid plants with another predatory mite, Neoseiulus californicus. Since P. persimilis is commonly used in the greenhouse where our strain of T. urticae was collected and strains of this pest are known to adapt to greenhouse environments, we hypothesize that there has been selection on the pest to recognize its enemy. We further hypothesize that there has been no selection to recognize N. californicus, as it has not been used against two-spotted spider mites in the greenhouse where our spider mites were collected. We discuss the implications of avoidance of predation by spider mites and non-lethal effects of predators for biological control of this pest in greenhouses.     

Relationships Between Attack and Escape Rates, Cannibalism, and Intraguild Predation in Larvae of Two Predatory Ladybirds

We compared incidences of cannibalism and intraguild (IG) predation (IGP) and quantified attack and escape rates—mechanisms which possibly account for the difference in incidences of these interactions—in laboratory experiments with Harmonia axyridis and Coccinella septempunctata. There was a tendency for H. axyridis to act as an IG predator and C. septempunctata as an IG prey. Cannibalism was also often observed in both species. The incidences of both IGP and cannibalism were different between the species. The average attack rates of C. septempunctata were less than 20%, but those of the more aggressive H. axyridis exceeded 50%. Larvae of both species attacked conspecifics and heterospecifics H. axyridis larvae successfully escaped when attacked by both conspecifics and by heterospecifics, while larvae of C. septempunctata escaped from attacks of conspecifics but not from those of heterospecifics. Thus the aggressive behavior of H. axyridis, in particular, of the third and fourth instars, negatively affects the larval survival of C. septempunctata. It may contribute to the dominance of H. axyridis in ladybird assemblages and its displacement of other ladybird species in several places in the world.