The control of Monomorium pharaonis (Hymenoptera: Formicidae) with Bacillus thuringiensis

In this study, the effect of different preparations made from Bacillus thuringiensis var. thuringiensis (strains: CCEB 555 and CCEB 058) on ants, Monomorium pharaonis, under laboratory conditions is reported. The different preparations tested consisted of (1) a liquid culture of the strain B. thuringiensis CCEB 555 (containing spores and exotoxin), (2) the supernatant of the culture broth of strain CCEB 555 (containing exotoxin), and (3) the biological preparation “Bathurin” prepared from the strain B. thuringiensis CCEB 058 (containing spores and inclusions, without exotoxin). The preparations were used either pure or in alternation with borax, i.e., 1 wk borax, 3 wk the respective preparation for several months. All preparations were found to be toxic to M. pharaonis and their effect was characterized by a slow extinction of the ant colony. Administration of “Bathurin” (1.3%) yielded a 100% mortality after 20 wk. Using a liquid culture of B. thuringiensis var. thuringiensis, 100% mortality was recorded after 21 wk, a period of time which did not differ from that obtained with the supernatant of the culture containing exotoxin. The alternation with borax was found to accelerate ant mortality by 9–10 wk after administration. In all experiments, the worker ants died first, the queen ants surviving them by 1–3 wk.In experiments employing worker ants only, a 100 and 98% mortality, respectively, occurred within 3 wk after administration of a liquid culture of B. thuringiensis and “Bathurin” supplemented with borax.     

Larvae of the green lacewing <Emphasis Type="Italic">Mallada desjardinsi</Emphasis> (Neuroptera: Chrysopidae) protect themselves against aphid-tending ants by carrying dead aphids on their backs

Larvae of the green lacewing Mallada desjardinsi Navas are known to place dead aphids on their backs. To clarify the protective role of the carried dead aphids against ants and the advantages of carrying them for lacewing larvae on ant-tended aphid colonies, we carried out some laboratory experiments. In experiments that exposed lacewing larvae to ants, approximately 40% of the larvae without dead aphids were killed by ants, whereas no larvae carrying dead aphids were killed. The presence of the dead aphids did not affect the attack frequency of the ants. When we introduced the lacewing larvae onto plants colonized by ant-tended aphids, larvae with dead aphids stayed for longer on the plants and preyed on more aphids than larvae without dead aphids. Furthermore, the lacewing larvae with dead aphids were attacked less by ants than larvae without dead aphids. It is suggested that the presence of the dead aphids provides physical protection and attenuates ant aggression toward lacewing larvae on ant-tended aphid colonies.     

Behavior of red wood ants (Hymenoptera,Formicidae) during interaction with different symbiont partners

A comparative analysis of the behavior of Formica polyctena Först during interaction with different symbionts (free-living aphids Aphis grossulariae Kalt. and hidden larvae of the sawfly Blasticotoma filiceti Klug) was carried out. Red wood ants demonstrate different levels of functional differentiation in relatively constant groups of foragers collecting honeydew. A deep “professional” specialization with clear division of a number of tasks among foragers was studied in groups of ants tending aphids. Four professional groups of foragers with different tasks were revealed: “shepherds,” “guards,” “transporters,” and “scouts” (or “coordinators”). The groups of foragers caring for sawfly larvae mainly consist of unspecialized ants. Only few ants (about 5%) remain on duty on the fern plant near B. filiceti larvae and protect the food resource from competitors, especially from other ants. In addition, the ants demonstrate simpler behavior while collecting the larval excretion, resembling that at the sugar feeders. On the whole, the behavior of red wood ants is rather flexible. The level of functional differentiation in groups of foragers collecting honeydew is determined not only by the colony size and requirements but by the nature of their interaction with trophobionts, particularly, by the possibility of direct contact.     

Density-dependent predation of the antLasius niger (Hymenoptera: Formicidae) on two attended aphidsLachnus tropicalis andMyzocallis kuricola (Homoptera: Aphididae)

The antLasius niger was observed collecting honeydew and preying on the two aphid speciesLachnus tropicalis andMyzocallis kuricola on the chestnut treesCastanea crenata. Observation determined how the antL. niger controlled their predation on the aphids in response to the density and honeydew-productivity of the aphids.Lachnus tropicalis was a better honeydew source thanM. kuricola forL. niger in terms of the amount of honeydew collected per unit time by the ants. The number of foraging workers on a tree increased with the number ofL. tropicalis on the tree, but not with the number ofM. kuricola. The density ofL. tropicalis perL. niger worker on a tree had a positive effect on the predation activity ofL. niger on both aphids, whereas the density ofM. kuricola per ant did not have any significant effect. The predation pressure by the ant which increased withL. tropicalis density, however, directed toM. kuricola rather than toL. tropicalis. These facts suggest (1) thatL. niger control their predation activities on aphids with regards to the densities of the attended aphids per worker, and (2) that the ants prey on the aphid species producing less honeydew. The effects of the ant predation on aphids and the importance of these predation effect in antaphids interactions were discussed.     

Occurrence of ant (hymenoptera, formicidae) and aphid (homoptera, aphidinea) syninclusions in Saxonian and Rovno ambers

Ant species found as syninclusions with aphids in Rovno and Saxonian ambers are listed for the first time. In a collection of 143 pieces of Rovno amber with worker ants, aphids were found in only one syninclusion with Ctenobethylus goepperti (Mayr). In a collection of Saxonian amber, 53 aphids of the genus Germaraphis were found in 152 amber pieces with ant workers; five syninclusions with aphids contained 19 specimens of C. goepperti; two syninclusions contained ten specimens of Lasius schiefferdeckeri Mayr; and three syninclusions contained one specimen each: one Camponotus mengei Mayr, one Anonichomyrma constricta (Mayr), and one Formica flori Mayr. Considering that the Rovno and Saxonian collections are nearly equal in the number of amber pieces, the fundamental difference in the number of syninclusions is striking. The number of C. goepperti in syninclusions is disproportionately high; the proportion of syninclusions with aphids and C. goepperti is 1.6 times that of all inclusions of workers of this species. They are found in 31% of examined inclusions of workers in the collection of the Saxonian amber, but in 50% of syninclusions of the entire collection. This strongly suggests that C. goepperti and Germaraphis formed an association during their lifetime.     

Effect of grazing intensity on the interaction between Shijimiaeoides divinus asonis (Matsumura, 1929) and its attendant ants

This study was carried out in the natural herbaceous grassland of Mt. Aso, which had been almost completely grazed, and which is subjected to routine grassland burning every spring (February or March) to conserve Shijimiaeoides divinus asonis (Matsumura, 1929). We clarify that ants protect the larvae of this butterfly and evaluate the effects of grazing intensity on the attendant ant population. The results obtained are summarized as follows: (i) Five species of attendant ant were identified, with the dominant species being Formica japonica (Motschulsky, 1866) and Camponotus japonicus (Mayr, 1866). (ii) The number of attendant ants decreased at night time; however, only one or two ants attended the larva until the following morning. (iii) Ten species of insect excluding ants and three species of spiders that approached the larva were recorded on the larval host‐plant. Formica japonica and C. japonicus fought off most newcomers of other insects including the natural enemies of these butterfly larvae. (iv) The number of ants in this butterfly habitat under regular grazing intensity was significantly higher than during low grazing intensity and non‐grazing periods. (v) A positive correlation was found between the number of attendant ants and the number of butterfly larvae on the host‐plant. We concluded that the interaction between this butterfly and attendant ants is one of facultative mutualism because the attendant ants protect the butterfly larva. Therefore, the numbers of this butterfly species may decrease if the number of attendant ants decreases due to the cessation of pasturage.     

Efficiency of herbivore exclusion by ants attracted to aphids on the vetch <Emphasis Type="Italic">Vicia angustifolia</Emphasis> L. (Leguminosae)

The efficiency of herbivore exclusion by ants on the vetch Vicia angustifolia L. (Leguminosae) with extrafloral nectary, mediated by ant attraction to aphids was investigated in a field census and laboratory experiments. In the field, workers of Lasius japonicus Santschi and Tetramorium tsushimae Emery frequently visited plants of the vetch parasitized by aphids of Aphis craccivora Koch, but only a few workers visited plants without aphids. An increase in the number of ants visiting a plant with increasing numbers of aphids caused a decrease in the number of larvae of the weevil, Hypera postica Gyllenhal. Therefore, the efficiency of herbivore exclusion by ants was higher on plants parasitized by Ap.craccivora aphids than that on plants unparasitized by aphids. In the laboratory experiments, L.japonicus workers frequently patrolled not only shoots with Ap.craccivora aphids but also shoots without them. However, T.tsushimae workers visited mainly shoots with Ap.craccivora aphids but less frequently on shoots without aphids. Therefore, L.japonicus workers excluded herbivores more efficiently on plants of the vetch than T.tsushimae workers. Consequently, the efficiency of herbivore exclusion by ants on the vetch can be influenced directly by differences in ant species and indirectly by the presence of aphids on plants. The present study highlights the significance of indirect interactions between ants and plants with extrafloral nectary, mediated by ant attraction to aphids for herbivore exclusion of plants.     

Myrmecophilous aphids produce cuticular hydrocarbons that resemble those of their tending ants

Aphid-tending ants protect aphids from natural enemies and collect honeydew secreted by the aphids. However, ants also often prey on the aphids they attend. Aphids, therefore, like social parasites of ants, may well have evolved chemical mimicry as an anti-predation strategy. In this study, we aimed to determine whether the aphid Stomaphis yanonis actively produces cuticular hydrocarbons (CHCs) that resemble those of the tending ant Lasius fuji. In the wild, ants put their CHCs on the aphids that they are tending, so in this study we analyzed “ant-free” aphids. Mature aphids that exuviated in the absence of ant attendance had almost all of the hydrocarbon components that the ants’ CHCs had. Moreover, hydrocarbons artificially applied to the aphids’ body surface were lost by exuviation. Taken together, these findings indicate that mature aphids actively produced ant-like CHCs, and they constitute the first documentation of a chemical resemblance between aphids and ants in a specific aphid–ant association.     

Isolation and characterization of actinobacteria ectosymbionts from Acromyrmex subterraneus brunneus (Hymenoptera, Formicidae)

The ectosymbiont actinobacterium Pseudonocardia was isolated from the integument of Acromyrmex leaf-cutter ants and seems to play a crucial role in maintaining asepsis of the nest. Currently, there has been an intensive search for Pseudonocardia associated with several attine species, but few studies have indicated that other actinobacteria may be associated with these ants as well. We therefore characterized the culturable actinobacteria community associated with the integument of the fungus-growing ant Acromyrmex subterraneus brunneus Forel, 1893 (Hymenoptera: Formicidae). Ectosymbionts were isolated using four different media and characterized by morphological and molecular (16S rDNA) methods. A total of 20 strains were isolated, of which 17 were characterized as Streptomyces spp., and one isolate each as Pseudonocardia, Kitassatospora and Propionicimonas. Unlike other Acromyrmex species, A. subterraneus brunneus is associated with a diversity of actinobacteria. Even though Pseudonocardia is present on this leaf-cutting ant’s integument, the number and diversity of Streptomyces spp. found differs from those of previous studies with other attine ants and suggest that different culturing approaches are needed to characterize the true diversity of microbes colonizing the integument of attine ants. Moreover, understanding the diversity of the culturable actinobacteria associated with A. subterraneus brunneus should increase our knowledge of the evolutionary relationship of this intricate symbiotic association.     

The eversible tentacle organs of Polyommatus caterpillars (Lepidoptera,Lycaenidae): Morphology,fine structure,sensory supply and functional aspects

In their late (3rd and 4th) larval stages, caterpillars of the myrmecophilous lycaenid (Lepidoptera) species Polyommatus coridon and Polyommatus icarus, possess on their 8th abdominal segment two eversible so called tentacle organs (TOs). Previous histological and behavioural results have proposed that the TOs may release a volatile substance that elicits “excited runs” in attendant ants. In our study we investigated for the first time the temporal in- and eversion pattern of TOs. Using nerve tracing, Micro-CT, light- and electron microscopy techniques we studied (i) the histology of the 8th abdominal segment, (ii) the fine structure of the cuticular and cellular apparatus of the TOs, (iii) the attachment sites of the retractor muscle of each TO and (iv) the fine structure of the long slender tentacle hairs which are exposed to the outside, when the TOs are everted and fold back into the TO-sac during inversion. Our data show that the tentacle hairs are typical insect mechanoreceptors, each innervated by a small bipolar sensory cell with a tubular body in the tip of the outer dendritic segment. The latter is enclosed by a cuticular sheath previously called the “internal cuticular duct” and misinterpreted in earlier studies as the space, where the tentacle hairs actively secrete fluids. However, we found no glandular structures nearby or in the wall of the TO-sac. Also we did not reveal any conspicuous signs of secretory activity in one of the enveloping cells belonging to a tentacle hair. Although highly unusual features for an insect mechanoreceptor are: (a) the hair-shaft lumen of tentacle hairs contains flocculent material as well small vesicles and (b) the thin cuticular wall of the hair-shaft and its spines possess few tiny pores. Our data do not support the assumption of previous studies that volatile substances are released via the tentacle organs during their interactions with ants which in turn are supposed to cause excited runs in ants.     

Host plant preference in the protected myrmecophilous Transylvanian Blue (<Emphasis Type="Italic">Pseudophilotes bavius hungarica</Emphasis>) butterfly (Lepidoptera: Lycaenidae) and its relationship with potential ant partners

When selecting specific host plants, caterpillars of many lycaenid butterflies, such as the protected Pseudophilotes bavius hungarica, are known to engage in various interactions with ants, which help them survive. Although P. bavius is a protected species, data about its host plant selection is very scarce, and little information is available on its myrmecophilous relationships. Our aim was to identify the host plant characteristics that determine the occurrence of the caterpillar and to clarify the specificity of its myrmecophily. We conducted a series of field surveys regarding host plant characteristics. Laboratory experiments were carried out to investigate the nature of interactions between the caterpillar and its potential ant partners. Control experiments involving non-visiting ants were also performed. On the basis of our findings, the physical characteristics of host plants do not seem to influence host plant choice, but the absence of aphids and the presence of different ant species proved important. According to the results of behavioural assays, neutral reactions to the caterpillars were recorded in the case of ant species that regularly visited the host plant (Lasius paralienus, Camponotus aethiops), in contrast to Tapinoma subboreale, which was not observed at all on the host plants and which behaved aggressively towards the larvae. Therefore, the caterpillar is expected to show a certain ant host selectivity. The study constitutes an essential contribution to our knowledge of the natural history of a protected butterfly species, which can be used as a basis for more appropriate management strategies, while also shedding light on aspects of myrmecophilous relationships in Lycaenidae in general.     

Role of xylem consumption on osmoregulation in Macrosiphum euphorbiae (Thomas)

Aphids are phloem feeders that occasionally ingest xylem sap. The duration of xylem consumption by Macrosiphum euphorbiae (Hemiptera: Aphididae) was positively correlated with the level of dehydration of alate aphids of different ages after a period of starvation, supporting the hypothesis that aphids ingest xylem sap to replenish their water balance. However, the duration of xylem sap ingestion but not phloem sap consumption varied in unstarved alate adults of different ages. Furthermore, both alate and apterous aphids ingested xylem sap at the end of their life, when aphids were not dehydrated but when fecundity started to decrease. Fecundity was negatively correlated with the proportion of time spent ingesting xylem sap, and that over the entire reproductive life of alate and apterous aphids. The lower proportion of xylem ingested by apterous than by alate aphids during the first few days of adult life may be related to a higher symbiont density in apterous morphs. As previous studies have demonstrated a relationship between sucrose assimilation, which is directly influenced by fecundity and symbiont density, and osmoregulation, we suggest that xylem consumption may play a role in the osmoregulation of haemolymph of aphids.     

Bioassay of Entomophthora against the spotted alfalfa aphid Therioaphis trifolii f. maculata

A method for the bioassay of Enthomophthora spp. against aphids is described. Twenty-four isolates comprising five species of fungi were screened for activity against Therioaphis trifolii f. maculata. The only isolates with a high level of activity were those of E. sphaerosperma obtained from aphids. The initial bioassays with E. sphaerosperma indicated that aphids, starved for 24 hr during inoculation with E. sphaerosperma primary spores, were less susceptible than those removed from the plants, for just the period of exposure to the primary spore shower. Using the latter procedure, five assays of the most pathogenic isolate gave a mean LC₅₀ of 11.3 primary spores/mm² while bioassays of four other isolates gave LC₅₀ values ranging from 15.7 to 27.3 spores/mm². The potential of E. sphaerosperma as a microbial control agent for T. trifolii f. maculata in Australia is discussed.     

Prey preference and prey switching in Anthocoris minki Dohrn (Hemiptera: Anthocoridae)

The understanding of host specificity of predacious insects is one of the most important problems in the investigation of biological control. Prey preference and the potential for prey switching in Anthocoris minki Dohrn (Hem.: Anthocoridae) was assessed in two- choice experiments under environmentally controlled conditions. Manly's preference index was used to compare A. minki predation on Agonoscena pistaciae Burckhardt and Lauterer (Hem.: Psyllidae) versus Psyllopsis repens Loginova (Hem.: Psyllidae) but showed no statistically significant evidence for preference. Anthocoris minki did prefer psyllids (Agonoscena pistaciae and Psyllopsis repens) over aphids (Forda hirsuta Mordvilko (Hem.: Pemphigidae) and Aphis craccivora Koch (Hem.: Aphididae)). The predator also consumed significantly more 2nd instar than 4th instar nymphs of P. repens. Switching behavior experiments showed that A. minki did not preferentially attack the most abundant prey when the prey combination was either biased toward A. pistaciae or toward F. hirsuta. Linear regression between preference indices and prey ratios indicated an overall preference for psyllids over aphids. The results suggest that a psyllid-specialized feeder such as A. minki is more likely to be effective biocontrol agent against psyllid species and less effective at reducing the numbers of other pest species that might co-occur with psyllids.     

The effects of the invasive Argentine ant (Linepithema humile) and the native ant Prenolepis imparis on the structure of insect herbivore communities on willow trees (Salix lasiolepis)

Abstract 1. We examined the relative effects of the invasive Argentine ant, Linepithema humile, and a common native ant, Prenolepis imparis, on the community of herbivorous insects occurring on willow trees, Salix lasiolepis in Northern California, U.S.A. 2. Using paired control and treatment branches from which we excluded ants and other non‐volant predators, we found that effects of Argentine ants on the herbivore community were generally similar to those of P. imparis. Argentine ants and P. imparis suppressed the damage by skeletonising insects by 50%, but had little effect on most other external‐feeding or internal‐feeding guilds. 3. The abundance of aphids was 100% greater in the presence of Argentine ants, but there was no effect on aphid numbers in the presence of P. imparis. Late season aphid numbers were substantially higher in the presence of Argentine ants, but not P. imparis. 4. The effects of Argentine ants on skeletonising insects and aphids combined with the overwhelming abundance of Argentine ant workers, suggests that they may have substantial, but often overlooked, effects on the herbivore communities on other plant species in or near riparian habitats in which they invade.     

Laurus novocanariensis essential oil: Seasonal variation and valorization

We investigated the qualitative and quantitative seasonal variation of the leaf and fruit oils of the Macaronesian endemism Laurus novocanariensis and their plant defensive potential. The monoterpene fraction dominated the leaf (74%) and berry essential oils (73–44%, ripe–unripe). The insect antifeedant effects of these oils were species- and season-dependent against the aphids (Myzus persicae and Rhopalosiphum padi). Overall, the biological effects of these oils correlated with the oxygenated terpene fraction. Among the pure components tested, β-caryophyllene and its oxide were strong antifeedants to Leptinotarsa decemlineata and Spodoptera littoralis. The aphids responded to β-ocimene, β-pinene, 1,8-cineole, linalool (antifeedants) and linalool oxide (attractive to M. persicae). The antifungal effects of the leaf oils on Fusarium spp. were season-dependent. β-Caryophyllene oxide proved to be a strong antifungal. L. novocanariensis oils inhibited Lactuca sativa germination and radicle elongation, the leaves being more effective. Linalool also inhibited seed germination.     

Comparative study of the structure of the reproductive system of dwarfish males of Glyphina betulae (Linnaeus, 1758) and Anoecia (Anoecia) corni (Fabricius, 1775) (Hemiptera, Aphididae)

The morphology and ultrastructure of the male reproductive system of dwarfish males of the monoecious aphid species Glyphina betulae (subfamily Thelaxinae) and the heteroecious species Anoecia (Anoecia) corni (subfamily Anoeciinae) are described. The testicular follicle of these species has the form of a single sac, the proximal parts of the vasa deferentia are slightly (G. betulae) or strongly (A. (A.) corni) expanded, the accessory glands are sack-shaped, and in G. betulae asymmetric and strongly elongated, whereas the ejaculatory duct is short.In both species only mature spermatozoa have been found within the testicular follicles, i.e. the consecutive stages of spermatogenesis have not been observed in adult males. Our studies also show that the testicular follicle, vasa deferentia, accessory glands and ejaculatory duct are histologically very simple. They are composed of more-or-less flattened epithelium of a secretory type, and thin muscle fibres. The epithelial cells are rich in rough endoplasmic reticulum, mitochondria and small vacuoles. The vasa deferentia, especially in G. betulae, are filled with an electron-dense secretion which, as was shown by histochemical staining, contains proteins and polysaccharides. We suggest that the maximum secretory activity of these epithelial cells occurs, as does spermatogenesis, during larval stages, so that the short living adult males are immediately ready for copulation as in other aphids with normal-sized males.     

Associative learning in ants: Conditioning of the maxilla-labium extension response in Camponotus aethiops

Associative learning has been studied in many vertebrates and invertebrates. In social insects, the proboscis extension response conditioning of honey bees has been widely used for several decades. However, a similar paradigm has not been developed for ants, which are advanced social insects showing different morphological castes and a plethora of life histories. Here we present a novel conditioning protocol using Camponotus aethiops. When the antennae of a harnessed ant are stimulated with sucrose solution, the ant extends its maxilla-labium to absorb the sucrose. We term this the “maxilla-labium extension response” (MaLER). MaLER could be conditioned by forward pairing an odour (conditioned stimulus) with sucrose (unconditioned stimulus) in the course of six conditioning trials (absolute conditioning). In non-rewarded tests following conditioning, ants gave significantly higher specific responses to the conditioned stimulus than to a novel odour. When trained for differential conditioning, ants discriminated between the odour forward-paired with sucrose and an odour forward-paired with quinine (a putative aversive stimulus). In both absolute and differential conditioning, memory lasted for at least 1 h. MaLER conditioning allows full control of the stimulation sequence, inter-stimulus and inter-trial intervals and satiety, which is crucial for any further study on associative learning in ants.     

Expression of the <Emphasis Type="Italic">zga</Emphasis> agglutinin gene in tobacco can enhance its anti-pest ability for peach-potato aphid (<Emphasis Type="Italic">Myzus persica</Emphasis>)

The effect of introduction of the Zephyranthes grandiflora agglutinin gene (zga) to tobacco on its anti-pest ability for peach-potato aphids was investigated. PCR analysis confirmed that the zga gene was integrated into the plant genome. The results from semi-quantitative RT-PCR and real-time PCR assays revealed that the zga gene was expressed at various levels in the transgenic plants. A bioassay with aphids indicated that transgenic plants conferred enhanced resistance to aphids. Compared with the controls, the average number of aphids fed with transgenic plants during a 20-day assay evidently decreased by 70.4% in leaf disc bioassay and 77.9% in whole plant bioassay. The average number of nymph was significantly reduced by 36.4% on zga-expressing plants in leaf disc bioassay and 35.6% in whole plant bioassay. The report indicated that the introduction of zga gene to tobacco plants is a useful method to improve its anti-pest ability for aphids.     

Larvae of the exotic predatory ladybird <Emphasis Type="Italic">Platynaspidius maculosus</Emphasis> (Coleoptera: Coccinellidae) on citrus trees: prey aphid species and behavioral interactions with aphid-attending ants in Japan

An aphidophagous ladybird, Platynaspidius maculosus (Weise) (Coleoptera: Coccinellidae), is originally distributed in China, Taiwan, and Vietnam. The ladybird has recently intruded into the southern and central parts of Japan. The present study found that the larvae of this ladybird preyed on three aphid species, Aphis spiraecola, Aphis gossypii, and Toxoptera citricidus (all Hemiptera: Aphididae), feeding on young shoots of various Citrus species in August to early October in Shizuoka Prefecture, central Japan. Laboratory rearing of the sampled larvae confirmed that the larvae completed their development (adult emergence) by consuming each of the three aphid species. The ladybird larvae were observed foraging in aphid colonies attended by one of the four ants, Lasius japonicus, Pristomyrmex punctatus, Formica japonica, and Camponotus japonicus (all Hymenoptera: Formicidae). Field observations revealed that the foraging/feeding larvae were almost completely ignored by honeydew-collecting ants even when they physically contacted each other. Thus, in Japan, the larvae of the exotic ladybird exploit colonies of the three aphid species attended by one of the four ant species on many Citrus species. On the basis of the results, I discuss the possibility of the ladybird’s reproduction on citrus trees in Japan, probable adaptations of the ladybird larvae to aphid-attending ants, and potential impacts of the ladybird on native insect enemies attacking ant-attended aphids on citrus.