Gallerucida bifasciata (Coleoptera: Chrysomelidae), a potential biological control agent for Japanese knotweed (Fallopia japonica)

Host specificity of foreign natural enemies are becoming more and more critical in classical biological control programs, as concerns about potential risk from introduced biocontrol agents have been increasing recently. Understanding the insect's fundamental and ecological host ranges is the first step in determining the potential for introduction of an insect to control invasive plants. Japanese knotweed, Fallopia japonica (Houttuyn) Ronse Decraene (Polygonaceae) is an invasive weed in the United States and Europe. A leaf beetle, Gallerucida bifasciata (Coleoptera: Chrysomelidae) is an important natural enemy attacking this plant in Asia. However, its host range records were ambiguous. This study examined the beetle's host specificity through a set of choice and no-choice tests in the laboratory and field in its native China. Gallerucida bifasciata larvae were able to complete development on seven of 87 plant species in larval development tests, while adults fed and oviposited on 10 plants in no-choice tests. Multiple choice tests showed adults strongly preferred Fallopia japonica, Persicaria perfoliata (L.) H. Gross and Polygonum multiflorum Thunb over all other plants. Open field tests and field surveys further revealed that these three species were in its ecological host range. The results of this study suggest that G. bifasciata is a potential promising agent for control of Japanese knotweed in the United States and Europe, although additional host specificity tests and risk assessment should be completed.     

Extreme differences in population structure and genetic diversity for three invasive congeners: knotweeds in western North America

Japanese, giant, and the hybrid Bohemian knotweeds (Fallopia japonica, F. sachalinensis and F. × bohemica) have invaded the western USA and Canada, as well as other regions of the world. The distribution of these taxa in western North America, and their mode of invasion, is relatively unresolved. Using amplified fragment length polymorphisms of 858 plants from 131 populations from British Columbia to California to South Dakota, we determined that Bohemian knotweed was the most common taxon (71 % of all plants). This result is in contrast to earlier reports of F. × bohemica being uncommon or non-existent in the USA, and also differs from the European invasion where it is rarer. Japanese knotweed was monotypic, while giant knotweed and Bohemian knotweed were genetically diverse. Our genetic data suggest that Japanese knotweed in western North America spreads exclusively by vegetative reproduction. Giant knotweed populations were mostly monotypic, with most containing distinct genotypes, suggesting local spread by vegetative propagules, whereas Bohemian knotweed spreads by both seed and vegetative propagules, over both long and short distances. The high relative abundance and genetic diversity of Bohemian knotweed make it a priority for control in North America.     

The Japanese knotweed invasion viewed as a vast unintentional hybridisation experiment

Chromosome counts of plants grown from open-pollinated seed from Japanese knotweed around the world have revealed the presence of extensive hybridisation with both native and other introduced taxa. These hybrids fit into three categories: inter- and intraspecific hybrids involving the taxa of Fallopia section Reynoutria (giant knotweeds), hybrids between Japanese knotweed and F. baldschuanica (Regel) Holub and hybrids between Japanese knotweed and the Australasian endemics of the genus Muehlenbeckia. In this minireview, the viability of the different classes of hybrid and the potential threats they pose are discussed in the context of recent examples of allopolyploid speciation, which generally involve hybridisation between a native and an alien species. Such wide hybridisations also challenge accepted taxonomic classifications. Japanese knotweed s.l. provides a fascinating example of the interplay between ploidy level, hybridisation and alien plant invasion. The octoploid (2n=88) Fallopia japonica var. japonica (Houtt.) Ronse Decraene is a single female clone throughout much of its adventive range, and provides an ideal system for investigating the potential for wide hybridisation.     

Green Frogs Show Reduced Foraging Success in Habitats Invaded by Japanese knotweed

Habitat loss is causing amphibian population declines worldwide, so there is increased attention to forces that degrade remaining habitats. Terrestrial habitats surrounding wetlands are critical foraging areas for temperate anurans. We investigated plant community changes in two old fields invaded by Japanese knotweed (Fallopia japonica) and the foraging success of Green frogs (Rana clamitans) in invaded and non-invaded portions of those fields. Within each field, vegetation data were recorded in quadrats located along two transects bisecting the invasion fronts. We placed frogs in ‘foraging buckets’ along transects and measured their change in mass over a 38 h period. There were significant changes in vegetation structure and composition associated with Japanese knotweed invasion. Diverse assemblages of native plants that covered non-invaded plots were absent from areas invaded by Japanese knotweed. There was also a significant change in vegetation architecture between invaded and non-invaded habitats. Change in frog mass declined significantly along transects, with most frogs in non-invaded plots gaining mass and no frogs in invaded plots gaining mass. Most frogs from non-invaded plots but only two from invaded plots defecated shortly after removal from foraging buckets (verification of recent feeding). We hypothesize that Japanese knotweed invasions degrade terrestrial habitat quality for frogs by indirectly reducing arthropod abundance. Nonnative plant invasions may be another factor contributing to amphibian population declines.     

Winter hosts of Aphalara itadori (Hemiptera: Psyllidae), a classical biological control agent of Fallopia japonica (Polygonaceae), in the UK

This study suggests that lack of suitable overwintering sites will not be a limiting factor to the establishment of Aphalara itadori, a classical biological control agent of Fallopia japonica, since it was able to overwinter on a common conifer species as well as non-conifers hosts.     

Candidates for biocontrol of Macfadyena unguis-cati in South Africa: biology, host ranges and potential impact of Carvalhotingis visenda and Carvalhotingis hollandi under quarantine conditions

Macfadyena unguis-cati (L.) Gentry (Bignoniaceae) was introduced as an ornamental in South Africa, but is fast becoming an important invasive plant in many areas. It is difficult to control the plant chemically and mechanically. The first biocontrol agent, the chrysomelid Charidotis auroguttata (Boheman), has been released. It established at some release sites, but numbers have so far remained low. Additional biocontrol agents were sought to augment C. auroguttata. The potential host ranges of two foliage feeding lace bugs, Carvalhotingis visenda (Drake and Hambleton) and C. hollandi (Drake) (Hemiptera: Tingidae) were evaluated on the basis of nymphal no-choice and adult multi-choice tests involving 23 plant species in 11 families. In no-choice tests, nymphs of both species were able to survive and complete development on M. unguis-cati only, and adults of both species only fed and oviposited on M. unguis-cati during the adult multi-choice tests. Host specificity tests thus confirm that the tingids are highly host specific biocontrol agents, and will not pose risk to any non-target plants in South Africa. A study to determine the potential impact of C. hollandi nymphal feeding on M. unguis-cati showed a significant decrease in the chlorophyll contents of leaves when compared to those of control plants. These studies indicate that, once released, the two lace bug species could contribute significantly to the biological control of M. unguis-cati in South Africa.     

Post-establishment assessment of host plant specificity of Arytainilla spartiophila (Hemiptera: Psyllidae), an adventive biological control agent of Scotch broom,Cytisus scoparius

Scotch broom, Cytisus scoparius (Fabaceae), is a shrub native to Europe that is invasive in the USA, New Zealand and Australia. The psyllid Arytainilla spartiophila has been purposely introduced to Australia and New Zealand as a biological control agent of C. scoparius, but is an accidental introduction to California. Lupines (Lupinus spp.) are the closest native taxon to Cytisus in North America, and are therefore considered to be at the highest risk for non-target damage. However, because no lupines are native to Australia or New Zealand, only one imported forage species was evaluated during prior host specificity testing. We conducted a laboratory nymphal transfer experiment, a field choice experiment and a field survey to assess risk to three lupine species (Lupinus albifrons, Lupinus bicolor and Lupinus formosus). In the laboratory, 20% of third-instar nymphs were able to develop to adulthood on L. formosus but not on the other lupine species, while 40% completed development on C. scoparius. In the field experiment, potted lupine and C. scoparius plants were placed beside large infested C. scoparius plants; oviposition occurred on all the potted C. scoparius plants, but on none of the lupines. In the field survey, no A. spartiophila eggs or nymphs were found on naturally occurring lupines growing adjacent to infested C. scoparius. The results indicate that A. spartiophila is not likely to damage or reproduce on lupines in the field. This study provides an example of how field studies can help clarify the host specificity of biological control agents.     

Development and survival ofOrius albidipennis andO. levigatus (Het.: Anthocoridae) on various diets

The effect of various diets on nymphal development and survival of two predaceous anthocorid bugs,Orius albidipennis (Reuter) andO. laevigatus (Fieber) was investigated in the laboratory. Five different diets were compared: eggs ofEphestia kuehniella Zeller; eggs ofE. kuehniella plus mixed flower pollen; only mixed flower pollen; pollen from sweet pepper (Capsicum annuum L. cv. Mazurka) flowers; and sweet pepper plants. A high percentage of predators successfully completed nymphal development on those diets containing lepidopterous eggs. When fed on sweet pepper pollen, the survival percentages were 65% forO. laevigatus and 38% forO. albidipennis. No nymphs of either species completed the nymphal stage on mixed flower pollen or on sweet pepper plants. Development was significantly faster on diets containing eggs ofE. kuehniella. Results are discussed in relation to the capability of the bugs to survive periods of prey scarcity and to the optimization of release strategies for these predators in the greenhouse.     

Reciprocally beneficial interactions between introduced plants and ants are induced by the presence of a third introduced species

Interspecific interactions play an important role in the success of introduced species. For example, the ‘enemy release’ hypothesis posits that introduced species become invasive because they escape top–down regulation by natural enemies while the ‘invasional meltdown’ hypothesis posits that invasions may be facilitated by synergistic interactions between introduced species. Here, we explore how facilitation and enemy release interact to moderate the potential effect of a large category of positive interactions – protection mutualisms. We use the interactions between an introduced plant (Japanese knotweed Fallopia japonica), an introduced herbivore (Japanese beetle Popillia japonica), an introduced ant (European red ant Myrmica rubra), and native ants and herbivores in riparian zones of the northeastern United States as a model system. Japanese knotweed produces sugary extrafloral nectar that is attractive to ants, and we show that both sugar reward production and ant attendance increase when plants experience a level of leaf damage that is typical in the plants’ native range. Using manipulative experiments at six sites, we demonstrate low levels of ant patrolling, little effect of ants on herbivory rates, and low herbivore pressure during midsummer. Herbivory rates and the capacity of ants to protect plants (as evidenced by effects of ant exclusion) increased significantly when plants were exposed to introduced Japanese beetles that attack plants in the late summer. Beetles were also associated with greater on‐plant foraging by ants, and among‐plant differences in ant‐foraging were correlated with the magnitude of damage inflicted on plants by the beetles. Last, we found that sites occupied by introduced M. rubra ants almost invariably included Japanese knotweed. Thus, underlying variation in the spatiotemporal distribution of the introduced herbivore influences the provision of benefits to the introduced plant and to the introduced ant. More specifically, the presence of the introduced herbivore converts an otherwise weak interaction between two introduced species into a reciprocally beneficial mutualism. Because the prospects for facilitation are linked to the prospects for enemy release in protection mutualisms, species introductions can have complex effects on existing species interactions, between both native and introduced species.     

Prey infected with nucleopolyhedrovirus is preferred by the predatory stink bug Eocanthecona furcellata without altering its field fitness

Preference tests using the hemipteran predator Eocanthecona furcellata Wolff showed that nymphs and adults showed a constant preference towards virus-infected Spodoptera litura larvae. Adults feeding on healthy larvae throughout their life handled and consumed their prey quickly as compared to those feeding on infected prey, the differences being significant. Developmental time, survival rate, pre-mating period, sex-ratio and incubation period of E. furcellata that were reared on experimental plants infested with healthy larvae of S. litura as lifetime prey for all stages (control), healthy larvae during nymphal stage and infected larvae for adult stage, infected larvae for nymphal and healthy larvae for adult stage and infected larvae as lifetime meal for all stages of E. furcellata did not vary significantly. However, a significant reduction in body weight, fecundity, longevity and percent egg hatchability was observed when E. furcellata were fed on infected larvae as lifetime meal. The results revealed that the virus-infected larvae, relative to healthy larvae, had significant effect on prey preference of predators, but feeding on virus-infected prey had no adverse effect on its field fitness except on lifetime meal, which may not hold true under field conditions.     

Short-distance dispersal behavior and establishment of the parasitoid Gonatocerus ashmeadi (Hymenoptera: Mymaridae) in Tahiti: Implications for its use as a biological control agent against Homalodisca vitripennis (Hemiptera: Cicadellidae)

The egg parasitoid Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae), was introduced into French Polynesia as a biological control agent to control the invasive plant feeding pest Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae). The short-distance dispersal of G. ashmeadi was monitored as part of the biological control program. G. ashmeadi showed exponential dispersal capacity with 47 m/day being a minimum estimate of its natural rate of spread at high host densities (>150 nymphs per minute of sweep net sampling) in urbanized areas at sea level, which were characterized by a high diversity of exotic ornamental plants. This rate of spread contrasted starkly with almost nonexistent establishment and dispersal where host densities were very low (<2 nymphs per minute of sweep net sampling) at high elevation (800 m) with relatively undisturbed native vegetation. Survey results across different altitudes revealed an effect of vegetative diversity and host density on the measurable mobility and establishment of G. ashmeadi. In contrast, no significant influence of wind direction was found on G. ashmeadi dispersal rate or direction. Survey results for G. ashmeadi from French Polynesia suggest that the best release establishment strategies for classical biological control of H. vitripennis are: (1) many small releases where host density is high, or (2) larger and fewer releases where host densities are low.     

Descriptions of two new and one little known species of the genus Cacopsylla (Hemiptera: Psyllidae) on Sorbus japonica (Rosaceae) and an observation of their biology

Three species of the genus Cacopsylla were collected from Sorbus japonica in Japan. Two of them, Cacopsylla elegans and Cacopsylla sorbicoccinea, are described as new species based on adults and fifth (final) nymphal instars. The third species, Cacopsylla midoriae, is newly transferred from the genus Psylla. The adult of C. midoriae is redescribed and the nymph is described for the first time. A key to the species feeding on S. japonica (adults and nymphs) is provided. The three species were observed to coexist on the same tree of S. japonica in northern Kyushu. Such coexistence of psylloid congeners is uncommon in Japan.     

Leaf Decomposition and Stream Macroinvertebrate Colonisation of Japanese Knotweed,an Invasive Plant Species

Japanese knotweed (Fallopia japonica Houtt. Ronse Decrane ) is a highly invasive exotic plant that forms monocultures in riparian areas, effectively reducing plant diversity. This change in riparian plant composition alters the allocthonous input of leaf litter into adjacent streams. A field experiment was completed to understand how leaf decomposition and macroinvertebrate colonisation associated with the incorporation of exotic leaf litter. Leaf packs of Japanese knotweed, native alder (Alnus incana L.), native cottonwood (Populus trichocarpa Torr . and Gray ), and two additional mixed pack types (alder and cottonwood; alder, cottonwood, and Japanese knotweed) were placed into a 50 m stream reach in Clear Creek, Idaho, and removed over a three‐month period. Leaf decomposition and macroinvertebrate assemblages were similar between leaf types, despite differences in nitrogen and phosphorus content. The diversity of leaf types within a given leaf pack also had no effect on leaf decomposition or macroinvertebrate dynamics. These findings suggest that allochthonous inputs of Japanese knotweed fulfill a detrital function similar to that of native leaf litter. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Some effects of pre-release host-plant on the biological control of Panonychus ulmi by the predatory mite Amblyseius fallacis

Amblyseius fallacis Garman has been selected for pyrethroid resistance and mass reared for experimental release as a biological control agent for tetranychid mites on a number of crops in Canada. Several releases of this predator onto apple and peach trees have failed to result in the establishment of A. fallacis, or in the biological control of Panonychus ulmi Koch. Here, we test the hypothesis that the change of host-plant at the time of release is a critical factor in the establishment of A. fallacis for biological control of P. ulmi. Functional and numerical response studies were undertaken on two populations of A. fallacis: a wild strain collected from the canopy foliage of an apple orchard near Vineland, Ontario; and a second strain reared on bean plants in a commercial insectary with Tetranychus urticae as prey. Each population consumed significantly more P. ulmi and produced significantly more eggs when on leaf disks from the plant species they were reared on, than on leaf disks from the novel host plant. A further experiment was conducted to determine if establishment and biological control of mass-reared A. fallacis could be affected by rearing a population for a short term on apple leaves prior to release on apple trees. Three release treatments were made into potted apple trees in a glasshouse, using predators commercially mass-reared on bean and T. urticae: A. fallacis released directly; A. fallacis reared in the laboratory for four weeks on bean and T. urticae; A. fallacis reared on apple leaves and T. urticae for four weeks. They were compared with a control treatment lacking predator release. Contrary to results of the functional and numerical response studies, no difference was observed between release treatments. All release treatments adding A. fallacis resulted in a similar, if limited, degree of biological control of P. ulmi. These results indicated that there may be short-term effects of host plant on the establishment of A. fallacis and biological control of P. ulmi, which in our study were observed as an initial reduction of the predatory response. However, in a test, the predators appeared to overcome these short-term effects and successfully established on the new host-plant to control P. ulmi.     

Intraguild interactions between the predator Macrolophus pygmaeus and the parasitoid Eretmocerus mundus,natural enemies of Bemisia tabaci

The native parasitoid Eretmocerus mundus Mercet and the predator Macrolophus pygmaeus Rambur are widely used to control Bemisia tabaci (Gennadius) in Mediterranean tomato greenhouses. An optimal biological control strategy for B. tabaci should take into account intraguild interactions between these natural enemies. In this study, predator's prey preferences and prey consumption were studied when offered different parasitoid and whitefly stages. The effect of the host plant on the adults of both natural enemies was also examined. M. pygmaeus preferred to consume B. tabaci over E. mundus when immature stages and adults of B. tabaci and E. mundus were offered. They consumed a larger amount of healthy B. tabaci nymphs and adults than of parasitised nymphs or E. mundus adults. The predator M. pygmaeus interfered with the reproduction of E. mundus females on cotton but not on tomato. However, B. tabaci nymphal mortality on tomato associated with parasitoid host feeding was also lower when the adult parasitoids coexisted with the predators. The joint release of M. pygmaeus and E. mundus adults did not increase the control of the whitefly B. tabaci.     

Factors affecting the efficacy of the leaf-spot fungus Mycosphaerella polygoni-cuspidati (Ascomycota): A potential classical biological control agent of the invasive alien weed Fallopia japonica (Polygonaceae) in the UK

Fallopia japonica, commonly known as Japanese knotweed, is an increasingly serious invasive alien weed in the UK and large parts of mainland Europe, as well as in North America. There is an urgent need to include classical biological control (CBC) into any integrated pest management strategy. The leaf-spot pathogen Mycosphaerella polygoni-cuspidati, a coevolved natural enemy of F. japonica present throughout its native Japanese range, is considered to have high potential as a CBC agent. In this study, the disease development of M. polygoni-cuspidati in the field and the optimum infection parameters under controlled conditions were investigated to elucidate the pathogen’s potential biocontrol efficacy against Japanese knotweed. Field observation in Japan showed that M. polygoni-cuspidati caused severe damage to its host plant. When sentinel knotweed plants from the UK were placed amongst naturally-infected field populations of F. japonica, disease incidence and severity were highest in July when monthly precipitation was also highest. In greenhouse inoculation tests, F. japonica was shown to be most susceptible at the young leaf stage (7–12 days after opening). Disease severity was highest after an initial dew period of 42–48 h, and severe defoliation followed inoculation at a temperature range of 15–25 °C. The optimum post-inoculation temperature after dew treatment for disease severity was 20–25 °C. In field inoculation tests, high disease incidence and severity indicate that the pathogen has the potential to control the plant effectively in the field. Humidity and temperature were shown to be the main factors influencing disease expression and lesion development of M. polygoni-cuspidati in a field situation. These results provide valuable information for any future use of M. polygoni-cuspidati as a CBC agent for management of Japanese knotweed in the UK.     

Prerelease efficacy test of the psyllid,Arytinnis hakani,a prospective biological control agent of the invasive weed Genista monspessulana

In weed biological control, conducting a prerelease efficacy test can help ascertain if prospective biological control agents will be capable of controlling the target plant. Currently, the phloem-feeding psyllid, Arytinnis hakani, is being evaluated as a prospective agent for the exotic invasive weed, Genista monspessulana, in the USA. Small potted plants were exposed to 0, 4, 8, 12 or 16 A. hakani second-instar nymphs which were allowed to develop on the plants for six weeks in an incubator at 18°C. Increasing A. hakani densities had a significant negative impact on the number of leaflets grown, per cent change in plant height and the final number of leaflets per centimeter of plant height. Increase in plant height and the number of leaflets relative to uninfested plants were reduced by 16 and 29%, respectively, at the highest infestation levels. Psyllids did not increase leaf senescence. One of the 10 plants that was exposed to 16 psyllids died, whereas none of the others died. Weekly nymphal mortality was 10–18% at the lowest density (four psyllids), but was at least 40% at the three highest densities. Although dead nymphs were replaced weekly, insect mortality may have reduced impact on the plants, especially at the higher densities. This insect is multivoltine, so prolongation of the infestation period may have a greater impact on the plant.     

Comparison of relative growth patterns of nymphal legs in relation to the oviposition habits of three heteropteran species

In most animals, morphological changes are closely correlated to changes in behavior during ontogeny. Females of the winter cherry bug, Acanthocoris sordidus (Thunberg), often oviposit away from host plants. First-instar nymphs of A. sordidus can develop to the second instar only with water. Thus, it is the second-instar nymphs that have to search for host plants when A. sordidus females oviposit off host plants, implying that movement is of greater importance during the second instar than during other instars. We investigated the relative growth pattern of nymphal legs of A. sordidus to determine whether the morphology of this species is adapted to changes in locomotion. For comparison, we examined the relative leg lengths of the nymphs of two species, Hygia opaca (Uhler) and Leptocorisa chinensis Dallas, that do not need to search for their host plants due to the species’ oviposition habits. As predicted, second-instar nymphs of A. sordidus had the longest relative leg length of all instars. On the other hand, the relative growth patterns of the nymphal legs of H. opaca and L. chinensis differed from that of A. sordidus, indicating that relative leg growth is more strongly affected by oviposition habits than by phylesis.     

From plant traits to invasion success: Impacts of the alien Fallopia japonica (Houtt.) Ronse Decraene on two native grassland species

Alien invasive plants threaten biodiversity, productivity and ecosystem functioning throughout the world. We examined the effect of Fallopia japonica on two native grassland species (Trifolium repens, Lolium perenne). We hypothesized that its negative effects on the native species are dependent on three mechanisms: (i) allelochemicals released and accumulated in soil with a history of invasion, (ii) altered soil biota and (iii) direct resource competition. We measured the response of the native species as the difference in their functional traits when grown under the three conditions. Our results demonstrate that neither allelochemicals nor soil biota from soil with history of F. japonica invasion had measurable effects on either species. Competition with the invader strongly reduced height, biomass and specific leaf area (SLA) of T. repens, while it had a lower effect on L. perenne. Furthermore, our results reveal that F. japonica took advantage of a positive plant–soil and plant–plant interaction. The results show that the prominent mechanism underpinning the invasion success of F. japonica in the grassland was the direct resource competition. This prominent role is also confirmed by the significant interactions between competition, allelochemicals and soil biota from soils with history of invasion of F. japonica on SLA of the native species.     

Development,survival and reproduction of Orius niger (Hemiptera:Anthocoridae) under different photoperiod and temperature regimes

The successful use of the predatory bug Orius spp. (Hemiptera:Anthocoridae) for the biological control of pests, namely western flower thrips Frankliniella occidentalis (Pergande), in greenhouses during winter depends on overcoming the obstacles of short photoperiods and low temperatures which limit the efficacy of the predators through diapause induction, slowed development, reduced survival and reproduction. Thus, research has focused on determining the insect species least vulnerable to these problems. This study investigated the effect of varying day-lengths (9, 11, 13 and 16 h) at 26°C as well as constant (18, 22, 26 and 30°C) and fluctuating (10/22°C) temperatures on biological characteristics of Orius niger (Wolff). Photoperiod did not induce reproductive diapause and did not significantly affect development, survival of eggs and nymphs, longevity, fecundity or sex ratio; with the exception of the 16 h day-length which led to a shorter duration of nymphal development. Fluctuating temperatures and a constant temperature of 18°C increased the incubation period, duration of nymphal development, total developmental time, generation time and longevity, compared to the constant temperatures of 22, 26 and 30°C, but decreased fecundity compared to 22 and 26°C. On the other hand, sex ratio was not influenced by 10/22 and 18°C. Egg hatch rate and nymph survival differed significantly between temperature regimes, being lower at 18 and 10/22°C. These results suggest that O. niger would be able to survive and reproduce under winter conditions in unheated Mediterranean greenhouses and is a candidate for use in the biological control of pests.