Parasitoids of obscure mealybug,Pseudococcus viburni (Hem.: Pseudococcidae) in California: establishment of Pseudaphycus flavidulus (Hym.: Encyrtidae) and discussion of related parasitoid species

To improve natural suppression of the obscure mealybug, Pseudococcus viburni (Signoret), the parasitoids Pseudaphycus flavidulus (Brèthes) and Leptomastix epona (Walker) (Hymenoptera: Encyrtidae) of Chilean origin were released in California's Central Coast vineyards from 1997 to 1999. A survey for parasitoids of P. viburni was conducted in the Edna Valley appellation wine grape region from 2005 to 2007, 6–8 years after classical biological control releases were discontinued. Two survey methods were used. First, field collections of obscure mealybugs from commercial vineyard blocks (2005–2007) and, second, placement of “sentinel mealybugs” on potted (1 L) grape vines (2006 only). From both survey methods, P. flavidulus was recovered, albeit levels of parasitism were low (less than 0.6%). We also placed longtailed mealybug, Pseudococcus longispinus (Targioni Tozzetti), on potted plants concurrent with placement of sentinel obscure mealybugs in the vineyard in order to measure parasitoid activity on this closely-related mealybug species. No P. flavidulus were recovered from P. longispinus. Other encyrtid parasitoids reared from either P. viburni or P. longispinus were Anagyrus pseudococci (Girault), Leptomastix dactylopii Howard, Leptomastidea abnormis (Girault), Coccidoxenoides perminutus Girault, and Tetracnemoidea peregrina (Compere). A hyperparasitoid, Chaetocerus sp., was also reared. The data are discussed with respect to biological control of vineyard mealybugs and newly developed controls for the Argentine ant, Linepithema humile (Mayr) (Hymenoptera: Formicidae). Because Pseudaphycus species reared from mealybugs are superficially very similar a taxonomic key and discussion of host relationships for selected Pseudaphycus species are provided.     

Density-Dependent Benefits in Ant-Hemipteran Mutualism? The Case of the Ghost Ant Tapinoma melanocephalum (Hymenoptera: Formicidae) and the Invasive Mealybug Phenacoccus solenopsis (Hemiptera: Pseudococcidae)

Although density-dependent benefits to hemipterans from ant tending have been measured many times, few studies have focused on integrated effects such as interactions between ant tending, natural enemy density, and hemipteran density. In this study, we tested whether the invasive mealybug Phenacoccus solenopsis is affected by tending by ghost ants (Tapinoma melanocephalum), the presence of parasitoids, mealybug density, parasitoid density and interactions among these factors. Our results showed that mealybug colony growth rate and percentage parasitism were significantly affected by ant tending, parasitoid presence, and initial mealybug density separately. However, there were no interactions among the independent factors. There were also no significant interactions between ant tending and parasitoid density on either mealybug colony growth rate or percentage parasitism. Mealybug colony growth rate showed a negative linear relationship with initial mealybug density but a positive linear relationship with the level of ant tending. These results suggest that benefits to mealybugs are density-independent and are affected by ant tending level.     

Spatial distribution of ants (Hymenoptera: Formicidae), vine mealybugs and mealybug parasitoids in vineyards

The mutualistic association between some ant species and honeydew‐producing Hemiptera has been shown to influence the distribution patterns and abundance of these hemipterans and their natural enemies. We studied the spatial distribution patterns of three ant species, mealybugs and mealybug parasitoids for two consecutive growing seasons on three wine grape farms in the Western Cape, South Africa. During the study period, no ant or mealybug controls were applied. Ant and mealybug monitoring was conducted on a total of 21 ha using a presence/absence sampling system, while parasitoids were collected from infested mealybug females. Spatial analysis by distance indices was used to analyse spatial distribution of insects and ArcView? was used to map the gap, patch and local association indices where significant association and disassociation occurred. Significant associations were found between some ants and parasitoids, while significant disassociations between the ants Crematogaster peringueyi and Linepithema humile; and also between Crematogaster peringueyi and Anoplolepis steingroeveri were found. Interspecific competition between ant species could play a role in the distribution of parasitoids and mealybugs. Our results stress the importance of monitoring for ants and mealybugs and further highlight the importance of restricted chemical applications against ants during the growing season.     

A survey of mealybugs and their natural enemies in horticultural crops in North Island,New Zealand,with implications for biological control

More than 1300 mealybugs and their natural enemies were collected from six crops (apples, pears, nashi, citrus, persimmon and grapes) at 91 sites. Pseudococcus longispinus and P. calceolariae were the commonest species in all crops, except in pipfruit in Hawkes Bay, inhabited almost exclusively by P. affinis. These three species accounted for more than 99% of all mealybugs collected. Mealybugs were attacked by 14 species of natural enemy, only two of which had been deliberately introduced to New Zealand. Six species of Encyrtidae were reared. Anagyrus fusciventris was recorded from New Zealand for the first time. Parectromoides varipes was newly identified as a primary parasitoid of mealybugs, and males of this species and Gyranusoidea advena, previously unknown, were found. Both species, together with Tetracnemoidea sydneyensis, T. peregrina and T. brevicornis, and Coccophagus gurneyi (Aphelinidae) and two species of Ophelosia (Pteromalidae) were widespread throughout the surveyed regions. Common predators included Cryptolaemus montrouzieri (Col: Coccinellidae), Cryptoscenea australiensis (Neuroptera: Coniopterygidae) and Diadiplosis koebelei (Dipt: Cecidomyiidae). Hyperparasitoids were extremely rare. Five species of ants were recorded tending mealybugs, but none is known to be disruptive to mealybug natural enemies. The implications of these data for biological control of mealybug pests in horticultural crops is discussed, and it is concluded that Pseudaphycus maculipennis (Hym: Encyrtidae) should be introduced against P. affinis. No other importations are recommended. The activity of existing species should be encouraged in future integrated pest management (IPM) programmes, by, for example, distributing A. fusciventris around the country, and commercializing the mass rearing and release of C. montrouzieri.     

Effects Of Tending By Solenopsis Invicta (Hymenoptera: Formicidae) On The Sugar Composition And Concentration In The Honeydew Of An Invasive Mealybug,Phenacoccus Solenopsis (Hemiptera: Pseudococcidae)

Mutualistic interactions between ants and hemipterans are mediated by the honeydew produced by the hemipterans. Previous works have demonstrated that the invasive mealybug Phenacoccus solenopsis produces abundant honeydew and attracts a large number of workers of the fire ant Solenopsis invicta. Mealybugs exhibit higher fecundity when tended by fire ants. The honeydew produced by P. solenopsis plays an important role in interactions between these two species. However, relatively few studies have focused on whether there is a cost to P. solenopsis mealybugs of being tended by S. invicta through changes in their excretion behavior and the quantity of honeydew produced. Our results indicated that the honeydew of P. solenopsis contains xylose, fructose, sucrose, trehalose, melezitose, and raffinose. The sugar concentration in the mealybug honeydew changed in an ant‐tended treatment. When tended by fire ants, the mealybugs generated honeydew with a significantly decreased xylose concentration. In contrast, the droplets showed a considerable increase in the melezitose concentration. P. solenopsis excreted honeydew more frequently when tended by S. invicta, but the weights of the droplets excreted by the ant‐tended mealybugs were significantly lower. In addition, S. invicta exhibited a significant preference for different sugars. Melezitose was visited more intensively than the other sugars in two choice tests. These results may suggest that, to attract more tending ants, mealybugs adjust their carbohydrate metabolism.     

Improving liquid bait programs for Argentine ant control: bait station density

Argentine ants, Linepithema humile (Mayr), have a positive effect on populations of mealybugs (Pseudococcus spp.) in California vineyards. Previous studies have shown reductions in both ant activity and mealybug numbers after liquid ant baits were deployed in vineyards at densities of 85-620 bait stations/ha. However, bait station densities may need to be <85 bait stations/ha before bait-based strategies for ant control are economically comparable to spray-based insecticide treatments-a condition that, if met, will encourage the commercial adoption of liquid baits for ant control. This research assessed the effectiveness of baits deployed at lower densities. Two field experiments were conducted in commercial vineyards. In experiment 1, baits were deployed at 54-225 bait stations/ha in 2005 and 2006. In experiment 2, baits were deployed at 34-205 bait stations/ha in 2006 only. In both experiments, ant activity and the density of mealybugs in grape fruit clusters at harvest time declined with increasing bait station density. In 2005 only, European fruit lecanium scale [Parthenolecanium corni (Bouché)] were also present in fruit clusters, and scale densities were negatively related to bait station density. The results indicate that the amount of ant and mealybug control achieved by an incremental increase in the number of bait stations per hectare is constant across a broad range of bait station densities. The results are discussed in the context of commercializing liquid ant baits to provide a more sustainable Argentine ant control strategy.     

Do additional sugar sources affect the degree of attendance of Dysmicoccus brevipes by the fire ant Solenopsis geminata?

Mutualistic interactions between ants and Hemiptera are mediated to a large extent by the amount and quality of sugar‐rich honeydew produced. Throughout the neotropics, the predaceous fire ant Solenopsis geminata (Fabricius) (Hymenoptera: Formicidae) is found in association with colonies of the pineapple mealybug, Dysmicoccus brevipes (Cockerell) (Hemiptera: Pseudococcidae), which they actively tend and protect from attack by natural enemies. In this study, we evaluate the effects of access to a sucrose solution on the mutualistic association between S. geminata and D. brevipes. Ten colonies of either species were established, with D. brevipes maintained on pumpkin, Cucurbita maxima Duchesne (Cucurbitaceae), in screen cages. Five of the S. geminata colonies were permitted access to vials with 20% sucrose solution and a pumpkin with 20 adult mealybugs. The remaining ant colonies were allowed access to mealybug‐infested pumpkins. Ant colonies with access to the sucrose solution attended mealybugs significantly less than those without additional sugar sources. Mealybug survival rates were similar under both treatments. Total body sugars and fructose were nearly twice as high in ants with access to honeydew and sucrose vs. those with access to honeydew and water. Fructose accumulated on the pumpkins over time in both treatments, suggesting that honeydew was not fully exploited by the ants. In conclusion, D. brevipes enjoy lower degrees of ant attendance when S. geminata have alternative sources of carbohydrates. We further discuss the significance of these findings for the conservation of predaceous ants and mealybug biological control.     

Ant–hemipteran association decreases parasitism of Phenacoccus solenopsis by endoparasitoid Aenasius bambawalei

1. Mutualism between ants and honeydew-excreting hemipterans is ubiquitous in the ecosystem. It is widely accepted that ant tending facilitates the colony growth of hemipterans by protecting them from predators and parasitoids. However, few studies have explored how ant tending helps defend against natural enemies. 2. Ghost ant Tapinoma melanocephalum and the invasive mealybug Phenacoccus solenopsis have a close mutual relationship. Previous studies have shown that ghost ant tending can definitely reduce parasitism and visit frequency of Aenasius bambawalei, the dominant endoparasitoid of P. solenopsis. However, the ghost ant workers seldom attack the parasitoids. It is still unclear how the ghost ant adversely affects parasitoids. This study explored the mechanism underlying the impacts of ants on natural enemies of the mealybugs. 3. Honeydew produced by P. solenopsis was an attractant to A. bambawalei. Parasitoids exhibited less searching activity, shorter longevity and lower parasitism when supplied with less honeydew. Aenasius bambawalei showed significant avoidance of pygidial gland secretions and visual cues of ghost ants. Parasitism in plants treated with 6-methyl-5-hepten-2-one, actinidine, and gland extracts was significantly lower than that in plants treated only with solvents (paraffin oil or double-distilled water). 4. It is concluded that honeydew consumption by ghost ants could negatively influence the performance of parasitoids. The pygidial gland secretions and visual cues of ghost ants also significantly inhibit the parasitism. These results may contribute to a better understanding of the regulation mechanism in ant–hemipteran–enemy interactions.     

Ant tending impairs performance of Aenasius bambawalei by manipulating the honeydew composition produced by Phenacoccus solenopsis

Honeydew produced by hemipterans is known as a possible kairomonal resource for parasitoids. The application of artificial honeydew effectively improves the performance of natural enemies. Aenasius bambawalei is a particularly dominant and aggressive endoparasitoid of the invasive mealybug Phenacoccus solenopsis. Our previous study showed that tending by the ghost ant Tapinoma melanocephalum significantly reduced the parasitism of A. bambawalei. We hypothesize that ghost ant tending influences host location of parasitoids by manipulating the composition of the honeydew produced by mealybugs. In this study, we tested whether the honeydew composition differs between treatments with and without ant attendance and whether changes in the honeydew influence the performance of A. bambawalei. Our results show that the sucrose concentration increased significantly in the ant‐attendance treatment but decreased when ant attendance was switched to an ant‐exclusion treatment; the inverse was true for the glucose concentration. Compared with the plastic honeydew treatment (mealybug with ant attendance), parasitoids spent much more time searching, had longer lifespans and showed higher parasitism on filter papers treated with natural honeydew (mealybug without any pre‐treatment) and those treated with convalescent honeydew (mealybug having experienced ant attendance and then switched to ant exclusion). These results support the hypothesis that ant tending influences the performance of parasitoids by manipulating honeydew composition.     

Influence of native ants on arthropod communities in a vineyard

• 1 Ants can have a range of effects on arthropods in crops, including suppressing herbivores such as caterpillars. However, ants can also increase hemipteran densities while reducing natural enemy numbers. In vineyard ecosystem, the effects of native ants and their interactions with other arthropods are poorly understood.
• 2 An ant‐exclusion experiment was designed to test the impact of native ants on both canopy and ground arthropods concurrently. The potential influence of ants on predation and parasitism of light brown apple moth (LBAM) eggs, a grape pest, was also examined. Adult grapevine scale insects and earwigs under bark were counted after a season of ant‐exclusion.
• 3 Among 23 ground ant species collected, six were found to forage in the canopy, with two Iridomyrmex species being the most commonly encountered.
• 4 There was no difference in the abundance of most arthropod orders and feeding groups between ant‐excluded and control vines, although ground spiders were more abundant under ant‐excluded vines, despite increased ground ant foraging pressure. LBAM egg parasitism and predation were low and probably affected by weather and other arthropods. Ant exclusion did not reduce survival of scale insects, although the distribution and abundance of scale insects were negatively associated with earwigs.
• 5 In conclusion, native ants did not consistently suppress arthropod assemblages, including natural enemies, and they did not promote the survival of scale insects. Interactions among native ant species within a vineyard might minimize their effects on other arthropods, although this needs further study.

     

Fire Ants Protect Mealybugs against Their Natural Enemies by Utilizing the Leaf Shelters Constructed by the Leaf Roller Sylepta derogata

The importance of mutualism is receiving more attention in community ecology. In this study, the fire ant Solenopsis invicta was found to take advantage of the shelters constructed by the leaf roller Sylepta derogata to protect mealybugs (Phenacoccus solenopsis) against their natural enemies. This protective effect of fire ant tending on the survival of mealybugs in shelters was observed when enemies and leaf rollers were simultaneously present. Specifically, fire ants moved the mealybugs inside the shelters produced by S. derogata on enemy-infested plants. Compared with that in plants without ants, the survival of mealybugs in shelters in the presence of natural enemies in plants with ants markedly improved. Both the protection of ants and the shelters provided by leaf rollers did not affect the survival of mealybugs in the absence of enemies in plants. Ants and leaf rollers significantly improved the survival of mealybugs in predator-infested plants, whereas no such improvement was observed in parasitoid-infested ones.     

Introduced ants reduce interaction diversity in a multi‐species,ant–aphid mutualism

Mutualisms contribute in fundamental ways to the origin, maintenance and organization of biological diversity. Introduced species commonly participate in mutualisms, but how this phenomenon affects patterns of interactions among native mutualists remains incompletely understood. Here we examine how networks of interactions among aphid‐tending ants, ant‐tended aphids, and aphid‐attacking parasitoid wasps differ between 12 spatially paired riparian study sites with and without the introduced Argentine ant Linepithema humile in southern California. To resolve challenges in species identification, we used DNA barcoding to identify aphids and screen for parasitoid wasps (developing inside their aphid hosts) from 170 aphid aggregations sampled on arroyo willow Salix lasiolepis. Compared to uninvaded sites, invaded sites supported significantly fewer species of aphid‐tending ants and ant‐tended aphids. At invaded sites, for example, we found only two species of ant‐tended aphids, which were exclusively tended by L. humile, whereas at uninvaded sites we found 20 unique ant–aphid interactions involving eight species of ant‐tended aphids and nine species of aphid‐tending ants. Ant–aphid linkage density was thus significantly lower at invaded sites compared to uninvaded sites. We detected aphid parasitoids in 14% (28/198) of all aphid aggregations. Although the level of parasitism did not differ between invaded and uninvaded sites, more species of wasps were detected within uninvaded sites compared to invaded sites. These results provide a striking example of how the assimilation of introduced species into multi‐species mutualisms can reduce interaction diversity with potential consequences for species persistence.     

Does the host plant affect the benefits from mutualisms? The invasive mealybug and ghost ant association

1. The benefits to trophobionte hemipterans are affected by the ant tending level, which is a widely accepted statement. The ant tending level is closely related to multiple factors. It is clear that the ant tending level can be affected by the temporal factor, age‐specific, the density of the hemipterans, and quantity and quality of honeydew produced by hemipterans. 2. Few studies of ant–hemipteran mutualisms have reported the patterns of host plants‐dependent effects, and whether host plants influence the ant tending level that is also unclear. As such, laboratory experiments were conducted to test whether the colony growth rate of an invasive mealybug Phenacoccus solenopsis Tinsley, parasitism of Aenasius bambawalei Hayat, an dominant parasitoid of P. solenopsis, are affected by tending by ghost ants (Tapinoma melanocephalum(Fabricius)], host plants (tomato and cotton), and interactions between the two factors. The difference in the ant tending level between the host plants was also determined. 3. The results showed that mealybug colony growth and parasitism were significantly affected by ant tending and host plant separately. There were significant interactions between the independent factors on the mealybug colony growth rate and percentage parasitism. These results suggest that benefits to mealybugs are host plant‐dependent.     

The effect of a mealybug infestation on the activity of amino acid decarboxylases in orchid leaves

Changes in the activity of lysine decarboxylase (LDC), tyrosine decarboxylase (TyDC), and ornithine decarboxylase (ODC) within orchid (Phalaenopsis × hybridum ‘Innocence’) leaves, infested by two mealybug species: Pseudococcus longispinus (Targ. Tozz.) and Pseudococcus maritimus (Ehrh.) were quantified. The pattern of changes was dependent on the insect species and duration of infestation. P. longispinus feeding increased LDC and TyDC activity after one week during the total period of observations. This species inhibited ODC activity after one week but increased later. P. maritimus decreased LDC activity in orchid leaves at all studied terms. TyDC action also went up during the first week of the infestation and was reduced after two weeks, while ODC was decreased after one day and induced later. The mechanism for the participation of analysed amino acid decarboxylases in local and/or systemic steps of orchid responses to mealybug infestation is discussed.     

Pseudococcus orchidicola (Hemiptera: Pseudococcidae), a newly found mealybug pest,confused with P. longispinus in Korea

Pseudococcus longispinus, a notorious cosmopolitan pest species of mealybugs, known to be distributed indoors in Korea since 2002, is found to be mixed with another species, Pseudococcus orchidicola. Finding P. orchidicola as a pest of tropical plants in Korea is rather unexpected because of their main distribution in the Pacific area and South Asia. However, all the available information from morphology, molecule and advice on identification from mealybug specialists indicates this is the best match for P. orchidicola. Morphological, molecular and some biological notes on P. orchidicola are provided with some adult and nymphal images, and compared with P. longispinus. A full discussion mainly on identity and distribution of P. orchidicola is given.     

Context‐dependence in an ant–aphid mutualism: direct effects of tending intensity on aphid performance

1. In ant–hemipteran mutualisms, ants receive carbohydrates in the form of honeydew, while hemipterans receive protection from natural enemies. In the absence of natural enemies, however, the direct effects of tending are generally less well known. We hypothesised that with increasing tending intensity (ant to aphid ratio), aphid performance would increase initially, then decrease at high tending levels due to the metabolic cost of producing high quality honeydew. 2. We tested our hypothesis in a greenhouse experiment by manipulating Argentine ant (Linepithema humile Mayr) colony size while holding constant the initial size of aphid (Chaitophorus populicola Thomas) aggregations. The two parameters associated with survival, aphid survivorship to maturity and longevity, declined with increasing tending intensity, whereas per capita birth rate and time to first reproduction showed no relationship to attendance. The intrinsic rate of increase declined only at relatively high tending levels, suggesting a nonlinearity in the effect of tending intensity. 3. Tending intensity measured in the experiment was similar to that observed in free‐living aggregations of C. populicola. Furthermore, the per capita recruitment rate of ants to free‐living aphid aggregations was negatively density‐dependent, indicating that small aggregations tend to experience the highest levels of tending intensity. This finding suggests that the aphid's intrinsic rate of increase may be positively density‐dependent, mediated by the aphid's mutualistic interaction with the ant. 4. In the Argentine ant–C. populicola interaction, experimental manipulation of colony size revealed a direct cost of ant attendance that was conditional upon tending intensity. Experiments that manipulate only ant presence or absence may yield an incomplete understanding of the mutualistic interaction if underlying nonlinearities exist.     

The influence of Lasius neoniger (Hymenoptera: Formicidae) on population growth and biomass of Aphis glycines (Hemiptera: Aphididae) in soybeans

In the United States, the soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), are often tended by the aphid-tending ant, Lasius neoniger Emery (Hymenoptera: Formicidae). In this study, we examined the effects of tending by ants on the density and biomass of soybean aphids on soybeans in Kentucky. We performed cage studies that limited access by ants and/or natural enemies. We used a split-plot design with natural enemy access as the main plot and ant attendance as the sub plot. We found that natural enemy access negatively affected aphid population density in the presence of tending ants, seen as a three- to four-fold increase in aphid density when natural enemies were excluded. In addition, we found that ant tending positively affected aphid biomass, both when natural enemies were given access to aphids or when natural enemies were excluded, seen by a two-fold increase in aphid biomass when ants tended aphids, both in the presence or absence of natural enemies. Biomass accumulation is seen as an important measurement for assessing aphid performance, and we argue that aphid-tending by ants can have an influence on natural field populations of soybean aphids. Agronomic practices that affect ant abundance in soybeans may influence the performance and hence pest outbreaks for this economically important pest.     

Survey of the natural enemies of Dysmicoccus mealybugs on pineapple in Hawaii

Surveys for mealybugs, associated natural enemies and ants were conducted in abandoned pineapple fields on the Hawaiian islands of Oahu and Maui from July 1992 to November 1993. Whole plant samples were taken, and mealybugs and ants found were identified. Mealybug-infested plant parts were isolated and held until natural enemies emerged from parasitized host material. At sample sites where only Dysmicoccus brevipes was present, its densities ranged from a mean of 23 to 157 mealybugs per plant, while in areas with mixed populations of this mealybug and Dysmicoccus neobrevipes Beardsley, densities ranged from a mean of 23 to 118 mealybugs per plant. Ants were present at all sample sites and on all dates. Pheidole megacephala (F.) was the most common ant species found. Anagyrus ananatis Gahan was the most common parasitoid reared. However, it attacked only D. brevipes, the dominant mealybug in the pineapple fields surveyed. Percent parasitization of D. brevipes by A. ananatis in the presence of ants ranged from 0.3 to 9.9%. Percent parasitization of D. brevipes and D. neobrevipes per plant by Euryrhopalus propinquus Kerrich ranged from 0.05 to 2.2%. Mean densities of the predators Lobodiplosis pseudococci (Felt), Nephus bilucernarius Mulsant and Sticholotis ruficeps Weise ranged from 0.05 to 5.75, 0.1 to 1.8, and 0.05 to 0.2 individuals per plant, respectively.     

Oviposition behaviour, influence of experience on host size selection, and niche overlap of the solitary Leptomastix epona and the gregarious Pseudaphycus flavidulus, two endoparasitoids of the mealybug Pseudococcus viburni

Oviposition behaviour and host size selection of the solitary parasitoid Leptomastix epona(Walker) and the gregarious Pseudaphycus flavidulus(Brèthes) [both Hymenoptera: Encyrtidae] were examined on five size classes of the mealybug Pseudococcus viburni(Signoret) [Hemiptera: Pseudococcidae]. The host size classes mostly consisted of one stage (first, second, third instar nymph, young adult and preovipositing adult) and were presented together to wasps of either parasitoid species. Both parasitoid species locate the host by drumming the surface of the patch with the antennae. Leptomastix eponaseems to use mainly the antennae to examine the host but P. flavidulusmay accept or reject a host for oviposition after antennation or insertion of the ovipositor. Leptomastix eponaattempts oviposition in all the host stages from second instar nymphs but P. flavidulusincludes first instar. Both parasitoid species select mainly larger hosts (>1 mm, third instar nymphs) to oviposit but P. flavidulusis able to parasitize more second instar nymphs compared to L. epona. Female wasps of L. eponamay host feed on small mealybugs (second and third instar nymphs) that they do not use for oviposition. Oviposition experience of either parasitoid species for 24 hours does not influence host size selection on patches with hosts of similar mixed sizes. Oviposition decisions are independent of the host sizes of the preceding ovipositions. Implications about stability of a single parasitoid – host system and the success of biological control of the mealybug were discussed in respect of the developmental refugia of the two parasitoid species. Niche overlap of the two parasitoid species was discussed with a view to giving an insight into a single or multiple introduction.     

Interactions between the ant Tapinoma nigerrimum (Hymenoptera: Formicidae) and the main natural enemies of the vine and citrus mealybugs (Hemiptera: Pseudococcidae)

Honeydew-excreting hemipterans, such as mealybug pests, can be protected from their natural enemies by tending ants in return for honeydew, thereby compromising the aims of biological control. In this respect, antagonistic interactions between the ant Tapinoma nigerrimum, native to the Mediterranean basin, and the main natural enemies of both the vine mealybug (VMB), Planococcus ficus, and the citrus mealybug (CM), Planococcus citri, were assessed in laboratory conditions. Parasitism of vine and CMs by their respective parasitoids, Anagyrus sp. nr. pseudococci and Leptomastix dactylopii, was negatively affected by the ant T. nigerrimum. Similarly, T. nigerrimum was shown to significantly disrupt the predatory potential of ladybird larvae, Cryptolaemus montrouzieri, when foraging on host CMs. By contrast, the presence of the ant did not negatively influence the predatory activity of C. montrouzieri adults when feeding on CMs. Consequently, the encyrtid parasitoids A. sp. nr. pseudococci and L. dactylopii and the larval stage of the predator C. montrouzieri may be considered as T. nigerrimum-sensitive, whereas the adults of C. montrouzieri may be regarded as T. nigerrimum-resistant predators. Accordingly, the ant T. nigerrimum constitutes a threat to the biological control of mealybugs by either the encyrtids A. sp. nr. pseudococci and L. dactylopii or the larval stage of the ladybird C. montrouzieri. Hence, adequate control of T. nigerrimum is highly recommended before any release of these mealybugs' natural enemies.