Key to the New Zealand species of Psyllaephagus Ashmead (Hymenoptera: Encyrtidae) with descriptions of three new species and a new record of the psyllid hyperparasitoid Coccidoctonus psyllae Riek (Hymenoptera: Encyrtidae)

Abstract  This paper records seven species of wasps in the genus Psyllaephagus (Hymenoptera: Encyrtidae) from New Zealand. All of these species are primary parasitoids of psylloids (Hemiptera: Psylloidea). Two are species previously described from New Zealand: P. acaciae Noyes and P. pilosus Noyes. Two are described Australian species which have established recently: P. bliteus Riek and P. gemitus Riek. Three new species are described here, from New Zealand: P. breviramus sp. nov., P. cornwallensis sp. nov. and P. richardhenryi sp. nov. All species are probably Australian in origin. A key to all seven Psyllaephagus species known from New Zealand is provided. An earlier first record of the Australian psyllid hyperparasitoid Coccidoctonus psyllae Riek (Hymenoptera: Encyrtidae), previously first recorded from New Zealand in 2006, is noted.     

Establishment of <Emphasis Type="Italic">Psyllaephagus parvus</Emphasis> and <Emphasis Type="Italic">P. perplexans</Emphasis> as serendipitous biological control agents of Eucalyptus psyllids in southern California

The free-living lemon gum psyllid, Cryptoneossa triangula Taylor, and the lerp-forming spotted gum psyllid, Eucalyptolyma maideni Froggatt (Hemiptera: Psyllidae) are invasive pests of eucalypts in California. In 2007, Psyllaephagus parvus Riek (Hymenoptera: Encyrtidae) was discovered parasitizing spotted gum psyllids and Psyllaephagus perplexans Cockerell was collected from lemon gum psyllids. While neither parasitoid species was purposely introduced, presence of the parasitoids was significantly associated with reduced intensity and duration of population peaks for both psyllid species. Spring peaks were reduced more than fall peaks. We estimated minimum rates of parasitism from the ratio of mummies to live nymphs. Higher parasitism was recorded in coastal than inland locations during the spring, while parasitism was similar for coastal and inland populations in the fall. Logistic regression models suggest parasitoids were the determining factor of psyllid population densities, although physical parameters, such as irrigation, may affect psyllid or parasitoid populations.     

Diapause strategy in the parasitoid Psyllaephagus pistaciae

The processes involved in the induction and termination of diapause in the parasitoid Psyllaephagus pistaciae Ferrière (Hymenoptera: Encyrtidae) were investigated under controlled laboratory conditions. Results indicated that the parasitoid P. pistaciae is able to overwinter successfully as a pupa within the mummified psyllid host Agonoscena pistaciae Burckhardt and Lauterer (Homoptera: Psylloidea), and the parasitoid clearly responded to a short photoperiod by entering diapause. The incidence of diapause increased significantly with decreasing temperature, indicating the existence of an interaction between low temperature and short‐day photoperiod that resulted in a 100% diapause. However, low temperature alone partly stimulated P. pistaciae to enter diapause. The incidence of diapause was likely to be independent from psyllid nymphal instar and the condition of its mother. This investigation showed that the termination of diapause in the overwintering parasitoid pupae and the duration of diapause completion or adult emergence is photoperiod independent, but dependent on temperature. However, chilling treatment was found to be unnecessary for diapause termination in P. pistaciae, although it decreased the duration of diapause maturation. The positive responses of parasitoids to environmental factors appears to act as a basic strategy to induce diapause in the parasitoid P. pistaciae against extreme climatic conditions.     

Behavioral responses of the parasitoid Psyllaephagus pistaciae (Hymenoptera: Encyrtidae) to host plant volatiles and honeydew

The behavioral responses of the parasitoid Psyllaephagus pistaciae, the major biocontrol agent of the common pistachio psylla, Agonoscena pistaciae, to volatiles emanating from its host plant and host honeydew, were examined using a four‐arm airflow olfactometer. In addition, the arrestment behavior of this parasitoid on clean and honeydew‐treated leaves of the pistachio, Pistacia vera, was monitored. The infested pistachio leaves were the most favored source of the volatile attracting the parasitoids. The parasitoid clearly distinguished and responded to infochemicals emitted by psyllid honeydew but at a lower level than to the volatiles from infested host plants. However, the searching time, locomotory behavior, antennal drumming and ovipositor probing were all affected when they encountered honeydew‐contaminated zones on pistachio leaves. These findings suggest that the psyllid honeydew releases kairomones that stimulate the parasitoids to greater searching activity, as well as providing a directional cue. The intensive searching activities in the presence of the volatiles tested were very similar to responses by the parasitoid females when encountering patches treated with psyllid honeydew. Such behavior could retain the parasitoid in a favorable area, thereby increasing the probability of additional host encounters.     

Host discrimination by the endoparasitoid Psyllaephagus pistaciae (Hymenoptera: Encyrtidae): A case of time-dependent ability

Host-discrimination behavior by the koinobiont parasitoid, Psyllaephagus pistaciae Ferrière (Hymenoptera: Encyrtidae), the major biocontrol agent of the common pistachio psylla, Agonoscena pistaciae Burckhardt & Lauterer (Hemiptera: Psyllidae), in Iran, was investigated in the laboratory. The results demonstrated that P. pistaciae quickly detected and avoided freshly parasitized hosts, either after antennating or by probing with their ovipositor. However, this discriminatory ability declined with time, probably influenced by both external and internal markers left by the previous ovipositing female. The results also confirmed that female P. pistaciae responded to changes in host quality associated with the parasitoid's larval development 4 days after the initial parasitization, clearly indicating that the second female could detect the presence of the larvae and adjust her host-selection decision. In addition, psyllid nymphs treated topically with a solution of Dufour's gland were rejected by the parasitoid, showing that Dufour's gland secretion had a significant effect as a host marking chemical. The current study also showed that superparasitism increased the host-mortality and that the rate of encapsulation decreased, suggesting that, when two eggs are laid in the same psyllid nymph, only one parasitoid develops to an adult.     

The biology of Psyllaephagus bliteus Riek (Hymenoptera: Encyrtidae), a parasitoid of the red gum lerp psyllid (Hemiptera: Psylloidea)

The red gum lerp psyllid, Glycaspis brimblecombei Moore (Hemiptera: Psylloidea), is native to Australia, where it feeds upon Eucalyptus species. It first appeared near Los Angeles, California, in 1998, and soon spread throughout the state. A biological control program directed against the psyllid was initiated and Psyllaephagus bliteus Riek (Hymenoptera: Encyrtidae) was imported from Australia and released in California. During quarantine screening, the taxonomic status of Psyllaephagus quadricyclus Riek was assessed by one of us (RLZ) and is proposed here as a new junior synonym for P. bliteus. The experiments discussed herein provide basic biological information on P. bliteus to supplement and improve the control program. We found that P. bliteus can oviposit into psyllid nymphs of any age but prefers third and fourth instars. Observations of host-handling behavior suggest that the large lerps of fifth instar psyllids increase host-handling time, thereby impeding oviposition and providing some protection from parasitism. Female P. bliteus were observed host-feeding on all psyllid nymphal development stages. Adults are relatively long-lived and, at constant temperatures of 17, 21, 23, 26, and 32 °C, longevity is a negative linear function of temperature. Females lived significantly longer than males. Adult females can live for several months, provided with hosts and held under glasshouse conditions (22 ± 3 °C), however, maximum egg deposition occurred within 22 days after adult emergence. Studies of larval development show that P. bliteus is a koinobiont and larval development is not initiated until the host reaches the late fourth or early fifth instar.     

Climate and the effectiveness of Psyllaephagus bliteus as a parasitoid of the red gum lerp psyllid

The encyrtid parasitoid Psyllaephagus bliteus Riek was introduced to California in 2000 to control the red gum lerp psyllid, Glycaspis brimblecombei Moore. Field experiments were conducted to determine the effect of regional summer climate (California coastal vs. interior valleys) on P. bliteus performance. Measurements of parasitoid performance, including reproduction and longevity, showed that P. bliteus was adversely affected by the extreme summer heat in the interior valley sites. The results suggest that regional climatic differences can contribute to the lower parasitism rates observed in California's warmer interior valleys. The results are discussed with respect to biological control of G. brimblecombei and the potential limitations of the parasitoid P. bliteus, as well as the possible impact of climate and intraguild predation as alternative hypotheses for the observed variable geographic performance of P. bliteus.     

Intraguild interactions between generalist predators and an introduced parasitoid of Glycaspis brimblecombei (Homoptera: Psylloidea)

Studies were conducted to evaluate potential impacts of generalist predators on the biological control of Glycaspis brimblecombei Moore (Homoptera: Psylloidea), a pest of eucalyptus trees in California, and its introduced parasitoid Psyllaephagus bliteus (Hymenoptera: Encyrtidae). Based on the results of the monitoring populations of G. brimblecombei and their natural enemies in inland and coastal California in the field, P. bliteus or Anthocoris nemoralis (Fabricius) (Heteroptera: Anthocoridae), alone or both species together were released into cages with G. brimblecombei to simulate predatory interference on the introduced parasitoid in the laboratory. Although coastal sites had higher densities of natural enemies as were percent parasitism and predation, the mean number of G. brimblecombei did not vary between inland and coastal sites. P. bliteus or A. nemoralis significantly reduced the psyllid densities in cages. However, when both species were together, the presence of A. nemoralis increased the parasitoid mortality relative to the mortality observed in the parasitoid-alone treatment. Moreover, the increase in parasitoid mortality was followed by the decrease in mortality of the psyllids. The current study also indicated that predation risk of parasitized hosts varies depending on the developmental stages of the psyllids.     

Limited protection of the parasitoid Pachycrepoideus vindemiae from Drosophila suzukii host‐directed spinosad suppression

Laboratory trials were conducted to determine whether the spotted wing drosophila, Drosophila suzukii (Matsumura) (Diptera: Drosophilidae), puparium can provide an effective physical barrier to protect immature stages of the pupal parasitoid Pachycrepoideus vindemiae (Rondani) (Hymenoptera: Pteromalidae) from spinosad treatments. Spinosad insecticides are currently an important suppression strategy for D. suzukii in organically managed fruit orchards although they are well known to cause mortality in hymenopteran parasitoids. High adult P. vindemiae female mortality (83%) occurred within 24 h of exposure to D. suzukii pupae treated with 10 mg a.i. l^?1 spinosad and female parasitoids did not avoid the pupae treated with similar low levels of spinosad in choice tests that included untreated pupae. Pachycrepoideus vindemiae develops as an idiobiont ectoparasitoid on host fly pupa within the sclerotized host puparium. Significant P. vindemiae survival and emergence was recorded when parasitized D. suzukii puparia were exposed to field treatment levels of spinosad; however, the parasitoid survival was dependent on the time of the spinosad treatment of the host post‐parasitization. Significant parasitoid survival occurred when the host puparia were treated at 2 weeks when the parasitoid was in the pupal stage but did not occur when the host puparia were treated at 1 week post‐parasitization, when the parasitoids were still in a larval stage. The parasitoid adults consumed or otherwise came in contact with residual degrading spinosad when they exited the treated host, and consequently high and low adult parasitoid mortality occurred when the adults emerged from puparia treated at 2 and 1 week(s), respectively. Our study indicates that generally the integration of P. vindemiae parasitism into a sustainable D. suzukii management program is not compatible with spinosad treatments, although P. vindemiae in the pupal stage inside sclerotized host puparia appear to be minimally impacted by spinosad treatments, provided that the spinosad degrades before parasitoid emergence.     

Occurrence of Ctenarytaina eucalypti (Maskell) (Hemiptera: Psyllidae) and its natural enemy Psyllaephagus pilosus Noyes (Hymenoptera: Encyrtidae) in Eucalyptus globulus in the state of Rio Grande do Sul, Brazil

Eucalyptus are hosts of several psyllid species. Among them, Ctenarytaina eucalypti (Maskell) is considered the most harmful to Eucalyptus globulus plantations. In Brazil, this pest has been observed in the state of Parana, and in 2005, it was found causing damage to plantations of E. globulus in Arroio Grande and Piratini cities, RS. The monophagous endoparasitoid Psyllaephagus pilosus Noyes was also observed parasitizing nymphs of C. eucalypti. This parasitoid has been shown to be highly efficient in controlling this psyllid species, making unnecessary the use of any other control method.     

The parasitoid complex of the citrus psyllaTrioza erytreae (Del Guercio) [Homoptera: Psyllidae]

Associated with the citrus psylla,Trioza erytreae (Del Guercio), in Salisbury, Rhodesia, are two primary parasitoids and 13 secondary and tertiary hyperparasitoids. One of the primary parasitoidsTetrastichus? radiatus remains external to the psyllid host whereas the other primary,Psyllaephagus pulvinatus, is an internal parasitoid. The major secondary hyperparasitoidAphidencyrtus cassatus, which was recorded in large numbers, attacks both primary parasitoids. Accounts are given of the biology of the two primary parasitoids and ofA. cassatus. The other hyperparasites in the complex are numerically unimportant and summaries of their biology are contained in a table. A diagram is given to illustrate the interrelationships of the parasitoids in the citrus psylla complex. It is concluded that, apart from phenological effects on the insects and host plant, the two primary parasitoids have considerable impact in controlling numbers of their host,T. erytreae, and that the secondary parasitoidA. cassatus through its attacks on the primary parasitoids is also of major importance in determining population levels achieved by the citrus psylla. Some characteristics of the parasitoids in the complex are discussed which might explain the high population levels of citrus psylla achieved in the Salisbury area. Several research avenues exist which would clarify the feasibility of biological control procedures againstT. erytreae.     

Natural enemies of the maize cob borer,Mussidia nigrivenella (Lepidoptera: Pyralidae) in Benin,West Africa

Mussidia nigrivenella Ragonot is a pest of maize cobs in West Africa. It significantly reduces maize yields and grain quality, with quantitative losses of 2-25%at harvest, and up to 10-15% indirect losses due to an increase in storage pest infestation levels. Infestation by M. nigrivenella also significantly increased the susceptibility of maize to Aspergillus flavus infection and subsequent aflatoxin contamination. Surveys conducted in different agro-ecological zones of Benin on cultivated and wild host plants during 1994-1997 revealed one egg parasitoid, three larval parasitoids and one pupal parasitoid attacking M. nigrivenella. Egg parasitism was scarce on all host plants sampled and in all four agro-ecological zones. Parasitism by larval and pupal parasitoids was usually less than 10%, and varied with host plant species. Both larval and pupal parasitoids were rare or absent in cultivated maize fields. The solitary chalcidid pupal parasitoid, Antrocephalus crassipes Masi, was the predominant species, contributing approximately 53% of the observed mortality. Logistic regression analysis indicated that this parasitoid was more prevalent on fruits of Gardenia spp. (Rubiaceae) than on the other host plant species including maize used by M. nigrivenella, and was most abundant between February and September. The differences in parasitoid diversity and parasitism between Benin and other regions suggest that there are opportunities for biological control through introduction of exotic parasitoids or using the 'new association' approach, which uses natural enemies of closely related host species that occupy similar ecological niches to the target pest.     

Seasonal biology and abundance of Psyllaephagus pistaciae,a biocontrol agent of the common pistachio psylla Agonoscena pistaciae

The solitary endoparasitoid Psyllaephagus pistaciae Ferrière (Hymenoptera: Encyrtidae), is the most widely distributed biological control agent of the common pistachio psyllid, Agonoscena pistaciae Burckhardt and Lauterer (Hemiptera: Psylloidea), in Iran. The pupation and overwintering sites of diapausing parasitoids and the psyllid were studied for 2 years using emergence traps in pistachio orchards in Rafsanjan, Iran. The psyllid mummies containing the overwintering parasitoid adhered to pistachio leaves and were carried on these leaves away from the tree when they latter senesced. The present results verified that plant litter which included dried grasses and old pistachio leaves tended to support a greater population of adult winter-form psyllid and psyllid mummies during the winter through early spring than other options examined. Adult parasitoids appeared in the field in early April, about 30 days after the emergence of adult psyllids, but almost at the same time as the hatching of the first generation psyllid nymphs in early April. Rates of parasitism of CPP were generally low throughout most of the year, ranging from 1 to 5%, but rose in late autumn to about 11%. Results suggest that the density of P. pistaciae is not great enough to keep pace with the psyllid populations in these orchards. They explain why growers consider it necessary to apply pesticides for this pest. However, this parasitoid undoubtedly does play an important role in the natural control of A. pistaciae late in the growing season, particularly in non-sprayed orchards. Conservation of these natural enemies should be one of the objectives in the development of sustainable pest management programs.     

Role of volatiles emitted by host and non-host plants in the foraging behaviour of Dentichasmias busseolae, a pupal parasitoid of the spotted stemborer Chilo partellus

The role of volatiles from stemborer host and non‐host plants in the host‐finding process of Dentichasmias busseolae Heinrich (Hymenoptera: Ichneumonidae) a pupal parasitoid of Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) was studied. The non‐host plant, molasses grass (Melinis minutiflora Beauv. (Poaceae)), is reported to produce some volatile compounds known to be attractive to some parasitoid species. The studies were conducted to explore the possibility of intercropping stemborer host plants with molasses grass in order to enhance the foraging activity of D. busseolae in such a diversified agro‐ecosystem. Olfactometric bioassays showed that volatiles from the host plants maize, Zea mays L., and sorghum, Sorghum bicolor (L.) (Poaceae), were attractive to the parasitoid. Infested host plants were the most attractive. Volatiles from molasses grass were repellent to the parasitoid. Further tests showed that volatiles from infested and uninfested host plants alone were preferred over those from infested and uninfested host plants combined with the non‐host plant, molasses grass. In dual choice tests, the parasitoid did not discriminate between volatiles from maize infested by either of the two herbivore species, C. partellus or Busseola fusca Fuller (Lepidoptera: Noctuidae). Volatiles from sorghum infested by C. partellus were preferred over those from C. partellus‐infested maize. The study showed that the pupal parasitoid D. busseolae uses plant volatiles during foraging, with those from the plant–herbivore complex being the most attractive. The fact that volatiles from molasses grass were deterrent to the parasitoid suggested that intercropping maize or sorghum with molasses grass was not likely to enhance the foraging behaviour of D. busseolae. Volatiles from the molasses grass may hinder D. busseolae's host location efficiency.     

Effects of parasitoid community structure upon the population dynamics of the honeysuckle leafminer,Chromatomyia suikazurae (Diptera: Agromyzidae)

Population dynamics of a leafminer,Chromatomyia suikazurae (Agromyzidae, Diptera) and its parasitoid community were studied for ten years at seven natural populations along an altitudinal gradient in Japan. This species which mines leaves of a forest shrub,Lonicera gracilipes (Caprifoliaceae), was attacked by 25 hymenopterous parasitoid species. Annually, the parasitoid community structure varied less within a population than among populations. The seven parasitoid communities were clustered into three groups corresponding to the altitudinal gradient: (a) lowland communities dominated by late-attacking, generalist pupal idiobiont eulophids and with highest species diversity, (b) hillside communities dominated by an early-attacking, specialist larval-pupal koinobiont braconid and (c) highland communities dominated by an early-attacking, generalist larval idiobiont eulophid. Annual changes of the host larval densities among the local populations were largely synchronous rather than cyclic. Among these populations, host density levels and mortality patterns greatly varied. By analyzing these inter-populational differences of host mortality patterns, the following conclusions were drawn: (1) The host mortality patterns were determined by the host utilization patterns of the locally dominant species. (2) The host pupal mortality but not larval mortality was related to species diversity but not to species richness itself of each parasitoid community. (3) Density dependence was detected only in pupal mortality at a lowland population dominated by late-attacking pupal parasitoids. These results suggest that interspecific interactions of parasitoids add additive effects to host population dynamics dissimilarly among local populations with different parasitoid communities.     

Effects of two insect growth regulators and a neonicotinoid on various life stages of <Emphasis Type="Italic">Aphytis melinus</Emphasis> (Hymenoptera: Aphelinidae)

The effect of two insect growth regulators and a neonicotinoid insecticide were tested on immature stages and adults of the parasitoid Aphtyis melinus DeBach (Hymenoptera: Aphelinidae), a key natural enemy of California red scale, Aonidiella aurantii (Maskell) (Hemiptera: Diaspididae), in California. No significant effects of the insect growth regulators on survival or development to the adult stage were found when the parasitoid was treated at any of the egg, larval, or pupal stages. The broad-spectrum neonicotinoid acetamiprid also showed no significant effect on the development of A. melinus to the pupal stage, probably because immature stages of this ectoparasitoid are protected under the cover of its armored scale host. However, 48 h exposure of adults to acetamiprid residues following emergence resulted in high levels of wasp mortality. Aphytis melinus adults treated with either of the two insect growth regulators as larvae survived 48 h exposure to pesticide residues as adults and showed levels of fecundity comparable with control insects. We conclude that the two insect growth regulators are compatible with augmentative releases of A. melinus but that treatments of acetamiprid should be avoided in situations where biological control by this parasitoid is important.     

Life history and demography of Psyllaephagus zdeneki (Hymenoptera: Encyrtidae), a potential candidate for biological control of olive psylla,Euphyllura pakistanica (Hemiptera: Psyllidae)

Life history and demographic parameters of Psyllaephagus zdeneki Noyes and Fallahzadeh (Hymenoptera: Encyrtidae) were studied on its host the olive psyllid, Euphyllura pakistanica Loginova (Hemiptera: Psyllidae). Experiments were conducted in a growth chamber at 20±1 °C, relative humidity of 60±5%, and a photoperiod of 16:8 (L: D) hours. Four different olive cultivars (Fishomi, Shenge, Oil and Yellow) were used to test possible host plant influence on parasitoid performance. The pre-imaginal developmental period of female P. zdeneki varied from 24.96 (on Fishomi) to 26.34 (on Shenge) days, and for males from 21.63 (on Fishomi) to 24.44 (on Yellow) days. Adult female longevity differed significantly among the four cultivars, ranging from 12.46 (on Fishomi) to 14.97 (on Shenge) days. For each cultivar, adult female longevity was significantly greater than male longevity. Life table parameters showed survival rates (l _x ) in newly emerged females were 84.61, 82.25, 85.71 and 78.12% on Fishomi, Yellow, Shenge and Oil, respectively. Female egg deposition was highest on Yellow (138.4 eggs per female) and lowest on Fishomi (116.3 eggs per female). The highest and lowest intrinsic rate of increase were 0.28 (on Shenge) and 0.24 (on Oil), respectively. The mean generation time ranged from 14.6 (on Shenge) to 16.7 (on Oil) days. These results are discussed with respect to the potential impact P. zdeneki as a natural enemy of E. pakistanica, the most important pest of olive in the Fars province of Iran, as well as the influence of olive cultivar on parasitoid life table parameters.     

Influences of plant species on life history traits of Cotesia rubecula (Hymenoptera: Braconidae) and its host Pieris rapae (Lepidotera: Pieridae)

The relative suitability of four plants was studied for larvae of Pieris rapae L. and its parasitoid Cotesia rubecula (Marshall). For unparasitized P. rapae, pupal dry weight and egg-pupa growth rate were higher on cabbage, radish and nasturtium than on Indian hedge mustard. Larval developmental rate and size were greatest for C. rubecula when its host was feeding on nasturtium. Wasp survival was not affected by the host insect/plant combination in which the parasitoid developed. These results indicate that the plant on which host larvae feed is an important factor in development of the parasitoid.     

Seasonal phenology and stage-specific parasitism of the apple ermine moth, Yponomeuta malinellus Zeller, in Korea

The apple ermine moth, Yponomeuta malinellus Zeller (Lepidoptera: Yponomeutidae), is a tent caterpillar that feeds on Malus spp. in Korea. Populations of the moth in native areas appeared to be regulated by the assemblage of parasitoids. Phenological associations between host stages and parasitoids, susceptible stage(s) of the host for each parasitoid, and stage‐specific parasitism were studied. The egg larval parasitoid Ageniaspis fuscicollis (Dalman) had highest parasitism of first instar larvae (24%), with 14% parasitism of other larval stages. Dolichogenidea delecta (Haliday) was recovered from all larval instars with the highest parasitism rate of second instar larvae (20.1%), followed by 19.9% parasitism of mid‐larval hosts. Herpestomus brunicornis Gravenhorst was reared from second instar larvae through to pupal collection, and had the highest parasitism rate (29.9%) at the pupal stage. The larval pupal parasitoid Zenillia dolosa (Meigen) was recovered from mid‐larval to pupal stages with the highest parasitism rate (5.5%) occurring in third to fourth instar larvae. The host stages for developing A. fuscicollis completely overlap with those of D. delecta, and with those of H. brunicornis to some degree. A statistically significant negative correlation exists between A. fuscicollis and these dominant parasitoids, indicating competitive interaction within the host.     

Superparasitism in Cotesia glomerata: response of hosts and consequences for parasitoids

Abstract.  1. Superparasitism occurs in Cotesia glomerata (Hymenoptera: Braconidae), a gregarious endoparasitoid of Pieris spp. (Lepidoptera: Pieridae). The response of P. brassicae larvae to superparasitism and the consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour.
2. Field surveys of a Swiss population revealed that C. glomerata brood sizes from P. brassicae larvae ranged from three to 158, and both the female ratio and the body weight of emergent wasps correlated negatively with brood size. In the laboratory, single oviposition on P. brassicae larvae did not produce any brood size larger than 62, but brood size increased with superparasitism.
3. Laboratory experiments demonstrated that both naive and experienced female wasps were willing to attack hosts that had been newly parasitised by themselves or conspecifics. Superparasitism reduced survivorship but increased food consumption and weight growth in P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae.
4. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater dry female mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitised hosts are rare in a habitat.