Temperature dependent functional response ofEncarsia formosa parasitizing the Poinsettia-strain of the cotton whitefly,bemisia tabaci, on Poinsettia

The control efficiency and performance ofEncarsia formosa Gahan (Hymenoptera: Aphelinidae) as influenced by the density of its host, the Poinsettia-strain ofBemisia tabaci Gennadius (Homoptera: Aleyrodidae), were investigated by laboratory experiments on Poinsettia (Euphorbia pulcherrrima Willd.).E. formosa showed a Type II functional response to fourth instar larvae ofB. tabaci, the response plateau increasing with temperature. A response model for randomly searching parasitoids incorporating temperature-dependent handling time and temperature-independent search rate was in accordance with the results, and gave an estimated search rate of 0.033 leaf·hour⁻¹ and handling times of 1.54, 2.86 and 20.1 h at 28°C, 22°C and 16°C, respectively. From the latter, the maximum number of hosts that can be parasitized at the three temperatures are 10.4, 5.6 and 0.8 larvae per day (provided the light period is 16 h). The number of hosts with ovipositor punctures was higher than the number of parasitized hosts, especially at 22°C and 28°C, implying thatE. formosa refrains from laying eggs in some of the hosts examined with the ovipositor. About 31% of the punctured larvae did not contain any eggs. Superparasitism occured during the experiment presumably originating from young, inexperienced parasitoids. Individual larvae were occasionally punctured several times, also by non-superparasitizingE. formosa. The resulting distribution of ovipositior holes was random, indicating thatE. formosa on the basis of antennal testing is unable to determine if a larva has previously been examined with the ovipositor. Almost fifty percent of the punctures were not followed by egg-depositions. Besides parasitizationE. formosa used hosts as food source. The number of hostfed larvae was independent of density, but varied with temperature being highest at 28°C (0.12 hostfed larvae per parasitoid per day).     

Host selection and survival of the parasitoidEncarsia formosa on greenhouse whitefly,Trialeurodes vaporariorum, in the presence of hosts infected with the fungusAschersonia aleyrodis

Successful control of greenhouse whitefly may be achieved by complementary activity of the parasitoidEncarsia formosa and the fungusAschersonia aleyrodis. One way to obtain an additive mortality effect of both entomopathogen and parasitoid would be achieved by the selection of healthy hosts by the parasitoid and rejection of fungus-infected hosts. Third and fourth instar larvae ofTrialeurodes vaporariorum which had been treated with a spore suspension ofA. aleyrodis 0, 4, 7, 10 or 14 days beforehand, were presented to female parasitoids. The parasitoids adopted the oviposition posture on untreated hosts as well as on treated hosts, irrespective of the different stages of infection in the hosts. However, significantly more hosts were parasitized byE. formosa in the control treatment than in the fungal treatment. The parasitoids offered treated hosts, showed rejection behaviour after probing on hosts showing detectable signs of infection (containing hyphal bodies or mycelium in the haemolymph). For instance, when hosts were offered seven days after spore treatment, the parasitoids showed an oviposition posture on a total of 83 (95.4%) out of 87 infected larvae, but laid only 4 eggs (4.6%). In contrast, on 48 (94.1%) out of 51 noninfected (or showing no detectable signs of infection) hosts an oviposition posture was adopted and 40 eggs (78.4%) were found after dissection. When infected hosts were encountered the oviposition posture lasted less than 1′40″ while rejection of non-infected hosts occurred after more than 1′40″. Other experiments were carried out offering treated hosts for 24 h to the parasitoids. The hosts were dissected afterwards. Again, significantly more eggs were laid in the non-infected hosts. When hosts were parasitized shortly after fungal spore treatment they were colonized by the fungus and the parasitoids did not develop. Transmission of the entomopathogen after probing infected hosts was observed to a limited extent. In conclusion,A. aleyrodis andE. formosa can be used together in a glasshouse situation. The parasitoid will be most effective when introduced more than seven days after application ofA. aleyrodis, because from that time onwards it is able to detect and reject fungus-infected hosts.     

Foraging behaviour of the whitefly parasitoidEncarsia formosa on tomato leaflets

IndividualEncarsia formosa parasitoids were observed continuously until the parasitoids flew away, either on clean tomato leaflets, on leaflets with honeydew, or on leaflets with unparasitized and parasitized whitefly larvae. Encounters with unparasitized and parasitized whitefly larvae, and contact with honeydew arrested the parasitoids on the leaflet. The walking speed increased linearly from 0.179 to 0.529 mm/s between 15 and 25–30°C. The walking activity showed another relationship with temperature: it was below 10% at 15 and 18°C, and increased to about 75% at 20, 25 and 30°C. It was not affected by host encounters or by 1 to 4 ovipositions. The total handling time of hosts was between 1.8–21.8% of the total time on the leaflet. Self-superparasitism was not observed. Conspecific-superparasitism did occur in 14% of the encounters with hosts containing a parasitoid egg, but was not observed anymore when the parasitoid egg had hatched. Experienced parasitoids superparasitized as often as naive females. The foraging behaviour ofE. formosa from landing on a leaf until departure has now been quantified and is discussed.     

Host-attacking behavior of a eulophid parasite,Kratochviliana sp., to the leaf mining host,Phytomyza ranunculi (Diptera: Agromyzidae) during its stay on the leaf

The present paper studies how the female parasite of Kratochviliana sp. visits and attacks its host larvae of Ranunculus leaf mining fly, P. ranunculi at a single leaf visit. The parasite visited its hosts at random on the leaf. The frequency of host visits was independent of the host density and the proportion of hosts survived from the parasite attack, in a leaf and its distribution was expressed as a single straight line. It almost always attacked living hosts at the first host visit after isolated from them for one day but with the rate of about 0.5 at the subsequent visits. In consequence, the relationships of the number of host attacks and killed hosts to the host density drew satulated curves in each. A model of host attack by this parasite at its single leaf visit was formulated by modifyingBakker et al.'s model (1972) basing upon these observations and the attack avoidance by the parasite to already attacked hosts previously reported.     

Host selection by the heteronomous hyperparasitoid Encarsia pergandiella: multiple-choice tests using Bemisia argentifolii as primary host

The ovipositional patterns of the heteronomous hyperparasitoid Encarsia pergandiella Howard (Hymenoptera: Aphelinidae) in the presence of its primary host Bemisia argentifolii Bellows & Perring (Hemiptera: Aleyrodidae), and in the presence or absence of conspecific and heterospecific secondary hosts (Encarsia formosa Gahan andEretmocerus mundus Mercet; Hymenoptera: Aphelinidae) were examined to assess host species preferences. Host preferences by heteronomous hyperparasitoids may affect the relative abundance of co-occurring parasitoid species and may influence host population suppression by the parasitoid community. Four combinations of hosts were tested: (1) B. argentifolii, E. mundus, and E. formosa, (2) B. argentifolii, E. formosa, and E. pergandiella, (3) B. argentifolii, E. mundus, and E. pergandiella, and, (4) B. argentifolii, E. mundus, E. formosa, and E. pergandiella. Arrays of hosts (24) were constructed in Petri dishes using leaf disks, each bearing one host. Thirty arrays of each host combination were exposed to single females for 6 h. All hosts were dissected to determine number of eggs per host. Encarsia pergandiella parasitized E. formosa hosts as frequently as E. mundus hosts. However, E. pergandiella parasitized either of these heterospecific hosts more frequently than conspecific hosts in treatments including two secondary host species. When a third parasitoid species was included in host arrays, E. pergandiella parasitized conspecific hosts as frequently as heterospecific hosts. Developmental stage of the hosts did not significantly influence host species selection by E. pergandiella. Our results indicate that host selection and oviposition by heteronomous hyperparasitoids like E. pergandiella, vary with the composition of hosts available for parasitization, and suggest a preference for heterospecific over conspecific secondary hosts.     

Influence of intrapatch experiences and temperature on the time allocation of the whitefly parasitoidEncarsia formosa (Hymenoptera: Aphelinidae)

The effect of experiences, such as contact with honeydew, rejections of hosts, and ovipositions in hosts, and of temperature on the time allocation of individualEncarsia formosa female parasitoids on tomato leaflets have been studied. Behavioral records were analyzed by means of the proportional hazards model. Analyses were carried out at two levels: (1) the tendency of leaving and (2) the tendency of changing from one leaf side to another. The patch-leaving behavior ofE. formosa can be described by a stochastic threshold mechanism, which is characterized by a certain tendency (probability per time) to leave. The median time from being placed on the leaflet or, if it occurred, from the latest encounter with a host until leaving was 18.6 min. The median time for changing from one leaf side to the other was initially 11.6 min and dropped to 5.7 min after both leaf sides had been visited. The effect of temperature, ranging from 20 to 30°C, was negligible. The presence of honeydew as well as the first oviposition in an unparasitized host decreased the tendency to leave, thus increasing the giving up time (GUT) since the latest encounter with a host. Encounters with parasitized hosts did not affect the GUT since latest encounter; as a result, the total residence time increased. After the first oviposition in an unparasitized host the tendency of changing from the lower leaf side on which hosts were present to the upper side was decreased. The presence of honeydew did not affect the tendency of changing leaf sides.     

Influence of Hairiness ofGerbera jamesoniiLeaves on the Searching Efficiency of the ParasitoidEncarsia formosa

To characterize the relationship between the leaf surface ofGerbera jamesoniiHooker cultivars and the searching behavior of the parasitoidEncarsia formosaGahan on a leaf, the trichome density and shape were described, and the walking behavior was quantified. Leaf hair density varied from 80 to more than 1000 trichomes/cm²and the hair shape varied from single erect trichomes to tapestries of entangled trichomes above the leaf surface. The walking activity, speed, and pattern of the parasitoid were tested on leaves with different hair density and shape. In spite of the large differences in leaf surface structure, the walking activity was similar and around 75% on most cultivars. The walking speed was between 0.2 and 0.3 mm/s on all cultivars and was not significantly different from the speed on tomato. On hairless sweet pepper leaves the walking speed is much higher (0.73 mm/s), so the hairs onGerberaleaves do hamper parasitoid females and result in a strong reduction of the walking speed, but the variation in hair density and shape onGerberadoes not lead to differences in walking activity, pattern, and speed. It appears that the hairs ofGerberaare less of an obstacle forE. formosathan the stiff, large hairs occurring in a high density on cucumber, where the walking speed of the parasitoid is less than 0.2 mm/s. A rise of temperature of 5°C gave a significantly higher walking speed (0.39 mm/s) ofE. formosafemales on all cultivars tested. The relative straightness of the walking track was high and the same on all cultivars. Consequences of the results of the searching behavior ofE. formosaonG. jamesoniicultivars are discussed with respect to other host plants. As parasitoid walking speeds are the same onGerberaand tomato, and whitefly reproduction is also similar on these two host plants, we conclude that biological control of whiteflies onGerberais a realistic option. This conclusion is confirmed by the strong increase of commercial biological control onGerberawhich occurred during the past 5 years.     

Nondevelopment ofCotesia kariyai(Hymenoptera: Braconidae) in Entomopoxvirus-Infected Larvae ofPseudaletia separata(Lepidoptera: Noctuidae)

Interaction between an entomopoxvirus (PsEPV) and a gregarious braconid endoparasitoid,Cotesia kariyai,inPseudaletia separatalarvae showed that infection of larvae with PsEPV was deleterious to the development and survival ofC. kariyai.The survival and development ofC. kariyaiin PsEPV-infectedP. separatalarvae depended on the length of time between parasitization and viral infection. No parasitoid larvae emerged from PsEPV-infected hosts when host larvae were exposed simultaneously to parasitization and PsEPV inoculation whereas more than 80% of the hosts produced parasitoids when PsEPV was administered 5 days postparasitization.C. kariyailarvae in PsEPV-infected hosts showed a retarded development, shrank, and died about 8 days after viral exposure. Virion-free plasma from PsEPV-infectedP. separatalarvae was toxic to the parasitoid larvae even up to a dilution level of 32 when it was injected intrahemocoelically into the host larvae. Development of parasitoids in hosts that were simultaneously parasitized and injected with the virion-free plasm never progressed beyond the egg stage. The parasitizedP. separatalarvae injected with the virion-free plasma did not pupate and died within 30 days after injection.     

GENETIC VARIATION IN ASTERIONELLA FORMOSA (BACILLARIOPHYCEAE): IS IT LINKED TO FREQUENT EPIDEMICS OF HOST‐SPECIFIC PARASITIC FUNGI?1

Understanding of the genetic basis for susceptibility and resistance is still lacking for most aquatic host–parasite systems, for instance, for phytoplankton and their fungal parasites. Fungal parasites can have significant effects on phytoplankton populations, mainly through their ability to decimate algal host populations during epidemics. We used random amplified polymorphic DNA (RAPD) and amplified fragment length polymorphism (AFLP) analysis to study levels of genetic variation within a population of the freshwater diatom Asterionella formosa Hassall in relation to parasitism by the obligate, host‐specific, fungal parasite Zygorhizidium planktonicum Canter. The level of genetic variation within the A. formosa population in Lake Maarsseveen, The Netherlands was found to be high despite the presumed absence or very low frequency of sexual reproduction in this species, the limited gene flow, and the severity of parasite attack that would purge the population from susceptible genotypes. RAPD analysis revealed four distinct banding patterns, with 3 of 21 markers (14%) being polymorphic. In AFLP analysis, every single isolate of A. formosa showed a unique banding pattern, and 120 of the 210 AFLP markers (57%) were found to be polymorphic. Furthermore, character compatibility analysis revealed that sexual reproduction may be one of the mechanisms that generates and maintains genetic variation in the A. formosa population in Lake Maarsseveen. The presence of genetic variation in A. formosa was reflected in infection experiments, which showed that genetically different A. formosa strains differed in their susceptibility to various Z. planktonicum strains and that parasite strains differed in their ability to infect particular host strains.     

Pre- and post-landing response of the parasitoid Encarsia formosa to whitefly hosts on Gerbera jamesonii

One of the factors that may complicate biological control of the greenhouse whitefly on Gerbera jamesonii by Encarsia formosa is the rosette shape of this ornamental, which differs from the vertical shape of most vegetable plants (cucumber, egg plant, tomato, etc.). Therefore, host-habitat location and the behaviour prior to landing on uninfested and infested leaves was studied. Attraction of E. formosa from a short distance by infested leaves could not be detected: the parasitoid females landed at random on uninfested and infested leaves. After the first landing, a redistribution of the wasps occurred on the leaves. After 24 h three times as many wasps were found on the infested leaves than on uninfested ones. In a dispersal experiment with four plants, E. formosa appeared to have no preference for landing on leaves of the medium age class, which is the age class on which most of the whiteflies in a suitable stage for parasitism occur. Twenty percent of the parasitoids were found on the plants 20 min after releasing them. These results were independent of the plant cultivar and the host density on the plants. In the course of 8 h, the number of E. formosa females recovered from plants increased linearly, and this increase was greater on plants where hosts were present and also greater on the plant cultivar with the lowest trichome density. After 24 h, the percentage of females was highest (56%) on plants with the highest host density. E. formosa females were arrested on leaves where hosts were present. Contrary to our expectation, the results from the two G. jamesonii cultivars that differed strongly in leaf hairiness were not significantly different in most experiments. Only at the high host density was parasitism found to be lower on the cultivar with the higher hair density. Parasitoids may walk on top of the `hair coverlet' of cultivars with high trichome density and, therefore, be hampered less than expected.     

The effect of nitrogen fertilizer applied to Euphorbia pulcherrima on the parasitization of Bemisia argentifolii by the parasitoid Encarsia formosa

More wasps of Encarsia formosa Gahan (Hymenoptera: Aphelinidae) were found on fertilized poinsettias, Euphorbia pulcherrima (Willd.) (Euphorbiaceae), than on non-fertilized plants. Parasitization of Bemisia argentifolii Bellows & Perring (Homoptera: Aleyrodidae) by E. formosa was higher on plants treated with calcium nitrate than with ammonium nitrate or on control plants. In a no-choice test, host feeding by E. formosa was higher when hosts were on fertilized plants than when hosts were on control plants. The nitrogen content of whitefly pupae reared on plants treated with ammonium nitrate was higher than those on calcium nitrate-treated plants.Variability in the parasitization of B. argentifolii by E. formosa appears to be due to host plant-mediated differences in the whiteflies. E. formosa may be influenced by the nutritional suitability of the host, which influences whether wasps continue to oviposit, feed, or disperse.     

The effect of population density on the reproduction ofTrichogramma japonicum Ashmead (Hymenoptera: Trichogrammatidae)

When a fixed number of the hosts, the eggs of the almond moth were exposed experimentally to various numbers of the parasites, Trichogramma japonicum, the following changes were observed with increasing parasite density:

1. The percentage of parasitism rises and approaches to 100 with gradually diminishing rate.
2. The number of parasite progeny increases and reaches a maximum, then decreases gradually.
3. The number of eggs laid per parasite female decreases gradually.
4. The proportion of hyperparasitized hosts progressively rises. The frequency distribution of parasite eggs in a host is of an intermediate type between random and uniform.
5. The competition among parasite larvae becomes severe. The progressive rise in mortality, the declining percentage of females in progeny and the emergence of stunted adults at the higher densities are observed.

In connection with both the nature of the parasitizing behaviour of adult and that of the competition among larvae, the nature of the density effect on the parasite population was discussed.     

Egg Size, Intrinsic Competition, and Lethal Interference in the Parasitoids Encarsia pergandiella and Encarsia formosa

Recent population dynamic theory predicts that disruption of biological control may occur when one parasitoid species' superiority in intrinsic competition is associated with a lower ability to find and exploit hosts (i.e., ability in extrinsic competition). One might expect such a trade-off, for instance, if parasitoids with larger (and fewer) eggs are more likely to prevail in intrinsic competition than species with smaller (and more numerous) eggs. We tested the idea that relative egg size could be used to predict the outcome of intrinsic competition in two closely related endoparasitoids, Encarsia pergandiella Howard and Encarsia formosa Gahan. Contrary to expectation, the parasitoid species with smaller eggs, E. pergandiella, prevailed in intrinsic competition, regardless of the order that hosts were exposed to the two species. In a literature survey, we found four studies of competing pairs of endoparasitoid species for which: (a) egg size estimates were available and (b) one species was consistently superior in intrinsic competition. In three of the four studies, the small-egged species prevailed in intrinsic competition, as we also found. Although E. formosa lost in intrinsic competition, this species negatively affected E. pergandiella's progeny production by host feeding on and killing hosts containing E. pergandiella eggs. E. formosa females also host fed on conspecific-parasitized hosts. As a mechanism of both intra- and interspecific interference competition, host feeding on parasitized hosts contradicts assumptions about the nature of interference competition in existing population dynamics models.     

Interactions between the ParasitoidAmetadoria misella(Diptera: Tachinidae) and the Granulovirus ofHarrisina brillians(Lepidoptera: Zygaenidae)

Interactions between a granulovirus (HbGV), a tachinid parasitoidAmetadoria misella,and their host, the western grapeleaf skeletonizerHarrisina brillians,were investigated. In field populations, the occurrence ofA. misellain HbGV-infectedH. brillianspupae was less frequent than would have been expected by random assortment of the virus and the parasitoid. Furthermore, enzyme-linked immunosorbent assay detected granulovirus less frequently and in lower concentrations in parasitized pupae than in nonparasitized pupae. Finally, in the host pupae that tested positive for virus, parasitoids were more likely to survive pupation than hosts. When laboratory-rearedH. brillianslarvae were exposed to naturally occurringA. misellain a field experiment, the parasitoid oviposited more often in older than in younger host larvae and more often in healthy than in HbGV-infected host larvae. These results are consistent with the hypothesis that selective oviposition byA. misellaleads to reduced overlap of the parasitoid and HbGV in hosts, resulting in greater parasitoid survival.     

Influence of Clavicipitaceous Endophyte Infection in Ryegrass on Development of the ParasitoidMicroctonus hyperodaeLoan (Hymenoptera: Braconidae) inListronotus bonariensis(Kuschel) (Coleoptera: Curculionidae)

Laboratory experiments were conducted to examine the effect of ryegrass infection by the endophytic fungusAcremonium loliiLatch, Christensen and Samuels onMicroctonus hyperodaeLoan, a parasitoid ofListronotus bonariensis(Kuschel). Progression of parasitoids through the larval instar stages was shown to depend on adequate nutrition of the weevil host. Compared to confinement on endophyte-free ryegrass, parasitized weevils held on nonpreferred diets comprising leaf segments from endophyte-infected ryegrass and switchgrass contained parasitoid larvae with retarded development. Similarly, development of parasitoid larvae was retarded in hosts feeding on artificial diet containing diterpenes and alkaloids ofA. loliiorigin. Several diterpenes incorporated into the diet reduced survival of the parasitoid larvae. Attack rate of parasitoids was reduced when the quality of potential host weevils was compromised by confinement on nonpreferredA. lolii-infected ryegrass or without food for 14 days.     

Analysis of foraging behavior of the whitefly parasitoidEncarsia formosa (Hymenoptera: Aphelinidae) in an experimental arena: A simulation study

Foraging ofEncarsia formosa was analyzed using a stochastic simulation model of the parasitoid's behavior. Parasitoids were allowed to search during a fixed time in an experimental arena with immatures of the greenhouse whitefly,Trialeurodes vaporariorum. The model simulates host searching, selection, and handling behavior and the physiological state (egg load) of the parasitoid. The simulated number of hosts encountered, parasitized, or killed by host feeding agreed well with observations on leaf disks. The hypothesis of random host encounter seems to be correct. The number of ovipositions on the leaf at low host densities was strongly affected by the parasitoid's walking speed and walking activity, the probability of oviposition after encountering a host, and the initial egg load. At high densities, the initial and maximum egg load were most important. A strong temperature effect was found at 18°C or below. The number of encounters, ovipositions, and host feedings increased with host density to a maximum of 25, 6.5, and 1.5, respectively, during 2 h at 25°C. The shape of the curves resembled a Holling Type II, which may be the result of the experimental procedure, where a parasitoid was confined to a patch during a fixed time.     

The area of discovery ofApanteles glomeratus (Hymenoptera: Braconidae),Pteromalus puparum (Pteromalidae) andBrachymeria regina (Chalcididae)

The areas of discovery ofApanteles glomeratus, Pteromalus puparum andBrachymeria regina were calculated using two different models. Increasing host or parasite density generally resulted in an initial increase followed by a decrease in area of discovery.     

Host-age discrimination during host location byCotesia glomerata, a larval parasitoid ofPieris brassicae

Some parasitoids are restricted with respect to the host stage that they attack and even to a certain age within a stage. In this paper we investigate whether the parasitoidCotesia glomerata can discriminate between old and young caterpillar instars of its host,Pieris brassicae, before contacting these hosts, since contacts with older instars are very risky with a chance of being killed, due to the aggressive defensive behaviour of the caterpillars. Flight chamber dual choice tests showed that volatile chemicals emitted by Brussels sprouts plants (Brassica oleracea var. gemmifera) after feeding damage by 1st and 5th larval instars are equally attractive to the wasps. Simulated herbivore damage by 2nd and 5th larval instars, obtained by treating mechanically damaged leaves with carterpillar regurgitant, was also equally attractive, even when the wasps were exposed to repeated experience on different larval instars to increase their discriminatory ability. In contrast, single choice contact bioassays showed that the time spent searching on a leaf with feeding damage of 1st instar larvae was significantly longer than the time spent on 5th instar feeding damage or on mechanically damaged leaves. Both flight and contact bioassays did not show any effect of egg-related infochemicals. The results demonstrate thatC. glomerata can discriminate between young and old larval instars ofP. brassicae, without contacting the caterpillars. This is not done through volatile herbivore-induced synomones but through cues that are contacted after arrival at a caterpillar-infested leaf.     

Poinsettia (Euphorbia pulcherrima Willd Ex Koltz) Cultivar-Mediated Differences in Performance of Five Natural Enemies of Bemisia argentifolii Bellows and Perring, n. sp. (Homoptera: Aleyrodidae)

Laboratory evaluations of five natural enemies of the silverleaf whitefly, Bemisia argentifolii Bellows and Perring, n. sp., were conducted to determine their potential as biological control agents in greenhouse poinsettia ranges. Adult longevity, prey consumption or host feeding and parasitism rates, and parasitoid emergence were measured for one predator, Delphastus pusillus LeConte, and four parasitoids, Encarsia formosa Gahan, Encarsia luteola Howard, Encarsia pergandiella Howard, and Encarsia transvena (Timberlake), as possible indicators of efficacy. Characterization of each parameter was performed on two poinsettia cultivars: the first, ′Annette Hegg Brilliant Diamond,′ has trichome densities on the leaf undersurfaces approximately 15% less than the trichome densities on the leaf undersurfaces of the second cultivar, ′Lilo.′ Adult longevity varied significantly between natural enemies (ranging from an average high of 85.2 days for female D. pusillus feeding on B. argentifolii nymphs to an average low of 2.8 days for the Canada colony of E. formosa), but not between cultivar. Prey consumption and oviposition by D. pusillus varied between prey type (nymphs consumed > eggs consumed) and poinsettia cultivar (′Annette Hegg Brilliant Diamond′ > ′Lilo′). Host feeding, parasitism and total number of B. argentifolii nymphs killed varied significantly among Encarsia spp., but no single wasp performed better than the rest across all three parameters. Host feeding, parasitism, and total number of nymphs killed were greater on ′Annette Hegg Brilliant Diamond′ than on ′Lilo′ and this difference was consistent among the four parasitoid species. Among parasitoid species differences in percentage emergence were consistent between the two poinsettia cultivars with emergence from parasitized nymphs on ′Lilo′ being greater than emergence on ′Annette Hegg Brilliant Diamond.′ Results from these evaluations suggest that the probability of achieving successful augmentative biological central will be greater on poinsettia cultivars with fewer trichomes. In addition, achieving biological control is likely to be difficult with releases of E. transvena, but a greater chance for success may be possible through releases of D. pusillus when whitefly densities are high or through releases of E. formosa (Beltsville colony) or mated E. pergandiella independent of whitefly densities.     

Time allocation of the parasitoidAphidius colemani (Hymenoptera: Aphidiidae) foraging forAphis gossypii (Homoptera: Aphidae) on cucumber leaves

The time allocation of individualAphidius colemani female parasitoids foraging forAphis gossypii nymphs on cucumber leaves has been investigated. Apart from experiences on the current leaf (such as density of hosts on the current leaf, density of hosts on a neighboring leaf and encounters with hosts on the current leaf), the effect of previous leaf visits on the time allocation was studied. Behavioral records were analyzed by means of the proportional hazards model, to determine the tendency of leaving the current leaf. The leaving tendency decreased only on leaves with a high host density (100 aphids), thus increasing the giving up time since the latest encounter. Rejection of aphids had no influence on the leaving tendency. To assess the effect of the number of hosts encountered on the leaving tendency, we considered three classes: 0–30 encounters, 31–100 encounters, and 100 or more encounters with hosts. The effect of the number of hosts encountered differed at different aphid densities. When fewer than 10 aphids were present the leaving tendency was much greater after 30 encounters than beforehand. At a density of 100 aphids the leaving tendency was lower than at the other aphid densities and increased only after 100 encounters. The density of hosts on a neighboring leaf, ranging from 0 to 100 hosts, had a negligible effect on the leaving tendency. Repeated visits to leaves with 10 unparasitized aphids resulted in an increase in the leaving tendency after 10 visits. It is argued that the parasitoids have some innate expectancy of host availability and that they concentrate on high-density patches.