Leaf age preference for oviposition by three species of whitefly on lemon

The leaf age preferences for oviposition by three species of whitefly (Dialeurodes citri [Ashmead],Aleurothrixus floccosus [Maskell], andParabemisia myricae [Kuwana]) were tested on lemon,Citrus limon (L.). Four categories of leaf age were tested (I: incompletely expanded, growing leaves; II: young leaves that are completely expanded but not yet dark green; III: completely mature leaves [dark green]; IV: completely mature leaves judged to be older than III as determined by the age of the twigs to which they were attached). Paired choice experiments were conducted in the laboratory using excised leaves and in the field using intact leaves. In both laboratory and field tests,P. myricae preferred I over all other leaf ages. It rarely oviposited on II and never oviposited on III or IV, even when they were the only two choices. The order of preference forD. citri in laboratory and field tests was II>I>III>IV. In laboratory tests,A. floccosus exhibited the same order of preferences asD. citri, but in the field, II was preferred over all others, III was slightly preferred over IV, and I varied greatly among replicates in comparisons against III or IV. In related experiments, the choice between I & III byA. floccosus andD. citri was not affected by confining the whiteflies to either I or III for three days prior to testing.

---

Sélection pour la ponte des feuilles deCitrus limon en fonction de l'âge par trois espèces d'aleurodes

Résumé Des feuilles de 4 tranches d'âges appartenant àCitrus limon L.: 1) en cours de croissance et incomplètement développées; 2) développées mais pas encore vert sombre; 3) complètement développées (vert sombre) 4) plus âgées que le lot 3 d'après l'âge du rameau, ont été proposées pour pondre à 3 espèces d'aleurodes (Dialeurodes citri Ashmead,Aleurothrixus floccosus Maskell,Parabemisia myricae Kuwana). Les femelles étaient isolées dans des cages dont les ouvertures sur deux côtés opposés étaient obstruées par la face inférieure de feuilles de citronniers d'âges différents. Les feuilles utilisées au laboratoire étaient coupées, les expériences dans la nature étaient faites avec des feuilles intactes. Récoltées au bout de 2 jours, leurs ufs étaient dénombrés.Dans les 2 cas,P. myricae a préféré significativement le lot 1, les pontes sur lot 2 étaient rares, et nulles sur les lots 3 et 4 même en l'absence de choix. Les préférences significativement différentes deD. citri au laboratoire et dans la nature étaient les suivantes: 2>1>3>4.A. floccosus a présenté au laboratoire les mêmes préférences queD. citri (P0,0004), mais certains résultats étaient contradictoires. Au laboratoire et dans la nature, le lot 2 était préféré; de même le lot 3 était préféré au lot 4. Cependant de nombreuses expériences de choix dans la nature entre les lots 1 et 3 et 1 et 4 ont montré la même tendance qu'au laboratoire, mais pour d'autres, les résultats étaient inversés. Après regroupement des résultats dans la nature, le lot 3 a été préféré au lot 1 (résultats à la limite du seuil de validité); aucune différence significative n'est apparue entre les lots 1 et 4. Les choix deA. floccocus etD. citri entre les lots 1 et 3 n'ont pas été modifiés par la présence de l'un des lots pendant les 3 jours précédant l'expérience.

     

Stylet penetration by larvae of the greenhouse whitefly on cucumber

Probing behaviour of Trialeurodes vaporariorum (Westwood) larvae was monitored using the DC electrical penetration graph (EPG) technique on the host plant cucumber. EPGs were recorded for 16 h, simultaneously with honeydew excretion using a honeydew clock. Three waveforms were distinguished: a pathway waveform (C), and two phloem waveforms, one with a high (H), and one with a low frequency (L) signal. The C waveform mainly occurred in the crawler stage of the 1st instar larvae. EPGs recorded from larvae during and after moulting indicated that the process involves stylet withdrawal; hence the stylets of each new instar need to penetrate again from the leaf surface to the phloem.All sessile stages, from L1 to pre-pupa, spent almost their entire time in waveforms H and L. These waveforms alternated more frequently in the early instars than during the later ones, in which the H waveform became predominant. The H waveform was highly correlated with honeydew excretion and thus phloem sap ingestion. The L waveform was not related to honeydew excretion but EPGs indicated that the stylet tips remain in a sieve element during both waveforms. Periods of honeydew production demonstrated a delay of 30–40 min in relation to the onset and end of H and L waveforms. This delay is presumably related to the time needed for food passing through, or emptying of, the insect's gut. From the 1st instar to the pre-pupa, the frequency of excreted honeydew droplets decreased but their size increased, causing a net increase of the excretion rate.     

Stylet penetration behavior resulting in inoculation of a semipersistently transmitted closterovirus by the whitefly Bemisia argentifolii

The electrical penetration graph (EPG) technique was used to determine what part of stylet penetration behavior by the whitefly vector, Bemisia argentifolii Bellows & Perring (Homoptera: Aleyrodidae), is lettuce chlorosis virus (LCV) inoculated in the host plant Malva parviflora L. LCV is a semipersistently-transmitted closterovirus. Since closteroviruses generally are found in the phloem of their plant hosts, this study tested the hypothesis that virus inoculation occurs during the phloem phase of stylet penetration behavior. Virus-exposed whiteflies were allowed to feed on uninfected host plants, and the whiteflies were divided into two experimental groups: group 1 attained phloem phase on the uninfected plants, and group 2 did not attain phloem phase. Two series of tests were conducted, one where whiteflies were manipulated so that the amount of time spent in non-phloem phase stylet penetration behaviors was similar between group 1 and group 2, and a second series of tests where whiteflies were manipulated so that the number of intracellular punctures made during stylet penetration was similar between group 1 and group 2. Both series of tests indicated that virus inoculation took place primarily during phloem phase. Considering only individual whiteflies shown to be capable of transmitting virus, 11 of 23 whiteflies (48%) in the phloem phase treatment successfully inoculated the virus whereas only one of 19 whiteflies (5%) in the non-phloem phase treatment successfully inoculated the virus (P=0.00008).     

Identification of phloem sieve elements as the site of resistance to silverleaf whitefly in resistant alfalfa genotypes

Experiments were conducted to locate the plant tissue where resistance is expressed against silverleaf whitefly, Bemisia argentifolii Bellows and Perring (Homoptera: Aleyrodidae), in alfalfa, Medicago sativa L. (Fabaceae), genotypes previously shown to have high levels of resistance against this pest. Previous work demonstrated that resistance in the resistant alfalfa genotypes was expressed primarily as high first‐instar mortality; consequently this study focused on first‐instar nymphs. Examination of stylets in cleared leaf tissue indicated that first‐instar nymphs located vascular bundles with equal success on resistant and susceptible alfalfa genotypes. Furthermore, direct current electrical penetration graphs (DC‐EPG) indicated that sieve elements were penetrated and phloem ingestion behavior was initiated with equal success on resistant and susceptible genotypes. Thus, the mechanism of resistance does not reside in tissues encountered by the stylets prior to penetrating a phloem sieve element. Honeydew production (as a proxy for ingestion) was greatly reduced on two resistant genotypes compared to the two susceptible genotypes. The frequency distribution of honeydew production was bimodal, indicating that most individuals on the resistant genotypes produced little or no honeydew while some produced as much honeydew as whiteflies on the susceptible genotypes. This indicates that expression of resistance is an all‐or‐nothing phenomenon; an individual nymph either encounters resistance and cannot sustain ingestion or it does not encounter resistance and ingests just as well as on a susceptible plant. Intermediates are rare. DC‐EPGs indicate that phloem ingestion behavior is significantly reduced on two of the resistant genotypes compared to the susceptible genotypes. The primary reason for this appears to be more frequent termination of phloem ingestion behavior on at least one of the resistant genotypes. On one of the resistant genotypes, the productivity of EPG‐measured phloem ingestion behavior (honeydew produced per min of phloem ingestion behavior) was reduced compared to a susceptible control.     

Design and use of a simulation model to evaluate germplasm for antibiotic resistance to the greenhouse whitefly (Trialeurodes vaporariorum) and the sweetpotato whitefly (Bemisia tabaci)

SARAH (Software for theAssessment of antibioticResistance toAleyrodidae inHost plants) is a deterministic simulation model of whitefly population growth based on whitefly life-history components determined on individual plants. The life-history components recorded were oviposition rate, adult survival, pre-adult survival, developmental period, and sex ratio. The simulation model serves as a tool to combine these components and to obtain a single criterion for (antibiotic) resistance. The criterion used was the decrease in simulated intrinsic population growth rate, r _s , relative the r _s value determined on a susceptible control genotype. This model-based evaluation method was tested using the greenhouse whitefly,Trialeurodes vaporariorum Westwood, on tomato and the sweetpotato whitefly,Bemisia tabaci Gennadius, on tomato, eggplant, collard, and pepper. To study its consistency over time, the evaluation method was repeated six times forT. vaporariorum on a susceptible and a resistant tomato cultivar. Simulated intrinsic population growth rate was more consistent in indicating resistance than any of the individual life-history components. Of tenL. hirsutum accessions tested for resistance toT. vaporariorum, three exhibited r _s values that were significantly lower than those for the susceptible control. In addition, on these tenL. hirsutum accessions, a significant positive correlation was observed between r _s and sex ratio (# females/# males). Four host plant species (tomato, collard, eggplant, and pepper) were evaluated for resistance toB. tabaci. All life-history components and r _s values varied among host species, while a negative r _s value was observed forB. tabaci on pepper. A high correlation was found between results from a sensitivity analysis of SARAH and results from a sensitivity analysis of a validated whitefly population simulation model by Yanoet al. (1989a). Significant correlations were found for the relationships between oviposition rate, adult survival, or pre-adult survival and r _s , indicating that none of these life-history components can be omitted from the test procedure. This model-based evaluation method offers a standardized way to quantify levels of antibiotic resistance to whiteflies and will enhance efficiency in breeding programs.     

Pathway phase waveform characteristics correlated with length and rate of stylet advancement and partial stylet withdrawal in AC electrical penetration graphs of adult whiteflies

A technique was developed for measuring the length of stylet insertion during adult whitefly probing. The distance that the labium shortens during a probe was shown to be equal to the length of stylets that were inserted into the plant tissue. The length of labial shortening then was measured in high-magnification video recordings of adult female silverleaf whitefly, Bemisia argentifolii, in conjunction with recording electrical penetration graphs (EPGs – AC method). Using a split-screen device, video images of the whitefly's labium during a probe and the EPG waveforms produced during the probe were recorded simultaneously on the same video tape. On playback, changes in labial length could be measured during specific EPG waveforms to determine the length of stylet insertion that occurred during the waveforms. The focus of the study was on two characteristics of the pathway phase sawtooth waveform: the frequency of voltage peaks and the increase in voltage level that occurs over time during sawtooth waveforms. The rate of stylet penetration was significantly and positively correlated with frequency of sawtooth waveform voltage peaks (r ²=0.33) and the length of stylet penetration was significantly and positively correlated (second-order polynomial) with the relative difference in voltage level between the beginning and end of the sawtooth waveform (r ²=0.43). Stylet advancement did not appear to occur during the few low-flat waveforms (unknown behavioral correlation) and high-flat waveforms (phloem phase) that were observed. Voltage drops occur sporadically during sawtooth waveforms, and these were associated with partial stylet withdrawal (indicated when the labium increased in length, but the probe was not terminated) with an accuracy of 99%.     

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).     

Survival of the parasitoidEncarsia formosa after treatment of parasitized greenhouse whitefly larvae with fungal spores ofAschersonia aleyrodis

The interaction between the entomopathogenic fungusAschersonia aleyrodis and the parasitoidEncarsia formosa on greenhouse whitefly as a host organism was studied, in particular, the survival of the parasitoid after treatment of parasitized hosts with fungal spores. The mean number of parasitized black pupae per parasitoid produced at 25°C was significantly reduced after spore treatment in the first three days following parasitization. Spore treatment four, seven or ten days after parasitization resulted in a mean number of parasitized pupae not significantly different from the number of black pupae in the control. The rather sudden change from low to high survival of parasitized hosts when treated with spores four days after parasitization in spite of high numbers of infected unparasitized larvae, coincided with the hatching of the parasitoid larva from the egg inside the host. Possible reasons for this decrease in susceptibility to infection after parasitoid egg hatch, such as induced changes in host cuticle or haemolymph, are discussed. Parasitoids emerged from treated hosts did not show differences in reproduction compared with parasitoids emerging from untreated hosts. Both natural enemeies of whitefly are compatible to a great extent.     

Encarsia formosa parasitizing the Poinsettia-strain of the cotton whitefly,Bemisia tabaci, on Poinsettia: bionomics in relation to temperature

Adult longevity, developmental time and juvenile mortality ofEncarsia formosa Gahan (Hymenoptera:Aphelinidae) parasitizing the Poinsettia-strain ofBemisia tabaci (Gennadius) (Homoptera: Aleyrodidae) on Poinsettia (Euphorbia pulcherrima Willd.) were investigated in laboratory experiments at three temperatures: 16 °C, 22 °C and 28 °C. Furthermore, the parasitoid's preference for different larval stages of the whitefly was determined at 24.5 °C. The lifespan ofE. formosa decreased with temperature from one month at 16 °C to nine days at 28 °C. A lower temperature threshold of 11 °C for adult development was found. The development of juvenile parasitoids inB. tabaci lasted more than two months at the lowest temperature, but was only 14 days when temperature was 28 °C. The lower temperature threshold for immature development was 13.3 °C, yielding an average of 207 day-degrees for the completion of development into adults. Juvenile mortality was high, varying from about 50% at 16 °C to about 30% at 22 °C and 28 °C.E. formosa preferred to oviposit in the 4th instar and prepupal stages ofB. tabaci followed by the 2nd and 3rd instars. The preference for the pupal stage was low. The parasitoid used all instars of the whitefly for hostfeeding, with no apparent differences between the stages. The average duration of the oviposition posture was four minutes. Demographic parameters were calculated from life tables constructed from the data. The intrinsic rate of increase (r _m) and the net reproductive rate (R ₀) increased with temperature from 0.0279 day⁻¹ at 16 °C to 0.2388 day⁻¹ at 28 °C and from about 12 at 16 °C to about 66 at 28 °C, respectively.     

Host age, sex, and pathogen exposure level as factors in the susceptibility of Musca domestica to Entomophthora muscae

Adult Musca domestica were exposed to conidial showers from cadavers that were killed by Entomophthora muscae. Of the 4 host ages (14 days, 7 days, 3 days and <1 day), older flies were more likely to die without postmortem signs of infection, while younger flies were more likely to die with signs and produce primary conidia. The youngest fly group also succumbed to infection significantly sooner than older flies. Total mortality did not differ for the 2 sexes, but females died significantly more often with signs in one trial. Males died significantly earlier than did females, but differences were not significant when the sexes were paired by weight. Increased levels of pathogen exposure significantly increased both total mortality and the number of flies dying with signs; it also decreased incubation period. The potential significance of these findings in the epidemiology of the disease is discussed.

---

Zusammenfassung Die Mortalitätsrate von Musca domestica nach Behandlung mit Entomophthora muscae Sporen wurde in Laborversuchen in Abhängigkeit von Wirtsalter,-geschlecht und Expositionsdauer festgestellt. Fliegen verschiedenen Alters (14 Tage, 7 Tage, 3 Tage und jünger als 1 Tag) wurden gemeinsam in 237 ml Kartonbehältern gehalten. Die Behälter besassen Netzdeckel, auf denen frisch an einer Entomophthora Infektion gestorbene Fliegen ausgelegt wurden. Auf diese Weise wurden die Versuchstiere 8 Stunden (low exposure) bzw. 48 Stunden (high exposure) lang mit Entomophthora Sporen kontaminiert. Mit Hilfe von am Boden der Versuchsgefässe ausgelegten Objektträgern wurde die relative Menge Entomophthora Sporen festgestellt, der die Versuchstiere ausgesetzt waren. Der Totenfall wurde täglich für die Dauer von 10 Tagen registriert; dabei wurde zwischen Konidienträgern (external signs) und Leichen ohne Konidien (no external signs) unterschieden.Die Versuchsgruppen, die 48 Stunden lang mit Entomophthora Sporen behandelt worden waren, wiesen im Vergleich zu den kürzer behandelten Versuchsgruppen höhere Mortalität und einen höheren Anteil an Konidienträgern auf. Darüber hinaus wurde das Auftreten von Konidienträgern bei den länger behandelten Gruppen früher beobachtet. Im Vergleich zu unbehandelten Kontrollgruppen erhöhte die Behandlung mit Entomophthora Sporen auch die Rate nicht konidientragender Leichen.Bei gleicher Expositionsdauer zeigten Weibchen und Männchen ähnliche Mortalitätsraten. Ein höherer Anteil Konidienträger bei den Weibchen wurde bei einem von zwei entsprechenden Versuchen beobachtet. Konidienträger traten bei Weibchen später als bei Männchen auf. In Bezug auf die Inkubationszeit konnte kein geschlechsspezifischer Unterschied festgestellt werden, wenn Versuchsgruppen mit gleichem Gewicht verglichen wurden. Bei beiden Geschlechtern erfolgte die Bildung von Konidien bei grösseren Individuen später als bei kleineren Individuen.Das Alter der Versuchstiere beeinflusste sowohl die Mortalitätsrate als auch die Häufigkeit und das zeitliche Auftreten von Konidienträgern. Jüngere Fliegen wiesen einen höheren Anteil Konidienträger auf; bei Fliegen mit einem Alter von weniger als einem Tag wurde das Auftreten von Konidienträgern früher beobachtet als bei älteren Individuen.Die untersuchten Faktoren können auch für die Epidemiologie von Entomophthora freilebender Musca domestica Populationen eine wichtige Rolle spielen. In Populationen mit niedrigerem Durchschnittsalter wird sich eine Infektion rascher ausbreiten, da jüngere Fliegen eine kürzere Inkubationszeit aufweisen und auch nach kürzerer Kontaminationsdauer Konidien an den getöteten Individuen ausgebildet werden. Darüber hinaus wird sich eine Infektion dann rascher ausbreiten, wenn die Population mit einer grösseren Sporenmenge infiziert wird, da dann eine kürzere Inkubationzeit und ein höherer Anteil Konidienträger zu erwarten ist.

     

Suitability of common weed species as host plants for the potato leafhopper,Empoasca fabae

The suitability of 13 weed species as host plants for the potato leafhopper,Empoasca fabae (Harris) (Homoptera: Cicadellidae) was compared to alfalfa,Medicago sativa L., and broad bean,Vicia faba L., in laboratory and greenhouse tests. Nymphs developed and adults reproduced on five of the weeds: smartweed (Polygonum pensylvanicum L.), pigweed (Amaranthus retroflexus L.), shepherdspurse (Capsella bursa-pastoris (L.) Medic.), dandelion (Taraxacum officinale Weber), and carpetweed (Mollugo verticillata L.). Adults survived and oviposited on all tested weeds, suggesting that nymph survival was the limiting factor in host suitability. Yellow nutsedge (Cyperus esculentus L.), crabgrass (Digitaria sanguinalis (L.) Scop.), and foxtail (Setaria faberi Herrm.) were least suitable for adult and nymph survival. Large nymphs (instar III–IV) were able to complete development on weed species that did not support small nymph (instar I–II) development to adulthood. Densities of nymphs on weeds from alfalfa fields as well as the laboratory studies suggested that the weed species are less suitable as hosts and thus weeds probably do not affect population increases of potato leafhopper within weedy alfalfa fields by serving as hosts. Weeds may however, interact with the crop to alter leafhopper colonization, dispersal, and/or reproduction. In addition, since some broadleaf weeds are suitable hosts, the leafhopper may have a secondary beneficial role for biological weed control. These interactions should be considered when evaluating pest management practices.

---

Eignung häufiger Unkrautarten als Wirte vonEmpoasca fabae

Zusammenfassung Der Wirtskreis derEmpoasca fabae (Harris) (Homoptera: Cicadellidae) wurde im Labor und im Freiland untersucht. Dreizehn Unkrautarten, wie auch Luzerne (Medicago sativa L.) undVicia faba L., wurden im Laborversuchen geprüft. Die Entwicklung der Nymphen und die Eiablage setzten auf fünf Unkrautarten fort: aufPolygonum pensylvanicum L.,Amaranthus retroflexus L.,Capsella bursa-pastoris (L.) Medic.,Taraxacum officinale Weber undMollugo verticillata L. Die Eiablage und das Weiterleben der Imagines geht auf allen untersuchten Unkräutern weiter. Dies weist darauf hin, dass die Lebensdauer der Nymphen die Eignung der Wirte begrenzt. Das Weiterleben der Imagines und der Nymphen waren aufCyperus esculentus L.,Digitaria sanguinalis (L.) Scop. undSetaria faberi Herrm. am stärksten verkürzt. Die Altnymphentwicklung (3. und 4. Stadien) bis zur Imago war auf Unkräutern gewährleistet, dagegen erfolgte die Jungnymphentwicklung (1. und 2. Stadien) bis zur Imago nicht. Populationsproben von Luzerne im Freiland, wie auch die Ergebnisse der Laboruntersuchungen, zeigten, dass Unkräuter nicht im Wirtskreis der Nymphen vonE. fabae sind und dass Unkräuter in Luzernefeldern keinen Einfluss auf die Populationszunahme in solcher Feldern haben. Unkräuter in Luzerne dürften aber die Kolonisierung, Verteilung der Population im Feld wie auch das Vermehrungspotential vonE. fabae verändern. Weil einige Unkräuter Wirte der Imago vonE. fabae sind, können sie als Ausweichpflanzen dienen, wenn die Luzerne, z.BB. nach dem Schnitt, für das Ueberleben der Zikaden ungeeignet ist.

     

Biological activity of maize leaf extracts against the African armyworm,Spodoptera exempta

Maize (Zea mays L.) leaf tissue of cv Bastille and cv Michoacan 12 was extracted with n-hexane. The extracts were bioassayed against 5th instar African armyworm,Spodoptera exempta (Walker)(Lepidoptera: Noctuidae), by feeding the larvae on agar based media or sucrose impregnated glass fibre discs. The hexane extract of the ‘resistant’ cv Bastille exhibited feeding deterrency and toxicity which were not shown by the ‘susceptible’ cv Michoacan 12. The hexane extract of cv Bastille was adsorbed onto silica gel, the solution filtered off and the adsorbed component taken up into ethyl acetate. Bioassay of these fractions indicated that the toxic and deterrent action was retained in the ethyl acetate fraction. Preparative thin layer chromatography of the ethyl acetate fraction isolated two biologically active constituents. These were both growth inhibitors and lethal by ingestion to the 5th instar African armyworm. Implications for resistance in maize varieties to insect pests are discussed.     

Feeding behavior: selection of both wet and dry food for increased growth in Schistocerca gregaria nymphs

An apparent fluctuating preference in acridids for food of different water contents is confirmed for Schistocerca gregaria nymphs. Short- and long-term preference tests reveal that previous diet, through its effect on locust water balance and overall food intake, can influence feeding behavior on high-water content (wet) and low-water content (dry) food. Locusts given only wet or dry food for 3 days prior to testing initially chose the food not received in pretreatment when given a choice of the two. They had decreased latencies to feed, increased meal length and decreased switching between food types as compared with locusts given the option to regulate water intake before testing. It is suggested that the behavior of the single-diet locusts has several related causes, among them, deterrence of the food, disturbed water balance in the locust, and chronic hunger. Long-term preference tests reveal that fifth-instar locusts given a continuous choice between wet and dry food take significantly more of their total dry matter from dry food each day until the last feeding day of the instar (day 9), when they take 97% of their total dry matter from wet food. This behavioral regulation of water and dry matter intake is adaptive: locusts with access to a mixed diet of wet and dry food have increased growth as compared with ones given only wet food. The growth increase is due largely to increased dry matter consumption, with some contribution from increased digestive efficiency and reduction in respiratory rate.

---

Résumé L'étude de larves de dernier stade de Schistocerca gregaria confirme les observations sur les acridiens d'une préférence apparemment labile pour les aliments à différentes teneurs en eau. Des observations sur les choix à court et à long termes rélèvent que le régime alimentaire antérieur, par son action sur le régime hydrique du criquet et sur sa prise totale de nourriture, peut influer sur le comportement alimentaire face à des aliments à forte teneur en eau (humides ou frais) ou à faible teneur (secs). Des criquets, approvisionnés uniquement en aliments secs ou frais pendant les 3 jours précédant l'expérience, choisissent de préférence la catégorie d'aliments non fournie pendant ce prétraitement. Par comparaison avec des criquets ayant eu la possibilité d'ajuster leur apport en eau avant l'expérience, ils présentaient une diminution du temps de latence avant de s'alimenter, une prolongation de la durée des repas et une diminution du nombre de changements de type d'aliments. II est suggéré que le comportement des criquets, à régime alimentaire composé d'un seul type, a plusieurs causes associées, en particulier la répulsion pour le végétal, la perturbation de l'état hydrique du criquet, et la faim chronique. Des tests sur les préférences à long terme montrent que des larves de 5ème stade ayant eu un choix continu entre aliments frais et secs prirent significativement plus de leur matière totale d'aliments secs jusqu'à ce qu'elles parviennent au dernier jour d'alimentation du stade (jour 9), alors elle prélevèrent 97% de leur matière sèche totale d'aliments frais. Cette régulation comportementale de l'absorption d'eau et de matière sèche est adaptative: des criquets ayant accès à une alimentation mixte (nourriture sèche et fraîche) ont unt croissance supérieure à ceux nourris exclusivement d'aliments frais. La croissance supérieure est due pour beaucoup à une consommation accrue de matière sèche, avec une légère contribution de l'amélioration de l'efficacité digestive et de la réduction du taux de respiration.

     

Adaptation of a generalist moth, Operophtera brumata, to variable budburst phenology of host plants

The adaptation of three allopatric populations of a generalist moth, Operophtera brumata (L.), on two tree species, Prunus padus (L.) and Quercus robur (L.) which represent the extremes of the timing of budburst in spring, was studied in Finland and Sweden. The synchrony of the hatching and budbursting was monitored, and its importance to dispersal and growth of larvae was assessed by rearing cohorts of larvae, whose hatching dates were manipulated, on both hosts. In addition, the realised heritability of the hatching time was estimated.Experimental populations hatched in approximate synchrony with the budburst of their original host species. As a result of the manipulation of the hatching dates of larvae, the growth rates of larvae decreased and the dispersal rates increased on both hosts in relation to the ageing of foliage. The realised heritability of hatching times was rather high (0.63). There were fewer differences in the host use efficiency and behaviour of the experimental populations than in the hatching phenology. The synchrony of hatching with the budburst of the local dominant host plant is probably a result of stabilising selection.     

Genetic variability in host plant adaptation of the green peach aphid, Myzus persicae

The green peach aphid, Myzus persicae (Sulz.), is polyphagous on over 400 plant species in more than 50 families. Phenotypic plasticity of individuals and genetic variability in the population presumably contribute to this polyphagy. The genetic variability in field populations of M. persicae was assessed with respect to their adaptation to sugar beets and potatoes. An analysis of more than 1 000 clones, sampled during 1980, 1981 and 1982 from different host plants in the field, revealed a wide genetic variability in host plant adaptation to sugar beets as well as to potatoes. Both traits seem to be inherited independently from each other and do not correlate with clone-specific host plant preference of apterous adults. The aphid M. persicae can be characterized as a polyphagous insect species with a wide, continuously distributed variability and a broad phenotypic plasticity. A general differentiation of herbivorous species into generalists and specialists tends to ignore the genetic component in the complex of insect-plant relationships.

---

Zusammenfassung Die Grüne Pfirsichblattlaus Myzus persicae (Sulz.) lebt polyphag an über 400 Pflanzenarten in mehr als 50 Pflanzenfamilien. Ein breites Nahrungsspektrum einer Art ergibt sich jedoch aus der phänotypischen Plastizität des Individuums oder einer Klonlinie und der genetischen Variabilität der Population. Felpopulationen der Grünen Pfirsichblattlaus wurden auf ihre genetische Variabilität bezüglich der Wirtspflanzenanpassung an Rübe und Kartoffel untersucht. Eine Analyse von mehr als 1 000 Klonen, die über die Jahre 1980, 1981 und 1982 im Rheinland gesammelt wurden, lassen eine breite Variabilität in der Wirtspflanzenanpassung der Population erkennen. Beide Merkmale scheinen unabhängig voneinander vererbt zu werden und zeigen keine Beziehung zum Wirtswahlverhalten adult apterer Läuse der entsprechenden Klone. Die Art M. persicae kann daher charakterisiert werden als eine polyphage Insektenart mit einer breiten genetischen Variabilität und einer grossen phänotypischen Plastizität. Eine generelle Differenzierung von Herbivoren in Generalisten und Spezialisten vernachlässigt die genetische Komponente in der komplexen Beziehung zwischen Insekten un ihren Wirtspflanzen.

     

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.     

The mechanism of trichome resistance in Anaphalis margaritacea to the meadow spittlebug Philaenus spumarius

This study demonstrated how simple, lanate trichomes on the stem of Anaphalis margaritacea (D.C.) inhibit feeding by young nymphs of Philaenus spumarius (L.). Nymphs were caged on stems which were subsequently sectioned and examined for feeding marks. The lack of stylet punctures at sites where nymphs were unable to feed indicated that trichomes prevented the start of stylet penetration rather than interfering with the progress of the stylets after penetration had begun. Trichome resistance declined progressively with spittlebug instar from complete resistance to first instar nymphs through complete absence of resistance to fifth instar nymphs. Discriminant analysis of the effect of number of trichomes per mm², vertical density, and height on trichome resistance to third instar nymphs revealed that trichome height was the critical factor in resistance. Variation in the percent of nymphs in each instar able to feed when caged on the stem was (1) positively correlated with degree to which the average rostrum length of the instar exceeded the trichome height, (2) independent of trichome density, which was uniformly high on all stems. Trichomes prevented stylet penetration where trichome height exceeded the length of the nymph's rostrum, probably because the necessary stimuli were not received when the sensilla on the tip of the rostrum could not touch the surface of the stem. A similar mechanism may explain resistance of simple trichomes to crop pests in the Cicadellidae.

---

Résumé Ce travail montre comment des trichomes simples et laineux sur les tiges d'Anaphalis margaritacea (D.C.) inhibent la prise d'aliments par les jeunes larves de Philaenus spumarius L. Les larves ont été encagées sur des tiges qui ont été ultérieurement coupées et les piqûres alimentaires dénombrées. L'absence de piqûres par les stylets aux lieux où les larves ont été incapables de s'alimenter a montré que les trichomes ont empêché le début de la pénétration des stylets plus qu'ils n'ont entravé leur progression après le début de la pénétration. La résistance des trichomes diminue progressivement: totale pour le premier stade larvaire, elle est devenue nulle au cinquième stade. Une analyse discriminante des effets de la densité des trichome par mm², de la densité verticale, de leur hauteur sur la résistance des trichomes aux larves de 3ème stade, a révélé que la hauteur du trichome est le facteur de résistance. La variation dans le pourcentage de larves de chaque stade encagées sur la tige, capables de consommer, présentait une relation positive avec le degré par lequel la longueur du rostre dépasse la hauteur du trichome, quelle que soit la densité des trichomes qui est toujours élevée sur toutes les tiges. Les trichomes ont empêché la pénétration des stylets quand leur hauteur a dépassé la longueur du rostre, probablement parce que les stimuli nécessaires n'ont pas été reçus quant l'extrémité du rostre ne pouvait pas atteindre la surface de la tige. Un mécanisme semblable peut expliquer la résistance de trichomes simples aux Cicadellidae.

     

Characterization and distribution of esterase electromorphs in the whitefly,Bemisia tabaci (Genn.) (Homoptera: Aleyrodidae)

Esterase profiles were examined for over 40 populations of the whitefly,Bemisia tabaci, obtained from native and cultivated plant hosts worldwide. Twelve unique electromorphs were identified from distinct populations concentrated largely in Central America, Africa, and India. One electromorph, type B, has recently been proposed as a separate species,Bemisia argentifolii, and has recently spread throughout much of the world. When considered with evidence from mating studies and the ability to induce phytotoxic disorders (squash silverleaf disorder), our data suggest that the single taxonBemisia tabaci may actually represent a species complex.     

Post-release survey to assess impact and potential host range expansion by Amitus hesperidum and Encarsia perplexa, two parasitoids introduced for the biological control of the citrus blackfly, Aleurocanthus woglumi in Dominica

Four years after the release of two exotic parasitoids, Amitus hesperidum Silvestri (Hymenoptera: Platygasteridae) and Encarsia perplexa Huang and Polaszek (Hymenoptera: Aphelinidae) for the classical biological control of the citrus blackfly (CBF), Aleurocanthus woglumi Ashby (Hemiptera: Aleyrodidae) in Dominica, a survey was conducted to assess establishment as well as potential nontarget effects especially on Aleyrodidae and other related taxa. CBF populations were low to non-existent in 50 of 51 field sites examined. At the site where CBF was encountered, both E. perplexa and A. hesperidum were present and CBF populations were declining. The two parasitoids were not among the several species collected on nontarget Aleryodidae and Hemiptera. It is concluded that E. perplexa and A. hesperidum have kept CBF populations under effective biological control in Dominica and there is no evidence of any nontarget effects on other Aleyrodidae or their natural enemies. Handling Editor: Dirk Babendreier.