The effects of parasitism by an encyrtid parasitoid, Anagyrus pseudococci on the survival, reproduction and physiological changes of the mealybug, Planococcus citri

Survival, reproduction and some biochemical changes in the citrus mealybug, Planococcus citri (Risso) parasitized by the endoparasitoid, Anagyrus pseudococci (Girault) are reported. The parasitized younger mealybugs were more likely to die than be successfully parasitized. The mean duration (±SE) required for mortality other than mummification were 1.85 ± 0.06, 2.48 ± 0.14 and 2.89 ± 0.17 days for first, second and third instar P. citri respectively. Hosts parasitized in the first instar did not produce any mummies while the second and third instar nymphs and adult stages of the mealybugs were mummified on average 8 to 11 days after attack. Parasitization caused cessation of normal fecundity as well as induction of early egg maturation of the mealybugs. The mealybugs parasitized in the adult stage produced a few eggs but the fecundity of unparasitized mealybugs was about 40 fold higher than that of parasitized mature adults. Parasitized pre-ovipositing and mature adults laid eggs within 24 h of attack. SDS-PAGE analysis of the body content of the parasitized mealybugs after 24 h and 48 h of parasitization and before egg hatch revealed an increase in the concentration of some proteins. Response to wound reactions was not visible either as formation of extra protein bands or increased density of the existing bands. The possible effect of parasitism on the changes of the reproductive physiology of the host is discussed.     

Functional response of a parasitoid Ganaspidium utilis （Hymenoptera： Eucoilidae） on the leafminer Liriomyza trifolii （Diptera： Agromyzidae）

Functional response of a solitary, larval-pupal endoparasitoid of Liriomyza leafminers, Ganaspidium utilis Beardsley, was estimated on Liriomyza trifolii Burgess at three temperatures （17℃, 25℃, 29℃ ） and host densities. A type Ⅱ random parasitoid equation （RPE） was used to estimate instantaneous search rate and handling time. The instantaneous search rate increased as temperature increased. All of the RPE regressions obtained for functional response of G. utilis at different temperatures were significant （P〈0.01）. The slope of RPE regression lines was lower across the temperatures. At 29±2℃, the maximum number of larvae parasitized was 7.8 per day. It decreased to 7.2 larvae parasitized at 25±2℃. At 17±2℃, no significant increment of parasitization was observed due to the host density increments. The estimated handling time was lowest at 17±2℃ and highest at 25 ± 2℃, respectively. The ability of G. utilis to find and parasitize L. trifolii over a wide range of temperatures makes them a good candidate for biological control of Liriomyza leafminers.     

Host feeding and ovipositor insertion as sources of mortality in the mealybug Phenacoccus herreni caused by two encyrtids, Epidinocarsis diversicornis and Acerophagus coccois

Females of Epidinocarsis diversicornis (Howard) killed cassava mealybugs, Phenacoccus herreni Cox & Williams, through host feeding and piercing (i.e. ovipositor insertion and its consequences). Parasitoids fed on 9% of 592 hosts in which ovipositor insertion was achieved. For hosts which were attacked but not fed on, mortality in the first 72 h was significantly greater than that of controls and was highest in the first nymphal instar. No host feeding by Acerophagus coccois Smith was observed in 494 attacks. Mortality due to piercing was significantly greater than control values, but did not differ between host life stages.

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Résumé La prise de nourriture et les piqûres (c'est-à-dire l'insertion de la tarière et ses conséquences) de Epidinocarsis diversicornis tuent Phenacoccus herreni. Les adultes s'alimentent sur 9% des 592 cochenilles dans lesquelles la tarière a pénétré. La mortalité au cours des 72 premières heures est beaucoup plus élevée chez les hôtes attaqués mais qui n'avaient pas subi une prise de nourriture, que chez les témoins; elle était particulièrement élevée chez les cochenilles du premier stade larvaire. Pour 494 attaques par Acerophagus coccois, aucune prise de nourriture n'a été observée; la mortalité était plus élevée que chez les temoins, sans qu'il y ait eu de différences suivant les stades.

     

Heritable differences in the response of the braconid waspMicroplitis croceipes to volatile allelochemicals

The heritable nature of differential responses by Microplitis croceipes(Cresson) (Hymenoptera: Braconidae) to airborne allelochemicals was investigated. Four isofemale lines were tested for three generations in a flight tunnel. Flight response was found to be strongly dependent on the familial origin, with two highly responsive lines being clearly differentiated from two less responsive ones. Each isofemale line could also be recognized by typical behaviors that suggested that M. croceipesfemales may inherit independent characters of sensory acuity to plant and to host chemicals and of learning ability.     

Predicting development of the mealybug parasitoids Anagyrus pseudococci, Leptomastix dactylopii and Leptomastidae abnormis under glasshouse conditions

Developmental rates of three encyrtid mealybug parasitoids were examined under constant, cycling and glasshouse temperatures. Under constant conditions the rate of development increased with rising temperature up to 35°C, but a constant 40°C was lethal. Under cycling conditions, 12 h periods at 40°C could be tolerated. Developmental rates under cycling conditions could be calculated from constant temperature data, by incrementing development on an h/h basis. A new technique was developed to determine the lower thresholds for development, using cycling regimes and to calculate the rates of development over short periods at high temperature. A previously untested polynomial regression technique was used to predict mean developmental periods under glasshouse conditions. These predictions were considerably more accurate than day-degree (D^o) predictions when substantial periods at extreme temperatures were involved, but not significantly different from day-degree predictions under hot-house conditions. In addition a method was developed to calculate a 95% range for emergence, which gave consistently good estimates of first and last eclosion observed under fluctuating conditions.

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Résumé Les vitesses de développement de 3 parasitoïdes de cochenilles ont été étudiées dans les conditions de température de serre, avec thermopériode ou en température constante. En conditions constantes, la vitesse augmente jusqu'à 35°C, mais est létale à 40°C. Avec thermopériode, une thermophase de 40°C peut être tolérée pendant 12 heures. Les vitesses de développement sous thermopériodes pourraient être calculées à partir des données obtenues en conditions constantes, modifiées en fonction des durées respectives des cryophases et thermophases. Une méthode originale est présentée pour obtenir les seuils inférieurs de développement, en utilisant des thermopériodes, et pour calculer les vitesses de développement à températures élevées.Une méthode de régression polynomiale, non vérifiée préalablement, a été utilisée pour prévoir les durées moyennes des développements sous serre. Ces estimations sont beaucoup plus sûres que les prédictions basées sur le concept de degré-jour (D^o), lorsque les températures extrêmes durent longtemps, mais en serre chaude elles ne présentent aucune différence significative. De plus, une méthode d'évaluation des 95% d'émergences, a donné de bonnes estimations des premières et dermières émergences en conditions périodiques.

     

Functional response of the predator <Emphasis Type="Italic">Scolothrips takahashii</Emphasis> to hawthorn spider mite, <Emphasis Type="Italic">Tetranychus viennensis</Emphasis>: effect of age and temperature

A leaf disc bioassay was employed to examine the effects of temperature and predator age on functional response of an acarophagous thrips, Scolothrips takahashii Priesner, to hawthorn spider mite, Tetranychus viennensis Zacher, in the laboratory. The results indicated that the predatory thrips exhibited type-II functional responses against the mites under various temperatures, and that females are more voracious than males. Analysis showed that temperature had significant effects on the predatory capacity of adult thrips over the range of 20–35 °C. Attack rate in females linearly increased with temperature while in males it was independent of temperature. Handling times in both males and females decreased linearly with increasing temperature. Extended response models describing the functional response with temperature incorporated as a parameter were developed, yielding an estimated maximum numbers of prey attacked at four temperatures were 38.38, 55.06, 71.74 and 88.42 eggs per day for females, and 15.11, 26.11, 37.11 and 48.01 eggs per day for males, respectively. The age of predator affected both the type of the functional response shown and the magnitude of predation by S.␣takahashii on the spider mite. Females of various ages exhibited Type-II functional responses with similar attack rates, but handling time prolonged linearly as age increased: the handling times in 15- and 18-d-old females were significantly longer than in 6-d-old thrips. However, Type-I functional responses were determined for males aged 12 d or more; the maximum number of prey eaten in 24 h decreased as age increased. The implications of the results for the management of hawthorn spider mite are discussed.     

Foraging behavior of Campoletis sonorensis in response to Heliothis virescens and cotton plants

Wind tunnel flight behavior of inexperienced female Campoletis sonorensis (Cameron) (Hymenoptera; Ichneumonidae) in response to its larval host Heliothis virescens (F.) (Lepidoptera: Noctuidae) feeding on the host plant cotton (Gossypium hirsutum L.) is described. The flight behavioral sequence was determined by quantification of frequencies of observed behaviors and probabilities of first-order behavioral transitions. Comparison of inexperienced C. sonorensis flights to undamaged and damaged cotton indicated that stimuli from undamaged plants alone are adequate to elicit the complete flight behavioral sequence observed in response to H. virescens feeding on cotton.Parasitoid foraging behavior was also analyzed after landing on the stimulus. This behavior appeared to be random in its initial stages, but became sequential after location of evidence of a host. Analysis of foraging on undamaged and 3 treatments of damaged cotton resulted in the determination that parasitoids tend to remain on damaged plants longer than undamaged plants although no significant difference was detected. C. sonorensis spent a greater percentage of their time foraging on host damaged plants than on undamaged plants.

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Résumé Les comportements d'évaluation de l'effectif d'hôtes potentiels et de la position des hôtes par C. sonorensis (Hyméno.: Iccheumonidae) ont été quantifiés pour déterminer les séquences significatives des événements comportementaux. La localisation de la colonie potentielle d'hôtes est apparue comme une séquence régulière des événements comportementaux. Une fois que le parasitoïde a localisé une colonie potentielle, on a constaté que la recherche au hasard des hôtes se poursuit jusqu'à ce qu'il ait la démonstration qu'il s'agit d'une plante attaquée. La localisation par le parasitoïde d'un hôte certain a consitué une autre séquence régulière des événements comportementaux terminant la localisation de l'hôte.L'influence de pieds de coton intacts, de pieds abîmés mécaniquement et de pieds occupés par des chenilles du 3ème stade de l'hôte et de plantes dont les chenilles ont été retirées juste avant l'expérience a été déterminée en modifiant la composition du complexe hôte/plante. Des femelles naïves de C. sonorensis ont montré en présence de pieds de coton intacts apparemment toutes les séquences comportementales de vol impliquées dans la localisation d'une colonie potentielle d'hôtes. Une fois que le parasitoïde a atteint la colonie potentielle d'hôtes, la présence de dégâts de l'hôte n'a pas modifié le temps passé sur la plante, mais a modifié le temps consacré à la prospection.

     

Host preferences of two encyrtid parasitoids for the Columbian Phenacoccus spp. of cassava mealybugs

In a choice test among six life stages of Phenacoccus herreni Cox & Williams, Epidinocarsis diversicornis (Howard) used its antennae to examine adult and 3rd stadium females more than other stages and preferentially attempted to oviposit in these plus 2nd stadium females. Success of ovipositor insertion was unaffected by host stage. The outcome of these behaviors was preferential oviposition by E. diversicornis in the large female host stages. Acerophagus coccois Smith also preferentially examined larger female mealybugs (second and third stadium nymphs and adults) more than other stages and successfully inserted its ovipositor in these stages more often than in second stadium male nymphs and male cocoons, resulting in a similar preference in this species for larger female host stages. When given a choice between adult female hosts of two species, P. herreni and Phenacoccus gossypii Townsend & Cockerell, E. diversicornis exhibited a clear preference for P. herreni; whereas A. coccois preferred P. gossypii.

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Résumé Epidinocarsis diversicornis (Howard), ayant la possibilité de choisir entre six stades différents de Phenacoccus herreni Cox & Williams, examine avec ses antennes plus particulièrement les adultes et les larves femelles du 3ème stade, et essaie de pondre de préférence dans ces stades et les larves femelles de second stade. L'insertion de la tarière s'effectue aussi bien quel que soit le stade de l'hôte. Il résulte de ces différents aspects du comportement que E. diversicornis pond de préférence dans les femelles des stades les plus avancés. Acerophagus coccois Smith préfère aussi examiner les cochenilles femelles les plus grosses (second et 3ème stade larvaires et adulte), et introduit sa tarière avec succès dans ces stades plus souvent que dans les larves mâles de second stade ou les cocons mâles; il en résulte aussi pour cette espèce une préférence pour les femelles des stades les plus gros.Quand on leur a donné le choix entre des femelles des deux espèces de cochenilles (P. herreni et Phenacoccus gossypii Towsend & Cockerell), E. diversicornis manifestait une nette préférence pour P. herreni, tandis que A. coccois préférait P. gossypii.

     

Foraging by the aphid parasitoid Ephedrus cerasicola for patchily distributed hosts

Females of the aphid parasitoid Ephedrus cerasicola were released into small glasshouses containing 20–25 paprika plants (Capsicum annuum L.), either aphid-free (controls) or infested with different numbers of Myzus persicae (Sulzer), i.e. from 0 to about 1000 aphids per plant. The number of parasitoids per plant were counted 1 h, 2 h, 3 h, 4 h, 5 h, 6 h, 8 h and about 24 h after the release. During the last inspection (24 h) the parasitoids were removed. In the aphid-infested houses, the parasitoids soon gathered on the most heavily infested plants (500–1000 aphids per plant), while the aphid-free and low-infested plants (50–200 aphids per plant) were almost free from parasitoids. In the aphid-free houses, a significantly lower portion of the released parasitoids were found on the plants, and they did not show any preference for certain plants. The percentage parasitism, based on mummies on the plants, was rather density independent after a slight peak on plants with about 50 aphids. No superparasitism was found by dissection of aphids.

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Résumé Des femelles de l'aphidophage Ephedrus cerasicola ont été lachées dans de petites serres contenant 20 à 25 pieds de paprika (Capsicum annuum), contaminés par des effectifs de 0 (témoins) à 1000 pucerons (Myzus persicae) par pied. Le nombre de parasitoïdes par plante a été rélevé 1 h, 2 h, 3 h, 4 h, 5 h, 6 h, 8 h, et environ 24 h après leur libération. Les E. cerasicola adultes ont été retirés lors de la dernière inspection (24 h). Dans les serres infestées, les femelles se rassemblent sur les plantes les plus contaminées (500 à 1000 pucerons par pied), tandis que les plantes sans pucerons ou faiblement contaminées (50 à 200 par plante) n'en avaient presque pas. Dans les serres sans pucerons, des femelles en proportion significativement plus faible ont été trouvées sur les plantes, sans qu'elles aient manifesté une préférence pour certaines d'entre elles. Le taux de parasitisme, d'après le nombre de momies par plante, est apparu plutôt indépendant de la densité, après un faible pic pour les plantes avec environ 50 pucerons. Les dissections de pucerons n'ont révélé aucun superparasitisme.

     

Seasonal variations in the encapsulation of eggs of the encyrtid parasitoid Metaphycus stanleyi by the pyriform scale, Protopulvinaria pyriformis

Encapsulation of eggs of the introduced parasitoid Metaphycus stanleyi Compere (Hymenoptera: Encyrtidae) by the pyriform scale, Protopulvinaria pyriformis (Cockerell) (Homoptera: Coccidae) under both greenhouse and field conditions, at Bet Dagan, was found to occur almost all year round (1986–1988). However, encapsulation rates varied considerably during the different seasons and were correlated with the ambient temperatures. The rates of efficient encapsulation (i) in scales infesting Hedera helix and Schefflera arboricola under greenhouse conditions, were lowest during December to May (6–17%) and highest during July to September (78–100%); (ii) in scales infesting avocado in the orchard, were lowest during October to May (0–11%) and highest during June to August (54–57%). Under greenhouse conditions, encapsulation rates did not differ in scales grown on H. helix and on S. arboricola, but were significantly lower in scales grown on avocado. Encapsulation by scales infesting S. arboricola was more frequent at temperatures ranging from 20–28 °C, than from 7–23 °C.The increased resistance by encapsulation of P. pyriformis to successful parasitization by M. stanleyi during the summer, may account for the inability of the parasitoid to prevent the autumn and winter outbreaks of the pest.     

Sex Allocation by a Parasitoid Wasp (Hymenoptera: Ichneumonidae) to Different Host Species: A Question for the Mechanism of Host Size Estimation

Sex allocation by the polyphagous solitary pupal parasitoid wasp Pimpla luctuosa Smith to a small host species, Galleria mellonella (L.), and a large host species, Mamestra brassicae L., was investigated to test whether female wasps responded to hosts of different sizes across different host species. In the experiments, both host species were presented to each test female wasp. Primary and secondary sex ratio experiments revealed that female wasps laid more female eggs in larger pupae of each host species, indicating that female wasps recognized size differences within host species. The wasp sex ratio (male ratio) from M. brassicae, however, was much higher than that expected on the basis of the sex ratio curve from different-sized G. mellonella. Larger hosts of each host species yielded larger wasps, indicating that the host size estimation by female wasps across different host species was incomplete or was not simple. These results suggested that P. luctuosa evaluated host size not only by physical measures such as dimension but also by other unknown measures. A possible explanation for the adaptiveness of different sex ratio responses by Pimpla luctuosa to different host species was discussed.     

Temperature-dependent growth rates for the cassava mealybug, Phenacoccus herreni, and two of its encyrtid parasitoids, Epidinocarsis diversicornis and Acerophagus coccois in Colombia

Developmental rates for Phenacoccus herreni Cox & Williams were determined at 18°, 20°, 22°, 25°, 30° and 35°C for the egg, all juvenile male and all juvenile female stadia. Longevity was determined for adult females and adult males. Developmental rates for the P. herreni parasitoid Epidinocarsis diversicornis (Howard) were determined at 18°, 20°, 25° and 30°C for the oviposition-to-mummy-formation period and the mummy-formation-to-adult-eclosion period. Developmental rates were determined for the P. herreni parasitoid Acerophagus coccois Smith for the same two life stages at 20°, 25° and 30°C. Least-squares-derived polynomial equations or logistic equations were fitted to each data set (except for A. coccois) so that rates could be interpolated for temperatures between observed points for use in an analysis of the impact of these parasitoids on population dynamics of P. herreni. Results of this analysis are presented separately.

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Résumé Les vitesses de développement des oeufs et de tous les stades larvaires mâles et femelles de P. herreni Cox & Williams ont été déterminées à 18°, 20°, 22°, 25°, 30° et 35°C. La longévité des adultes mâles et femelles a été déterminée. Les vitesses de développement de la ponte à la formation du cocon et de celleci à l'émergence de E. diversicornis Howard, encyrtide parasite de P. herreni ont été déterminées à 18°, 20°, 25° et 30°C. Il en a été de même pour un second encyrtide parasite A. coccois Smith à 20°, 25° et 30°C. Les équations polynomiales des derniers carrés dérivés et les équations logistiques ont été ajustées pour chaque lot de données (à l'exception de A. coccois) de façon à ce qu'elles aient pu être interpolées pour analyser l'impact de ces parasites sur la dynamique de population de P. herreni. Les résultats de cette analyse sont présentés à part.

     

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).     

Progeny production and adult longevity of the mealybug parasitoidsAnagyrus pseudococci,Leptomastix dactylopii,andLeptomastidea abnormis [Hym.: Encyrtidae] in relation to temperature

Progeny production increased and adult longevity decreased with rising temperature within the range 18°C to 30°C for the 3 mealybug parasitoidsAnagyrus pseudococci (Girault),Leptomastix dactylopii Howard andLeptomastidea abnormis (Girault). The Weibull distribution gave a good fit to survival curves for the 3 parasitoids and statistical comparison of Weibullb andc parameters at different temperatures allowed changes in the scale and shape of the curves to be detected. In general, ♀♀ lived longer than ♂♂ for all 3 species, except at high temperature. FemaleL. abnormis attained their maximum progeny production at 24°C and maintained this level up to 34°C. They lived longer than the other 2 parasitoid species at 30°C and showed a type I survival curve throuhout the range of temperatures examined.A. pseudococci andL. dactylopii both required high temperatures (30°C) to attain their maximal progeny production, but werepseudococci tended towards type II, with a larger proportion of the population dying within the first few days.L. dactylopii lived longest at 26°C, with ♀♀ showing a type I survival curve at all temperatures and ♂ survival curves changing from type I to type II at 30°C. The implications of these findings for the population dynamics of the different parasitoids are briefly discussed.      

Functional response of Euseius finlandicus and Amblyseius andersoni to Panonychus ulmi on apple and peach leaves in the laboratory

The functional response of adult females of the predatory mites Euseius (Amblyseius) finlandicus and Amblyseius andersoni to larvae and adult females of the fruit tree red spider mite Panonychus ulmi was determined on apple and peach leaf disks in the laboratory at 25°C and 16:8 (L:D). For adult females of P. ulmi the predation efficiency of E. finlandicus was higher on peach than on apple, whereas that of A. andersoni was higher on apple than on peach. Efficiency of predation on larvae of P. ulmi by either predator did not differ significantly between apple and peach. On both plants, A. andersoni had a higher predation rate than E. finlandicus on larvae of P. ulmi. It is concluded that in the laboratory the host plant has a substantial effect on predation efficiency of A. andersoni and E. finlandicus when they preyed on adults but not when they preyed on larvae of P. ulmi.     

Campoletis sonorensis [Hym.: Ichneumonidae] as a parasitoid ofSpodoptera frugiperda [Lep.: Noctuidae]: Host stage preference and functional response

The response of the parasitoidCampoletis sonorensis (Cameron) as a natural control agent of the fall armyworm,Spodoptera frugiperda (J. E. Smith), was evaluated at 6 host densities and 2 temperature regime. A type-II functional response was exhibited by femaleC. sonorensis in response to varying densities of all armyworn larvae at both temperatures. Significantly more larvae were parasitized at 25°C for the host densities of 60 and 75, than for the same densities at 30°C. Third instar fall armyworm larvae were found to be the preferred stage for parasitization byC. sonorensis. This research was supported by funds from HATCH project no. H-368 as allocated to the Georgia Agricultural Experiment Station.     

Induction of adult diapause and of low and high reproductive states in a parasitoid wasp,Ooencyrtus nezarae, by photoperiod and temperature

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) was reared on eggs ofRiptortus clavatus (Thunberg) (Heteroptera: Alydidae) at various temperatures under long-day (L16:D8) or short-day (L10:D14) conditions. There was no diapause during egg, larval or pupal stages under any set of conditions examined. However, at 15°C under short-day conditions, vitellogenesis was arrested in all adult females and they entered diapause. At 15°C under long-day conditions, or at 20°C under short-day conditions, some adult females entered diapause. Under the latter set of conditions, the adult females laid eggs but they laid fewer eggs than under long-day conditions, Even at 25°C, under short-day conditions, adult females laid fewer eggs than under long-day conditions, and this low rate of oviposition was attributed to the retarded development of ovaries. Diapause adults reared at 15°C were more resistant to low temperature than nondiapause adults reared at 25°C.