Competition between two primary parasitoids,Gyranusoidea tebygi Noyes and Anagyrus mangicola Noyes,attacking the mealybug Rastrococcus invadens Williams and the influence of a hyperparasitoid Chartocerus hyalipennis Hayat

Two primary parasitoids of the mealybug Rastrococcus invadens Williams, (Hemiptera: Pseudococcidae), Gyranusoidea tebygi Noyes and Anagyrus mangicola Noyes (Hymenoptera: Encyrtidae), were studied. Both primary parasitoids were capable of eliminating the mealybug host but on occasions the parasitoids went extinct before the mealybugs. Three of four parasitoids known to attack G. tebygi also attacked A. mangicola. The latter species was more heavily parasitized than the former, especially by the gregarious species Chartocerus hyalipennis Hayat. In competition experiments the presence of hyperparasitoids slightly slowed the speed of extinction of either the mealybug or the primary parasitoid. A. mangicola was heavily parasitized by C. hyalipennis and the primary went extinct while many suitable mealybug hosts were still available. There were two situations where the primary parasitoids were in direct competition; in the first G. tebygi was more successful than A. mangicola while the reverse was true for the second. It is likely that the superiority of either parasitoid would depend on particular conditions but the introduction of A. mangicola is unlikely to lessen the control being exerted in West Africa by G. tebygi.     

Establishment and spread of Gyranusoidea tebygi noyes and Anagyrus mangicola noyes (hymenoptera: encyrtidae), two biological control agents released against the mango mealybug Rastrococcus invadens williams (homoptera: pseudococcidae) in Africa

Two specific endophagous parasitoids Gyranusoidea tebygi and Anagyrus mangicola, of Indian origin, were mass‐reared at the International Institute of Tropical Agriculture in Cotonou and released against the mango mealybug Rastrococcus invadens, in collaboration with national biological control programmes. G. tebygi was released in the following countries: Benin, Gabon, Ghana, Nigeria, Sierra Leone and Zaire. In Togo, it had been released earlier and studied during another project. This parasitoid is now established in all areas infested by the mango mealybug. In addition, it established itself without previous release in Congo and Côte d'Ivoire. A. mangicola has been released in Benin, Gabon and Sierra Leone since 1991, and by mid‐1993 was recovered from a few sites. It seems locally established in southern Benin.     

Impact of Gyranusoidea tebygi Noyes (Hymenoptera: Encyrtidae) on the Mango Mealybug Rastrococcus invadens Williams (Homoptera: Pseudococcidae) in Nigeria

This study investigated the impact of released exotic mango mealybug parasitoid Gyranusoidea tebygi on mango mealybug Rastrococcus invadens in Nigeria. Observations were also made on the occurrence of the mealybug on other host plants in the surveyed areas. The monitoring exercise started in 1991 about 2 years after the first release in Ibadan. By 1997 and 1998, G. tebygi was found to have crossed all agro-ecological barriers to colonize the entire area of infestation nationwide on mango as well as other host plants. During this period, the populations of R. invadens had greatly decreased from between 11.0 and 98.0 mealybugs per leaf in 1991 to between 0.0 and 18.2 mealybugs per leaf in 1998. This fall was attributed to the activities of the released parasitoid. At many sampling sites in 1998, mealybugs were virtually absent on both mango and other host plants. Predators that were observed during the survey, were the coccinellids: Exochomus promptus Weise, Chilocorus nigritus (F) and Nephus spp. Larvae of chrysopid species, Ceratochrysa autica (Walker) and Plesiochrysa sp. The hyperparasitoids reared from mummies were Marietta leopardina Motsch (Aphelinidae), Chartocerus hyalipennis Hayat and Chartocerus subaeneus (Forster) (Signiphoridea).     

Impact of the Biological Control Agent Gyranusoidea tebygi Noyes (Hymenoptera: Encyrtidae) on the Mango Mealybug, Rastrococcus invadens Williams (Homoptera: Pseudococcidae), in Benin

The distribution of Rastrococcus invadens among different host plants and the impact of the mealybug on mango growth were investigated on 2067 trees in three surveys across all the ecological zones of Benin. The first survey started in 1989, less than 1 year after the first release of the exotic parasitoid Gyranusoidea tebygi. Within 3 years, G. tebygi had colonized the entire area of infestation, and was found on practically all infested mango trees as well as other infested host plants. By 1991, the incidence of R. invadens on the secondary host plants had declined significantly. The percentage of infested mango trees declined from 31.0% in 1989 to 17.5% in 1991, highest populations being found in the coastal savanna. During the same period, the mean percentage of infested mango trees having indigenous predators declined from 42.3 to 20.9%. Average mealybug densities declined steadily from 9.7 females/48 leaves in 1989, with 3.2% of all mango trees having densities above 100 mealybugs, to 6.4 females/48 leaves in 1991, with 1.3% of all trees having densities above 100 mealybugs. In multiple regression analyses, based on 23 meteorological, agronomic and plant variables, the duration of the parasitoid's presence proved to be a major factor. It influenced mealybug population densities and sooty mould incidence, which, in turn, affected the production of new leaves. In all analyses, the impact of rainfall, for example, on the sooty mould or the mealybug was less important than the effect of G. tebygi. The present study demonstrates for the first time on a large scale the impact of G. tebygi on R. invadens and, indirectly, on its main host plant, mango.     

Population Dynamics and Life Tables of the Mango Mealybug, Rastrococcus invadens Williams, and its Introduced Natural Enemy Gyranusoidea tebygi Noyes in Benin

Life table data for Rastrococcus invadens and its introduced natural enemy Gyranusoidea tebygi were obtained in the field and in the laboratory. The mealybug population's potential rate of increase ranged from 0.066/day to 0.078/day. The potential for increase of the parasitoid was double that of its host. Seasonal fluctuations in abundance of R. invadens were followed from 1988 to 1992 on mango trees in southern Benin. The population density of R. invadens decreased during the rainy seasons and peaked during the dry seasons. Mealybug field sex ratios were extremely variable, and the impact of such variability on the mealybug's potential rate of increase was analyzed. The populations of the exotic encyrtid G. tebygi, introduced into Benin in 1988 for control of the pest, were synchronized with the host populations. The spatial patterns of parasitism distribution in relation to the host population density were either independent or directly density-dependent, both at the tree level and for larger zones. However, reducing the scale of analysis resulted in different types of relationships. The impact of predators was a minor factor in the population dynamics of the mealybug. Four of the six species of hyper-parasitoids attacking mealybugs parasitized by G. tebygi developed high populations. In the two orchards studied, mealybug populations eventually collapsed and disappeared. This fact is discussed as being an indication that the biological control of the mango mealybug by G. tebygi was achieved by non-equilibrium local dynamics, and should be evaluated in a meta-population perspective.     

Influence of host plant on the mango mealybug,Rastrococcus invadens

Life history traits of the mango mealybug,Rastrococcus invadens Williams, were compared between two neighboring mango trees (Mangifera indica L.), one of which was heavily infested and the other slightly infested. On the infested tree, mealybug survival was high because of good feeding conditions and low escape. The pre-reproductive period of mealybugs on the heavily infested tree was shorter and total offspring production higher than that of mealybugs on the uninfested tree. This significantly affected the intrinsic rate of natural increase and explained the observed differences in population densities among the trees. The results obtained from an additional experiment using juvenile clones from the same two mango varieties, in which environmental factors were uniform, demonstrated the importance of plant genotype on mealybug size and survival.     

Interspecific discrimination and larval competition amongMicroplitis croceipes,Microplitis demolitor,Cotesia kazak (Hym.: Braconidae), andHyposoter didymator (Hym.: Ichneumonidae), parasitoids ofHeliothis virescens (Lep.: Noctuidae)

Interspecific host discrimination by adults, and larval competition among the endoparasitoidsMicroplitis croceipes (Cresson),Microplitis demolitor Wilkinson,Cotesia kazak (Telenga) andHyposoter didymator (Thunberg) were investigated usingHeliothis virescens (F.) as the host. In ovipositional choice tests, the mean number of encounters and ovipositions for unparasitized hosts was not significantly different from the mean number of encounters and ovipositions for parasitized hosts for each treatment combination (P>0.05). Thus, none of the parasitoid species discriminated between host larvae recently parasitized once by a female of another species und unparasitized hosts. However, in all but two cases, females did discriminate between unparasitized hosts and hosts in which an early first instar of the first-attacking species was developing.Cotesia kazak andH. didymator did not discriminate between unparasitized hosts and hosts parasitized by an early first instar ofM. demolitor. Larval competition among these parasitoid species was studied for three time intervals between the first and second species parasitization: 1) second species attack immediately (5–15 sec) after the first; 2) second species attack 24 h after the first; and 3) second species attack 48 h after the first. Time until egg eclosion was shortest forM. demolitor, thenC. kazak, thenM. croceipes, and longest forH. didymator. When the second parasitoid species attacked a host immediately after the first species, the species in which egg eclosion occurred first was the victor more frequently, except whenM. demolitor competed withC. kazak andH. didymator. With a 24 h delay between the first and second species to attack, the older first instar from the first parasitization usually outcompeted the younger first instar from the second attack. A first instar from the second species to attack generally outcompeted the second instar of the first species when the second parasitization had been delayed 48 h. Competiors were eliminated mainly by physical attack, butC. kazak andM. croceipes apparently also killedH. didymator eggs by physiological processes.     

Heterospecific larval competition and host discrimination in two species of aphid parasitoids: Aphidius ervi and Aphidius smithi

Females of the solitary aphid parasitoids Aphidius ervi Haliday and A. smithi Sharma & Subba Rao (Hymenoptera: Aphidiidae) discriminated between unparasitized pea aphids and those parasitized by the other species. Oviposition restraint varied with the attack sequence and the length of the interval between successive attacks. The tendency to reject a previously parasitized host increased with interval length; A. smithi females rarely oviposited in aphids that had been parasitized 30 h earlier by A. ervi. Early first-instar larvae of A. ervi physically attacked and killed older A. smithi larvae, and older A. ervi larvae killed younger A. smithi, possibly by physiological suppression. Neither species appeared to have a competitive advantage when their eggs hatched at the same time. The evolution of heterospecific host discrimination in A. ervi and A. smithi is discussed. It is suggested that avoidance of multiparasitism is adaptive for both parasitoid species: for A. smithi because it is the inferior larval competitor, and for A. ervi because immatures develop more slowly in multiparasitized than in initially unparasitized hosts.

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Compétition interspécifique et discrimination des hôtes chez deux parasitoïdes de pucerons: Aphidius ervi et A. smithi

Résumé Les femelles des parasitoïdes de pucerons: Aphidius ervi Haliday et A. smithi Sharma & Subba Rao (Hyméno. Aphidiidae) distinguent les pucerons du pois sains des parasités par d'autres espèces. La rétention de la ponte dépend de la séquence de l'attaque et du temps écoulé entre des attaques successives. La tendance au rejet d'un puceron précédemment parasité augmente avec l'importance du délai; A. smithi a rarement pondu dans des pucerons qui avaient été parasités 30 h avant par A. ervi. Les jeunes larves de premier stade de A. ervi ont attaqué physiquement et tué les larves plus âgées de A. smithi, et les larves plus âgées de A. ervi ont tué des larves plus jeunes de A. smithi par élimination physiologique. Aucune espèce ne semble avoir un avantage quand les oeufs ont éclos en même temps. L'évolution de la discrimination interspécifique de l'hôte chez A. ervi et A. smithi est discutée. On estime que la tendance à éviter le multiparasitisme est adaptative chez les 2 espèces: pour A. smithi parce qu'il est dominé dans la compétition larvaire et pour A. ervi parce que les larves se développent plus lentement dans un hôte multiparasité que dans un hôte initialement sain.

     

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.

     

Interaction betweenRastrococcus invadens [Hom.: Pseudococcidae] and two natural enemies

Two natural enemies of the mealybugRastrococcus invadens, the parasitoidGyranussoidea tebygi and the pathogenHirsutella cryptosclerotium were tested. Levels of parasitism by the parasitoid were reduced by the pathogen but overall mortality of the mealybug was greater both when agents were acting together.      

Interspecific competition between the principal larval parasitoids of the pine sawfly,Neodiprion sertifer (Geoff.) (Hym.: Diprionidae)

Summary Interspecific competition between the 3 principal larval parasitoids of the pine sawfly, Neodiprion sertifer, is of common occurrence when total larval parasitism, and hence multiple attack rates, are high. At the intrinsic level, the ectoparasitoid Exenterus abruptorius is superior to the 2 endoparasitoids, Lophyroplectus luteator and Lamachus eques, respectively, whereas L. luteator is superior to L. eques only. During mass outbreaks of the host competition between Exenterus and Lophyroplectus is most intense, whilst Lamachus fails to build up in the presence of its 2 competitors (Fig. 1). However, due to its superior host finding ability, Lamachus dominates in areas of low host densities where the 2 other species are less efficient (Table 1). L. luteator frequently sustains heavy losses in direct competition with E. abruptorius, but these can be largely compensated in the next generation because its fertility is about 5-times higher than that of Exenterus.     

Intraspecific host discrimination and larval competition inMicroplitis croceipes,Microplitis demolitor,Cotesia kazak (HYM.: Braconidae) andHyposoter didymator HYM: Ichneumonidae), parasitoids ofHeliothis virescens (LEP.: Noctuidae)

Intraspecific host discrimination and larval competition were studied forMicroplitis croceipes (Cresson),Microplitis demolitor Wilkinson,Cotesia kazak (Telenga), andHyposoter didymator (Thunberg), solitary endoparasitoids of the tobacco budworm,Heliothis virescens (F.). In ovipositional choice tests between unparasitized and parasitized hosts, the mean number of ovipositions for unparasitized hosts was significantly higher than the mean number of ovipositions for hosts parasitized once by a conspecific female forC. kazak andH. didymator, demonstrating that females of these two species discriminate against hosts recently (within a few seconds) parasitized by a conspecific female. No significant difference in oviposition occurred between these two kinds of hosts forM. croceipes andM. demolitor. Mean percent parasitization by a second conspecific female was determined at 24, 48, and 72 h delays in time between the first and second female attack, and with no delay. Except for the 0 h time delay forC. kazak andH. didymator, percent parasitization by a second conspecific female generally decreased as the delay in time between the first and second female attack increased. When the second parasitization immediately followed the first, one parasitoid larva always eliminated the other by physical combat. With a 24 or 48 h delay between the first and second parasitization, the younger larva was the victor over the older larva forM. croceipes, M. demolitor andC. kazak in at least 50% of the cases. Elimination of older larvae by younger larva was by physical attack. However, forH. didymator, the older instar was the victor, and elimination of younger larvae by older larvae was probably through physiological processes. Further, older larvae ofH. didymator apparently killed the eggs of the second female by physiological processes.      

Competition for hosts between two hyperparasitoids of aphids,Dendrocerus laticeps andDendrocerus carpenteri (Hymenoptera: Megaspilidae): The benefit of interspecific host discrimination

The two hyperparasitoids of aphids, Dendrocerus carpenteri(Curtis) and Dendrocerus laticeps(Hedicke), differ in their interspecific host discrimination behavior: D. carpenterireadily accepts hosts for oviposition previously parasitized by D. laticeps,whereas D. laticepsoften refrains from ovipositing on hosts previously parasitized by D. carpenteri.The two species also differ in their competitive abilities: D. carpenteriis competitively superior to D. laticepswhen the time between ovipositions is 1 h. D. carpenteri's superiority probably arises from (a) the action of the venom produced by the female to paralyze the host, as evidenced by the comparison of emergence patterns from naturally vs artificially parasitized hosts, and (b) its more rapid egg development. Because of its competitive abilities, D. carpenterigains only limited benefit from discriminating interspecifically, at least when encountering a host parasitized by a competitively inferior species, such as D. laticeps.In contrast, D. laticepsgains considerable benefit from recognizing hosts previously parasitized by D. carpenteri,as the probability to produce offspring from such hosts is greatly reduced.     

Decision to superparasitize based on larval survival: competition between aphid parasitoids Aphidius ervi and Aphidius smithi

Oviposition by a parasitoid wasp in an already parasitized host (= superparasitism) may result in larval competition and the loss of offspring. Consequently, in solitary species, the decision to superparasitize should be based on the probability of offspring survival. Females of Aphidius ervi Haliday and A. smithi Sharma & Subba Rao (Hymenoptera: Aphidiidae), two parasitoids of the pea aphid, discriminated between conspecific-and heterospecific-parasitized hosts. Both species showed partial preference for aphids previously attacked by A. smithi, a result predicted from the fact that A. ervi is superior to A. smithi in larval competition under most conditions. Females of A. smithi selectively attacked conspecific-parasitized pea aphids when given a choice between these and self-parasitized hosts. Results indicated that females of both species responded to an external pheromone-like marker to discriminate between selfconspecific-and heterospecific-parasitized aphids. In addition, A. smithi may use internal cues to recognize, and avoid oviposition in, aphids previously parasitized by A. ervi.

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Résumé L'oviposition par une guêpe parasite dans un hôte qui a déjà été parasité (= superparasitisme) pourrait conduire à une compétition entre larves et à une perte de progéniture. Par conséquent, dans les espèces solitaires, la décision de superparasiter devrait être basée sur la probabilité de survie de la progéniture. Les femelles de Aphidius ervi Haliday et de A. smithi Sharma & Subba Rao (Hymenoptera: Aphidiidae), deux parasites du puceron du pois, ont discriminé entre les hôtes parasités par des individus de la même espèce et ceux parasités par des individus d'une espèce différente. Les deux espèces de parasite ont montré une préférence partielle pour les pucerons préalablement parasités par A. smithi, résultat qui était prévisible du fait que, dans la majorité des cas, A. ervi est supérieur à A. smithi dans la compétition entre larves. Les femelles de A. smithi ont sélectivement attaqué les pucerons parasités par un individu de la même espèce lorsqu'il leur était donné de choisir parmi ceux-ci et ceux parasités par les même individu. Les résultats indiquaient que les femelles des deux espèces répondaient à une sorte de phéromone externe pour distinguer les pucerons parasités par le même individu de ceux parasités par un individu de la même espèce et ceux parasités d'une autre espèce. En plus, A. smithi pourrait utiliser des signaux internes pour reconnaître et éviter l'oviposition dans les pucerons préalablement parasités par A. ervi.

     

Efficiency,intrinsic competition and interspecific host discrimination of Copidosoma desantisi and Trichogramma evanescens,two parasitoids of Phthorimaea operculella

Trichogramma evanescens West. (Hymenoptera: Trichogrammatidae) and Copidosoma desantisi Annecke & Mynhardt (Hymenoptera: Encyrtidae) are potential parasitoids of the potato tuber moth (PTM), Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae) in Egypt. Discrimination of a parasitized host from an unparasitized host would prevent wasting of time, eggs and reduce competition with conspecifics or heterospecifics. Therefore, we evaluated interspecific host discrimination, multiparasitism and intrinsic competition between the two wasp species. In a choice test, females of T. evanescens showed high interspecific host discrimination only when they were offered 2-day-old C. desantisi parasitized and unparasitized PTM eggs. In contrast, C. desantisi showed high host discrimination and preferred unparasitized eggs to PTM eggs harboring 2-h- or 2-day-old T. evanescens’ eggs. We also evaluated the effect of different introduction sequences on the efficacy of the two wasps. Dissection data indicated that the two parasitoids had a negative impact on each other. There was a significant reduction in the total number of deposited eggs as well as total number of parasitized hosts by each parasitoid. Regarding the rearing experiment, the total number of T. evanescens-induced black eggs or C. desantisi formed mummies in combined treatments was significantly lower than in single parasitoid treatments (control). Moreover, C. desantisi was inferior and did not develop from any multiparasitized host regardless of oviposition order. It was suggested that combined release of the two wasps would not elevate rate of parasitism over that of single parasitoid treatments and competition between them would reduce their efficacy.     

Intraspecific host discrimination and larval competition inMicroplitis croceipes,Microplitis demolitor,Cotesia kazak (Hym.: Braconidae) andHyposoter didymator (Hym.: Ichneumonidae), parasitoids ofHeliothis virescens (Lep.: Noctuidae)

Intraspecific host discrimination and larval competition were studied forMicroplitis croceipes (Cresson),Microplitis demolitor Wilkinson,Cotesia kazak (Telenga), andHyposoter didymator (Thunberg), solitary endoparasitoids of the tobacco budworm,Heliothis virescens (F.). In ovipositional choice tests between unparasitized and parasitized hosts, the mean number of ovipositions for unparasitized hosts was significantly higher than the mean number of ovipositions for hosts parasitized once by a conspecific female forC. kazak andH. didymator, demonstrating that females of these 2 species discriminate against hosts recently (within a few seconds) parasitized by a conspecific female. No significant difference in oviposition occurred between these 2 kinds of hosts forM. croceipes andM. demolitor. Mean percent parasitization by a 2^nd conspecific female was determined at 24, 48, and 72-h delays in time between the 1^rst and 2^nd female attack, and with no delay. Except for the 0 h time delay forC. kazak andH. didymator, percent parasitization by a 2^nd conspecific female generally decreased as the delay in time between the 1^rst and 2^nd female attack increased. When the 2^nd parasitization immediately followed the 1^rst, one parasitoid larva always eliminated the other by physical combat. With a 24 or 48 h delay between the 1^rst and 2^nd parasitization, the younger larva was the victor over the older larva forM. croceipes, M. demolitor andC. kazak in at least 50% of the cases. Elimination of older larvae by younger larvae was by physical attack. However, forH. didymator, the older instar was the victor, and elimination of younger larvae by older larvae was probably through physiological processes. Further, older larvae ofH. didymator apparently killed the eggs of the 2^nd female by physiological processes.      

Interspecific interference between Apoanagyrus lopezi and A. diversicornis, parasitoids of the cassava mealybug Phenacoccus manihoti

The parasitoids Apoanagyrus lopezi De Santis and A. diversicornis (Howard) (Hymenoptera: Encyrtidae) have been introduced into Africa for the biological control of the cassava mealybug Phenacoccus manihoti Matile-Ferrero (Homoptera: Pseudococcidae). We have studied competition between these species to investigate if they can coexist. Here we report on the influence of the simultaneous presence of non-conspecific adult females on searching efficiency on patches. Wasps of either species foraged on discs of cassava leaf with mealybugs, while at the same time different numbers of non-conspecifics were also depleting the patch. Patch area per parasitoid and number of hosts available to each parasitoid were equal in all treatments.In both species, the presence of other foragers clearly affected several aspects of the parasitoids' behaviour. Patch residence time increased with the number of non-conspecifics in A. diversicornis. In both parasitoid species, the proportion of hosts left unparasitized after the patch visit decreased with increasing numbers of females on the patch. The proportions of super- and multiparasitism did not change with the number of females. Both species produced more offspring during a patch visit in the presence of more non-conspecifics. These behavioural changes did not, however, lead to a change in the offspring production rate on patches. A. diversicornis produced offspring at a rate three times that of A. lopezi when one A. lopezi and one A. diversicornis foraged simultaneously. This is the first report of an aspect of interspecific competition where A. diversicornis has an advantage over A. lopezi. Interference between adult females thus promotes coexistence of the two species on P. manihoti.     

Host discrimination and larval competition in the aphid parasite Ephedrus californicus

Naive and experienced females of Ephedrus californicus Baker (Hymenoptera: Aphidiidae) were tested for their ability to discriminate between parasitized and unparasitized pea aphids, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae). Attacks lasting 6 s generally resulted in oviposition; the average length was 11.8 s. The proportion of parasitized aphids that was rejected varied with the interval length between attacks. It is suggested that host discrimination is time-dependent and can be induced by a pheromone-like external marker left by a first-attacking female (0–9 h), or by changes in host quality associated with parasite development ( 14 h). Experienced, but not naive, females responded to the external marker, which became less effective with time. In superparasitized aphids, the older of two E. californicus larvae usually eliminated a younger competitor; but younger larvae survived under certain conditions. Mechanisms for the elimination of supernumerary larvae varied with the relative developmental stage of the competitors and included physical combat and physiological suppression. Host instar had no effect on larval competition or the female's ability to discriminate.

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Résumé L'aptitude à choisir entre Acyrthosiphon pisum Harris (Hom. Aphididae) parasités ou non a été examinée chez des femelles nouvelles ou expérimentées d'E. californicus Baker (Hym. Aphidiidae). Les attaques durant 6 s étaient généralement suivies de pontes; la durée moyenne d'une attaque était 11,8 s. La proportion de pucerons parasités refusés, variait avec le laps de temps écoulé entre des attaques. Les femelles nouvelles aussi bien qu'expérimentées rejetaient généralement les pucerons contenant les stades les plus âgés de parasites (13–15 h). Cependant, seules les femelles expérimentées évitaient de pondre dans des pucerons attaqués par une autre dans les 9 h précédentes. Quand des femelles expérimentées pouvaient choisir entre 20 pucerons parasités et 20 pucerons sains, en moyenne 73% des 20 premières attaques étaient sur pucerons sains pour des intervalles 9 h. Des attaques non suivies de pontes, c'est-à-dire durant 5 s, n'intervenaient que pour 3% parmi toutes les premières attaques sur pucerons sains; ces attaques atteignaient ultérieurement 17% pour les pucerons sains et 40% pour les parasités.On en a déduit que la sélection des hôtes est temporelle et peut-être induite par un marqueur externe type phéromone, laissé par la lère femelle attaquante (0–9 h), ou par un changement dans la qualité de l'hôte lié au développement du parasite. Les femelles expérimentées, et non les nouvelles, répondaient au marqueur externe, dont l'efficacité diminue avec le temps.Chez les pucerons superparasités, les mécanismes pour l'élimination des larves en surnombre ont varié avec le stade larvaire des compétiteurs, ils comportaient un combat physique et une suppression physiologique. La larve de E. californicus la plus âgée éliminait généralement une compétitrice plus jeune. Cependant les larves les plus jeunes ont survécu dans certaines conditions; c'est-à-dire quand les premiers stades mandibulés luttaient contre les 2e et. 3e stades sans mandibules. Le stade de l'hôte était sans effet sur la compétition ou sur l'aptitude au choix des femelles.

     

Interspecific host discrimination and competition in Apoanagyrus (Epidinocarsis) lopezi and A.(E.)diversicornis, parasitoids of the cassava mealybug Phenacoccus manihoti

Abstract.

• 1 Choice experiments on interspecific host discrimination in A.lopezi and A. diversicornis were carried out on discs of cassava leaf containing four hosts (P.manihoti) that had been parasitized by the other species and four unparasitized hosts.
• 2 A.lopezi accepted both host types equally for oviposition, whereas A.diversicornis accepted fewer hosts that had been parasitized by A.lopezi than unparasitized ones. A.diversicornis is therefore capable of interspecific host discrimination, but such a capability was not demonstrated for A.lopezi.
• 3 Survival probability in singly parasitized hosts was 0.85 for both parasitoid species. When the time interval between ovipositions was 2 h or less, survival in multiparasitized hosts was 0.68 for A.lopezi and 0.17 for A.diversicornis, irrespective of priority. Increasing A.lopezi priority to 24±2h did not increase A.lopezi survival. A.diversicornis survival, however, increased to 0.43 when A.diversicornis was given 24 ± 2 h priority. A.diversicornis eggs took 19 h longer than A.lopezi eggs to hatch. This could explain the difference in competitive abilities in multiparasitized hosts.
• 4 The observed difference in host selection behaviour between A.lopezi and A.diversicornis is in accordance with the different benefits of multiparasitism: A.lopezi gains more than A.diversicornis because of its superior within-host competitive abilities.
• 5 Neither species avoided multiparasitism completely. The low survival probability of A.diversicornis in multiparasitized hosts may partly explain its failure to establish when introduced into Africa as part of a biological control programme of P.manihoti.

     

Sunflower stem weevil and its larval parasitoids in the Central and Northern Plains of the USA

The sunflower stem weevil, Cylindrocopturus adspersus (LeConte) (Coleoptera:Curculionidae), is apest of cultivated sunflower (Helianthusannuus L) from the southern to the northernGreat Plains. The incidence of weevilinfestation in fields from the six differentstates sampled during 1996 and 1997 ranged from33% (Minnesota) to 100% (Kansas, Colorado,Nebraska). Weevil populations in the fieldssampled were statistically greater in thecentral Plains (Colorado, Kansas, Nebraska)with a mean of 12.3 and 19.5 larvae per stalkcompared with the northern Plains (North andSouth Dakota, Minnesota) of 0.7 and 1.3 larvaeper stalk in 1996 and 1997, respectively.Parasitization of weevils varied from field tofield ranging from 1 to 100%, but was usuallyless than 20%. The nine species oflarval parasitoids recovered were allHymenoptera and included: Nealioluscurculionis (Fitch), N. collaris(Brues), Bracon sp. (Braconidae); Neocatolaccus tylodermae (Ashmead), Chlorocytus sp., Pteromalus sp.(Pteromalidae); Quadrastichus ainslieiGahan (Eulophidae), Eurytoma tylodermatisAshmead (Eurytomidae); and Eupelmus sp.(Eupelmidae). Nealiolus curculionis wasthe most prevalent parasitoid reared from C. adspersus, and it was recovered from allstates sampled. Parasitoid species richness wasgreatest in the central Plains. Thereduced number of parasitoid species foundattacking C. adspersus in the northernPlains may be caused by low host populationlevels, slow migration by parasitoids into theregion, or lack of adaptation to climaticconditions. Additional work to understandthe population dynamics of the parasitoidcomplex associated with C. adspersus mayresult in improved biological control of thesunflower stem weevil in cultivatedsunflower.