The effect of superparasitism on development of the solitary parasitoid wasp, Venturia canescens (Hymenoptera: Ichneumonidae)

Abstract.

• 1 The effects of superparasitism on the rate of development, adult size and mortality of Venturia canescens (Hymenoptera: Ichneumonidae) were investigated. Parasitoids were reared from third (L3) and fifth (L5) instars of one of its hosts, Plodia interpunctella (Lepidoptera: Pyralidae) containing one, two or four parasitoid eggs.
• 2 Superparasitism increased the development time of Venturia reared from both instars, but the developmental delay was more evident in wasps from L5 hosts.
• 3 The size of parasitoids from L3 hosts was unaffected by egg number, but wasps from both superparasitized L5 treatments were significantly smaller than those from singly parasitized hosts.
• 4 Parasitoid mortality was significantly higher in L5 than L3, but within instars did not differ significantly with egg number.
• 5 The results confirm that superparasitism may affect the fitness of both the adult female wasp and her progeny, and should therefore be incorporated into models of superparasitism as an adaptive foraging strategy.

     

The effect of host size on quality attributes of the egg parasitoid,Trichogramma pretiosum

In a study of the quality ofTrichogramma pretiosum Riley (Hymenoptera: Trichogrammidae), we compared female wasps emerging from natural hosts, parasitized in the laboratory or the field with those emerging from factitious hosts used for commercial mass production. Females from the natural hosts were larger, more fecund, and longer lived than those from the factitious hosts. Compared to small females, large female wasps are substantially more fecund when honey (carbohydrate) is available but marginally more fecund when honey is unavailable. The size of a femaleT. pretiosum depends on two factors: the size of the host egg from which it emerges even when the wasp was gregarious, and the number of conspecifics that emerge with it. The similarities in the size distribution of female wasps emerging from natural hosts, in conjunction with the mechanism by whichTrichogramma measure host size and allocate eggs accordingly, suggests the hypothesis that size related components of fitness in femaleT. pretiosum are under strong selection in the field.     

Host behaviour and its influence on foraging and acceptance by the solitary parasitoid wasp,Venturia canescens (Hym: Ichneumonidae)

Behavioural interactions between the solitary koinobiont parasitoid,Venturia canescens, and two of its hosts,Plodia interpunctella andCorcyra cephalonica, were investigated. The response of both hosts to simulated antennation using a two-haired brush was examined over instars 3 (L3) to 5 (L5). YoungP. interpunctella larvae predominantly adopted escape tactics (writhe, trash) whereas L5P. interpunctella usually froze after the stimulus was applied. L3C. cephalonica larvae were more aggressive (headrear, flick) thanP. interpunctella in response to the application of the stimulus, but olderC. cephalonica responded less aggressively than in earlier instars. AlthoughV. canescens readily jabbed its ovipositor at both hosts after antennation,P. interpunctella was considerably more susceptible to parasitoid attack thanC. cephalonica, irrespective of size in the final (L5) instar.C. cephalonica, the larger, more aggressive host, actively resisted parasitism whereasP. interpunctella responded much more passively after parasitoid contact. Parasitoids examined and jabbed their ovipositors at dead hosts, but this behaviour was not sustained, implying that host movement stimulates parasitoid attack. On patches containingV. canescens, L5C. cephalonica andP. interpunctella, mostP. interpunctella larvae responded by freezing after parasitoid contact.P. interpunctella that froze usually avoided parasitism, whereas larvae that attempted to escape by crawling were pursued with vigour byV. canescens and usually parasitized. Irrespective of behaviour after parasitoid contact,C. cephalonia displayed more aggressive behaviour and had much greater success in warding off parasitoid attack. Host acceptance byV. canescens is clearly affected by the size and species of the host it attacks. The influence of host defensive behaviour is discussed in relation to the evolution of parasitoid counter-defences and oviposition strategies.     

Influence of initial egg density and host size on the development of the gregarious parasitoid <Emphasis Type="Italic">Bracon hebetor</Emphasis> on three different host species

Bracon hebetor (Say) (Hymenoptera : Braconidae) is a gregarious parasitoid that attacks a variety of important lepidopterous pests of stored product and in the field. In this study the effect of host species, size and larval competition on parasitoid size, survival and development were investigated. In laboratory studies, wasp eggs at a range of densities, were placed on larvae of different weight of three Lepidoptera host species namely Adoxophyes orana (Fischer von Röslerstamm, Tortricidae), Plodia interpunctella (Hubner, Pyralidae) and, Lobesia botrana(Dennis & Schiffermueller, Tortricidae). On A. orana survival of immature parasitoids was very low at all densities and different host weights. On L. botrana survival progressively reduced as egg density increased at both host weights examined for this host. Survival on P. interpunctella was significantly affected by egg density but not by host weight. Initial egg density had a significant effect on the size of emerging adults from each rearing host. Smaller adult parasitoids emerged as egg density per larva increased. Larval host weight of P. interpunctella and A. orana had a significant effect on the size of emerging adult parasitoids mainly at the higher egg densities used in these experiments. The above results of host quality on fitness of parasitoid are discussed.     

Reciprocal influences and costs of parasitism on the development of Corcyra cephalonica and its endoparasitoid Venturia canescens

Many endoparasitoids develop successfully within a range of host instars. Parasitoid survival is highest when parasitism is initiated in earlier host instars, due to age-related changes in internal (physiological) host defences. Most studies examining fitness-related costs associated with differences in host instar have concentrated on the parasitoid, ignoring the effects of parasitism on the development of surviving hosts that have encapsulated parasitoid eggs. A laboratory experiment was undertaken examining fitness-related costs associated with encapsulation of Venturia canescens (Hymenoptera: Ichneumonidae) eggs by fifth (L5) instar larvae of Corcyra cephalonica (Lepidoptera: Pyralidae). Growth and development of both host and parasitoid were monitored in C. cephalonica larvae containing 0, 1, 2, or 4 parasitoid eggs. Adult size and fecundity of C. cephalonica did not vary with the number of eggs per host. However, there was a distinct increase in host mortality with egg number, although most parasitoids emerged from hosts containing a single egg. The most dramatic effect on the host was a highly significant increase in development time from parasitism to adult eclosion, with hosts containing 4 parasitoid eggs taking over 2.5 days longer to complete development than unparasitized larvae. The egg-to-adult development time and size of adult V. canescens did not vary with egg number per host, as demonstrated in a previous experiment using a different host (Plodia interpunctella). The results described here show that there are fitness-related costs to the host associated with resistance to parasitism.     

Flexible larval development and the timing of destructive feeding by a solitary endoparasitoid: an optimal foraging problem in evolutionary perspective

1. In studying the evolution of life-history strategies in parasitoids, considerable attention has been paid to the relationship between host quality and parasitoid fitness. Various workers have reported that host quality influences parasitoid size, development time, and survival. Because body size is frequently correlated with fecundity, longevity, and host-finding ability in parasitoids, this parameter is usually considered to be the main target of selection. 2. In koinobiont parasitoids that consume the entire host before pupation, adult parasitoid size and development time are often strongly correlated with host size at the time when it is developmentally arrested through destructive feeding by the parasitoid larva. 3. Here, a mathematical model is proposed to describe the larval feeding behaviour of the solitary koinobiont endoparasitoid Venturia canescens in four larval stadia of its host Plodia interpunctella. In particular, the model describes how adult size, represented by an exponential growth rate, and development time are traded off when the parasitoid develops in nutritionally suboptimal second stadium hosts. 4. Using a graphical model, the different conditions faced by V. canescens during development in various host species of greatly differing mass are illustrated. 5. It is argued that the relative importance of size and development time on parasitoid fitness is determined by ecological and biological characteristics of both host and parasitoid, and it is suggested that there may be correlations between life-history traits and host-utilisation strategies among koinobionts.     

Differential cannibalism and population dynamics in a host-parasitoid system

The effects of host cannibalism on a host-parasitoid system were explored through experiment and modelling. In individual encounters between parasitized and unparasitized Plodia interpunctella larvae, parasitized larvae were more likely to be cannibalized. Inclusion of this differential cannibalism into a simple Lotka-Volterra-type model of host-parasitoid population dynamics generates alternative stable states-including stable coexistence and extinction of the parasitoid — which depend on starting conditions. Possible mechanisms for differential cannibalism, and its implications for studies of host-parasitoid populations and biological control programmes are discussed.     

The effect of host spacing on the clutch size and parasitization rate of Trichogramma minutum

The clutch size of the parasitoid wasp Trichogramma minutum Riley (Hymenoptera: Chalcidoidea) is in part adjusted in response to the spatial distribution of its insect egg hosts. This paper describes the effects on progeny allocation of differences in the distance separating single hosts, and a possible mechanism is proposed. The number of progeny laid into a single host decreases with reduced interhost distance. The effect is not due to superparasitism of more widely spaced hosts, since single hosts which the wasps are allowed to parasitize only once receive only as many eggs as the most widely spaced host. Furthermore, no correlation was found between the number of hosts parasitized and the mean clutch size for each wasp, indicating that the wasps do not simply reduce progeny allocation with successive host encouters. Instead, the wasps may use a measure of the frequency of host encouter, for example the time or distance between hosts, as a cue to set cluch size. Comparisons of clutch size for first and second hosts parasitized showed that there is an initial large reduction in clutch size, after which all subsequent hosts parasitized are allocated a constant, reduced number of progeny. The implications of changes in clutch size for the parasitization rate of the wasps are discussed.

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Die bestimmung der wirtsdichte durch die parasitische wespe Trichogramma minutum

Zusammenfassung Bei der parasitischen Wespe Trichogramma minutum hängt die Menge abgelegter Eier teilweise von der räumlichen Verteilung seines Insektenwirtes ab. In dieser Arbeit wird der Einfluss von unterschiedlichen Abständen zwischen den Wirtstieren auf die Anzahl von Nachkommen pro Ei beschrieben und mögliche Mechanismen zur Bestimmung der Wirtsdichte vorgeschlagen. Die Anzahl von Nachkommen pro Wirtsei verringert sich mit kleiner werdendem Abstand zwischen den Wirtseiern. Es wird vermutet dass die Wespen ein Mass für die Häufigkeit von Wirtsbegegnungen, z.B. Zeit oder Abstand zwischen Wirten als Schlüssel für die Bestimmung der Menge von Nachkommen benutzen. Diese Erscheinung kann nicht auf Superparasitismus von weiter entfernten Wirten zurückgeführt werden. Einzelne Wirte, die nur einmal von den Wespen parasitiert werden durften, erhielten die gleiche Anzahl von Eiern, wie die am weitesten verteilten Wirte. Darüber hinaus wurde kein Zusammenhang zwischen der Anzahl parasitierter Wirte und der durchschnittlichen Grösse der Nachkommenschaft pro Wespe gefunden. Das weist darauf hin, dass die Wespen nicht einfach mit jeder weiteren Wirtsbegegnung ihre Menge zugewiesener Nachkommen verringern. Schliesslich wird der Einfluss von unterschiedlicher Anzahl von Nachkommen auf die Wirtsmortalität diskutiert.

     

Lifetime Reproductive Success in the Solitary Endoparasitoid, Venturia canescens

Parasitoid wasps have long been considered excellent organisms in studies examining the evolution of reproductive and life-history strategies. In examining the lifetime reproductive success of parasitoids in the laboratory, most investigations have provided the insects with excess hosts and food, where they exist in a relatively constraint-free environment. Importantly, these conditions may not accurately reflect the true heterogeneity of natural systems, where suitable hosts and food sources are likely to be limiting. This study examines the influence of differences in host and food availability on reproductive and life-history parameters in an asexual strain of the solitary endoparasitoid, Venturia canescens (Hymenoptera: Ichneumonidae). Lifetime reproductive success in V. canescens was measured in response to temporal variations in host and food (honey solution) access. Cohorts of parasitoids were provided with 200 fifth-instar larvae of the Indian meal moth, Plodia interpunctella (Lepidoptera: Pyralidae), and food for variable periods daily after eclosion. V. canescens is synovigenic, and host-deprived wasps continued to mature eggs over the first few days after eclosion until the egg storage capacity was reached in the oviducts. When these wasps were subsequently provided with hosts, oogenesis resumed and continued until later in adult life. Constantly fed wasps lived longer and produced more progeny than wasps from cohorts which were alternately fed and starved or were starved from eclosion. Moreover, wasps with constant host and food access produced most progeny early in life and usually experienced prolonged periods of postreproductive survival. In contrast, the reproductive period of wasps with limited host access was more evenly distributed throughout the adult life. Consequently, the cumulative progeny production by V. canescens with constant food access was fairly uniform irrespective of host availability. Longevity and fecundity in V. canescens were positively correlated with adult size. However, variable host access had little effect on the longevity of wasps which were constantly supplied with honey. Over the first 2 days of adult life, variation in food access also had no effect on progeny production by V. canescens. We argue that manipulating temporal host and food access to parasitoids in the laboratory more closely approximates natural conditions, where these resources are likely to be spatially separated. Moreover, our findings suggest that many highly synovigenic parasitoids like V. canescens, which produce microtype (=hydropic) eggs, have a considerably higher reproductive potential than ovary dissections have revealed. Our findings are discussed in relation to life-history evolution in the parasitic Hymenoptera.     

砂藓RcPLD基因的抗旱功能分析

砂藓(Racomitrium canescens)是一种具有极强耐脱水性的苔藓植物,编码磷脂酶D的基因RcPLD能够在砂藓的脱水和复水过程中产生显著的表达响应,它可能参与了砂藓的强耐脱水性功能。该研究使用已克隆的RcPLD编码序列构建拟南芥(Arabidopsis thaliana)过量表达转基因株系rcpld-oe,初步考察过表达株系的干旱胁迫耐受能力及其相关的生理生化指标,分析RcPLD增强拟南芥抗旱性的机制。结果表明:(1)利用已克隆的RcPLD编码序列构建了植物中的过表达载体,成功构建了RcPLD的过表达转基因拟南芥株系rcpld-oe,并获得了多个T_3代rcpld-oe纯合体株系。(2)在正常生长条件下,rcpld-oe株系T_3代纯合体植株比野生型拟南芥植株体积小,但营养生长期较长,抽薹较晚,莲座叶衰老速率较慢;在干旱处理条件下,rcpld-oe株系表现出比野生型拟南芥更强的干旱耐受能力。(3)在干旱胁迫处理过程中,rcpld-oe株系莲座叶的水分散失速率降低,可能在一定程度上降低了干旱对膜完整性的损伤和光合作用的抑制,但其渗透调节物质含量的变化相对较小。研究发现,在干旱胁迫条件下,rcpld-oe植株莲座叶的水分散失速率和光合作用抑制程度显著降低,从而表现出明显强于野生型的干旱耐受能力,这为后续RcPLD功能的深入研究和更多砂藓抗旱功能基因的挖掘奠定了基础。     

Superparasitism in the solitary ectoparasitoid Aptesis nigrocincta: the influence of egg load and host encounter rate

Theory predicts that the acceptance of hosts already parasitized by a conspecific will depend both on egg load and the availability of hosts. In the present laboratory study, we tested the effect of egg load and host encounter rate on the propensity of superparasitism in the solitary parasitoid Aptesis nigrocincta Gravenhorst (Hymenoptera, Ichneumonidae), a synovigeneous ectoparasitoid of prepupae of the European Apple Sawfly. Parasitoid females carry few voluminous eggs at a time and the egg maturation rate is less than one egg per day. Egg load was manipulated by giving females access to hosts one week prior to the start of treatments and host availability by giving females access to either one host cocoon every day or every other day. In the first treatment where females had a high egg load of 5.3 egg in their ovaries and encountered host cocoons at low rates, we found that parasitized hosts were accepted to the same degree as healthy hosts. In females with significantly decreased egg load (3.8 eggs) encountering hosts at the same rate we found a slight but non-significant decrease in the acceptance of parasitized hosts compared with healthy hosts. In contrast, A. nigrocincta females accepted significantly fewer parasitized hosts at a high host encounter rate that would lead them to the point of egg limitation in the near future. Within the range of egg loads tested, the host encounter rate appears to be the most important determinant for a females decision to oviposit onto hosts already parasitized by a conspecific.     

Host-habitat conditioning in the eulophid egg parasitoid, Edovum puttleri: effects of herbivore's host and its food plant

The egg parasitoid, Edovum puttleri Grissell, was cultured from eggs of two Leptinotarsa hosts, L. decemlineata (Say) and L. texana Schaeffer, and experienced on eggs from three L. decemlineata food plants, namely potato, Solanum tuberosum tuberosum L., eggplant, S. melongena L., and tomato, Lycopersicon esculentum Mill. Host preference of the various cultures was determined using choice tests. parasitoid choices were influenced by the Leptinotarsa rearing host. Wasps, reared from L. decemlineata eggs from a mixture of all three food plants and experienced on egg masses from all three food plants, preferred to parasitize egg masses from potato. Wasps, reared from L. texana eggs from silver leaf nightshade, S. elaeagnifolium L., and experienced on egg masses from all three food plants, preferred to parasitize egg masses from tomato. Choices were also influenced by the herbivore's food plant that wasps were conditioned to. Wasps, reared from L. texana and experienced on egg masses from potato, preferred to parasitize egg masses from potato. Wasps, reared from L. decemlineata on potato and experienced on egg masses from potato, preferred to parasitize egg masses from potato. Wasps, reared from L. decemlineata on tomato and experienced on egg masses from tomato, did not display a distinct host preference. Within-egg mass parasitism (i.e., number of eggs per mass that were parasitized, probed, etc.) did not consistently differ among the three food plants. The implications of these data in parasitoid ecology and biological control programs for L. decemlineata are discussed.

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Résumé L'oophage, E. puttleri a été élevé à partir des oeufs de Leptinotarsa decemlineata et L. texana. Il a été mis en expérience sur l'une des 3 plantes consommées par L. decemlineata: Solanum tuberosum tuberosum, S. melongena et Lycopersicon esculentum. Des expériences de choix ont permis de déterminer ses préférences pour les différentes catégories d'oeufs. Le choix de l'oophage est influencé par l'espèce de Leptinotarsa sur laquelle il a été élevé. E. puttleri élevé sur oeufs de L. decemlineata récoltés sur un mélange des 3 solanées, et mis en présence d'un choix de groupes d'oeufs provenant de ces 3 plantes, a parasité préférentiellement les oeufs récoltés sur S. t. tuberosum; élevé à partir d'oeufs de L. texana récoltés sur S. elaeagnifolium, et mis en présence d'un mélange de pontes des 3 solanées, il a préféré parasiter les oeufs provenant de L. esculentum. Son choix a été influencé aussi par la plante qui l'avait conditionné. E. puttleri, élevé à partir de L. texana et mis en expérience sur des pontes provenant de S. t. tuberosum a préféré pondre sur les oeufs provenant de cette solanée. De même, élevé à partir d'oeufs de L. decemlineata pondus sur S. t. tuberosum, il a préféré pondre sur les oeufs provenant de cette plante. Cependant, les oophages élevés à partir d'oeufs de L. decemlineata provenant de L. esculentum, et mis en présence de pontes provenant de L. esculentum, n'ont pas préféré significativement ces oeufs au détriment de ceux provenant des autres solanées, bien que les pontes récoltées sur L. esculentum aient été parasitées plus fréquemment quand 2 des 3 pontes (une de chaque plante) avaient été sondées ou parasitées. Le parasitisme à l'intérieur d'une ponte (nombre d'oeufs de la ponte parasités, sondés, etc) ne varie pas significativement avec la solanée d'origine. La discussion a porté sur les conséquences de ces observations sur l'écologie de l'oophage et sur la lutte biologique contre L. decemlineata.

     

Dynamic effects of parasitism by an endoparasitoid wasp on the development of two host species: implications for host quality and parasitoid fitness

1. The study reported here examined growth and developmental interactions between the gregarious larval koinobiont endoparasitoid Cotesia glomerata (Hymenoptera: Braconidae) and two of its hosts that vary considerably in growth potential: Pieris rapae and the larger P. brassicae (Lepidoptera: Pieridae). At pupation, healthy larvae of P. brassicae are over twice as large, in terms of fresh body mass, as those of P. rapae. 2. Clutch size of C. glomerata was manipulated artificially, and the relationship between parasitoid burden and the maximum weight of the parasitised host (= host–parasitoid complex) was measured. In both hosts, the maximum complex weight was correlated positively with parasitoid burden. Compared with unparasitised hosts, however, the growth of P. rapae was increased at significantly lower parasitoid burdens than in P. brassicae. Emerging wasp size was correlated negatively with parasitoid burden in both host species, whereas development time was less affected. 3. After larval parasitoid egress, the weight of the host carcass increased slightly, but not significantly, with parasitoid burden, although there was a strong correlation between the proportion of host mass consumed by C. glomerata larvae during development and parasitoid burden. 4. Clutch size was generally correlated positively with instar parasitised in both hosts, and greater in P. brassicae than in P. rapae. Sex ratios were much more female biased in L1 and L2 P. rapae than in all other host classes. Adult parasitoid size was correlated inversely with host instar at parasitism, and wasps emerging from P. brassicae were larger, and completed development faster, than conspecifics emerging from P. rapae. 5. The data reveal that parasitism by C. glomerata has profound species‐specific effects on the growth of both host species. Consequently, optimality models in which host quality is often based on host size at parasitism or unparasitised growth potential may have little utility in describing the development of gregarious koinobiont endoparasitoids. The results of this investigation are discussed in relation to the potential effectiveness of gregarious koinobionts in biological control programmes.     

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

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

Factors determining the functional response of the parasitoid Venturia canescens

A series of experiments were undertaken to determine the factors responsible for the maximum number of hosts attacked by the parasitoid Venturia canescens (Gravenhorst) (Hymenoptera:Ichneumonidae) in the course of functional response experiments. Egg limitation is not a constraint on the maximum number of parasitoid progeny produced per day. Rather, the constraints arise from the requirement that eggs must be moved from the ovarioles to the oviducts and the rate at which this occurs, together with the existence of a positive relationship between the rate of parasitoid searching behaviour and the number of eggs in the oviducts.

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Résumé Le parasitoïde V. canescens produit 251±45 descendants au cours de sa vie. Les adultes ont à l'émergence 23,7±1,1 oeufs dans leurs oviductes ou descendant des ovarioles vers les oviductes à la vitesse de 1,78±0,06 oeuf/heure. II y avait une corrélation linéaire positive entre le temps dépensé par l'ichneumonide à sonder le substrat pour découvrir la larve d'un hôte et le nombre d'oeufs présents dans les oviductes. L'effectif d'oeufs pondus était proportionnel au temps passé à sonder. Le temps nécessaire à la maturation des oeufs et la relation entre le comportement de prospection et le nombre d'oeufs disponibles contribuent à limiter la vitesse de production de descendants au cours des expériences de réactions fonctionnelles. Ces facteurs ne limitent vraisemblablement pas le succès de la reproduction dans les conditions naturelles, à moins que le taux de rencontre des hôtes ne dépasse la vitesse de maturation des oeufs.

     

The mechanism by which the parasitoid wasp Trichogramma minutum responds to host clusters

Optimal strategies for utilizing a resource require the ability to assess the quantity available. To allocate its progeny appropriately, the parasitic wasp Trichogramma minutum Riley (Hymenoptera: Chalcidoidea: Trichogrammatidae) must respond not only to the size but also to the number of its insect egg hosts which are locally available.By allowing the wasps to oviposit into different clustered arrangements of hosts, it is shown that progeny allocation per host depends on the host number and position of contacting neighbours. In particular, hosts with more neighbours, and thus reduced exposed surface area, are allocated fewer progeny. It is argued that exposed surface area may be the cue used by the wasp to adjust its progeny allocation to the number of local hosts. This discrimination occurs in the absence of superparasitism. A simple model is described which accounts for previously reported counting responses.Observation of examination paths on glass bead models arranged in clusters showed that the frequency and number of edge turns change significantly with the number of neighbours. Changes in these parameters could be used to mediate the wasp's response to the number of local neighbours.The significance of host clustering effects in the use and rearing of Trichogramma for biological control is discussed briefly.

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Der mechanismus, mit dem Trichogramma minutum auf gruppen des wirtes reagiert

Zusammenfassung Optimale Strategien zur Nützung einer Reserve erfordern die Fähigkeit, die verfügbare Menge zu messen. Um die Nachkommen angemessen zu verteilen, muss die parasitische Wespe Trichogramma minutum Riley (Hymenoptera: Chalcidoidea: Trichogrammatidae) sowohl zur Größe, als auch zur Anzahl von Insekten Wirtseiern reagieren können.Die Wespen legten ihre Eier in verschiedene Gruppierungen von Wirtseiern. Es wurde gezeigt, dass die Anzahl der zugewiesenen Nachkommen von der Menge angrenzender Nachbarn und deren räumlicher Verteilung abhängt. Insbesondere werden Wirten mit mehreren Nachbarn, und demzufolge kleinerer freier Oberfläche, weniger Nachkommen zugewiesen. Auf Grund dieser Beobachtung wird die freie Oberfläche als Schlüssel für die Bestimmung der Menge von vorhandenen Wirten vorgeschlagen. Ein einfaches Modell zur Erklärung des vorher beschriebenen Zählverhaltens wird beschrieben. Es wurden Experimente mit Glaskugeln gemacht, die in unterschiedlichen Gruppen angeordnet waren. Es wurde gezeigt, dass einige Parameter des Weges bei der Wirtsuntersuchung (u.a. Häufigkeit und Anzahl der Richtungsänderungen) direkt von der Anzahl der Nachbarn abhängen. Änderungen in diesen Parametern könnten der Reaktion auf die Anzahl angrenzenden Nachbarn zugrunde liegen.Die Bedeutung des Einflusses von Wirtsgruppierung auf die Zucht und die Anwendung der Trichogramma bei der Schädlingsbekämpfung wird kurz diskutiert.

     

Changes in host acceptance and host suitability as an effect of rearing Trichogramma maidis on a factitious host

Host acceptance and suitability of Ostrinia nubilalis Hbn. for Trichogramma maidis Pint. et Voeg. (Hymenoptera: Trichogrammatidae) were studied, after rearing the parasitoids on O. nubilalis or Ephestia kuehniella Oliv. (Lepidoptera: Pyralidae) for various numbers of generations. In cages, containing two maize plants with in total 30 egg masses of the natural host (O. nubilalis), wasps continuously reared on either E. kuehniella or O. nubilalis found egg masses in equal proportions. The former however, did not parasitize the eggs successfully, due to either low degree of host acceptance or insufficient host suitability or both. Subsequently, host acceptance behaviour of individual females from different strains was studied by means of direct observations after releasing them into a small arena containing eggs of O. nubilalis or E. kuehniella. Wasps continuously reared on O. nubilalis or 1–3 generations on E. kuehniella accepted egg masses of the natural host better than wasps continuously reared on E. kuehniella. Acceptance of O. nubilalis by the wasps gradually decreased with increasing numbers of generations reared on E. kuehniella. By rearing T. maidis, previously cultured on E. kuehniella, for five generations on O. nubilalis, acceptance of egg masses of this host species did not change. Host suitability of O. nubilalis appears to correspond with host acceptance of the strains. Contrary to the observations on O. nubilalis, acceptance and suitability of E. kuehniella were not influenced by the host on which T. maidis was reared. Results of this study show that host acceptance and suitability of the natural host are important quality factors for T. maidis when mass produced on a factitious host.

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Zusammenfassung Wirtsakzeptanz und Wirtseignung von Ostrinia nubilalis Hbn. für den Eiparasitoiden, Trichogramma maidis Pint. et. Voeg., wurden untersucht nach der Zucht auf dem natürlichen Wirt, O. nubilalis, oder auf dem Ersatzwirt, der Mehlmotte, Ephestia kühniella Zell.. Versuche mit Käfigen, in denen je zwei Maispflanzen mit insgesamt 30 Eigelegen von O. nubilalis und 40 Weibchen von T. maidis eingesperrt waren, zeigten, dass Weibchen, die seit über 150 Generationen auf O. nubilalis bzw. E. kühniella gezüchtet worden waren, die Eigelege gleich gut fanden. Während die auf O. nubilalis gezüchteten Parasitoide die Eigelege zu 80% parasitierten, war die Parasitierungsrate der auf E. kühniella gezüchteten Parasitoide gleich Null. Die Ursachen dieses Phänomens wurden in weiteren Versuchen durch direkte Beobachtungen über Wirtsakzeptanz und Wirtseignung untersucht. Einzelne Weibchen von Stämmen, die nur auf O. nubilalis oder während einer zunehmenden Generationenzahl auf E. kühniella gezüchtet worden waren, untersuchte man während 15 Minuten unter der Stereoskoplupe in einer kleinen Arena, die ein Eigelege des Maiszünslers oder Eier der Mehlmotte enthielt. Dabei zeigte sich, dass Wespen, die ausschliesslich auf Maiszünslereiern oder nur 1–3 Generationen auf Mehlmotteneiern gezüchtet worden waren, die Eigelege von O. nubilalis besser akzeptierten als solche, die seit vielen Generationen auf E. kühniella vermehrt worden waren. Die Wirtsakzeptanz verschlechterte sich mit zunehmender Generationenzahl auf E. kühniella. Wenn T. maidis nach 23 Generationen auf E. kühniella wieder während 5 Generationen auf O. nubilalis gezüchtet wurde, verbesserte sich die Akzeptanz der Eigelege von O. nubilalis nicht. Die Wirtseignung der Eigelege von O. nubilalis, stimmt überein mit der Wirtsakzeptanz der verschiedenen Stämme. Dies ergaben Versuche mit 15 Minuten Direktbeobachtung und bei 24 Stunden Expositionszeit. Im Gegensatz zu den Beobachtungen auf Eigelegen von O. nubilalis wurde die Akzeptanz und Eignung der Eier von E. kühniella durch die Zucht auf verschiedenen Wirten nicht beeinflusst. Das Ergebnis dieser Untersuchung zeigt, dass Akzeptanz und Eignung des natürlichen Wirtes wichtige Qualitätsfaktoren für T. maidis in der Massenzucht auf Ersatzwirten sind.

     

The influence of female age and host size on the sex ratio of the parasitoid Opius concolor

The sex ratio of the progeny of single females parasitizing large hosts favoured the females (sex ratio=0.26); but on small hosts favoured the males (0.73). No differences in mortality of the sexes were detected. The sex ratio was independent of female age when large hosts were used. The percentage of males observed in the progeny of the first day of female oviposition was significantly greater than the mean, irrespective of the age at which female oviposition began. When females were exposed to small hosts, a greater percentage of females was observed in the progeny from the last days of oviposition.

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Résumé L'influence de la taille de l'hôteret de l'âge de la femelle sur le taux sexuel de la descendance a été étudiée sur le parasitoïde Opius concolor Szépl.Le taux sexuel de la descendance des femelles isolées est favorable aux femelles (t.s.=0,26) quand elles ont à leur disposition des hôtes de grande taille, tandis qu'avec des hôtes petits le taux sexuel est favorable aux mâles (t.s.=0.73). On n'a pas détecté de mortalité différentielle des sexes.Les pourcentage de mâles obtenu le premier jour de ponte des femelles sur les hôtes de grande taille est significativement différent de la moyenne, indépendamment de l'âge de la femelle à ce moment. Cependant, sur des hôtes petits, bien que restant favorable aux mâles dans l'ensemble, une plus grande proportion de femelles à partir des premiers jours de ponte a pu être observée.