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.

     

Host volume measurement by the parasitoid wasp Trichogramma minutum: The roles of curvature and surface area

The number of eggs laid into its insect-egg host by Trichogramma minutum Riley (Hymenoptera: Chalcidoidea: Trichogrammatidae) depends on the volume of its lepidopterous host. The wasps allocated fewer progeny into Manduca sexta (L.) (Sphingidae) egg hosts partially embedded in the substrate than into fully exposed hosts, both in daylight and in total darkness. Since the hosts are of identical diameter and surface chemistry, the mechanism of volume determination is neither chemical nor visual, but essentially mechanosensory, based on accessible surface area. Before oviposition, the wasp examines the host surface by walking over it, turning at the edge, and drumming with its antennae. The number of edge turns (N_t) made by wasps when they examined glass beads presented as model hosts was inversely dependent on surface area, but independent of diameter. Conversely, the examination time (T_e) was directly dependent on diameter, but independent of surface area.A mechanism is proposed for host volume measurement. While the wasp examines a potential host, the relative curvature (and hence diameter) is measured, possibly by antennal angle. If this measure is within an acceptable range, the wasp initiates its drumming walk. The measure is also used to set T_c and to establish an estimate of a maximum possible volume (and hence progeny number) to the host object. With T_c fixed, the value for N_t depends on accessible surface area, the wasp turning more often on restricted surfaces. N_t allows the wasp to correct the estimate of host volume such that an appropriate number of eggs is laid in non-spherical hosts. The use of curvature to set T_e increases the accuracy of the wasp's measure of N_t, while also improving its coverage of the surface. The use of curvature and surface area allows Trichogramma to optimize its progeny allocation for a variety of host sizes and shapes.

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Die bestimmung des wirtsvolumen durch die parasitische wespe Trichogramma minutum: Die rolle von krümmung und oberfläche

Zusammenfassung Bei dem Insekten-Eiparasiten Trichogramma minutum (Hymenoptera: Trichogrammatidae) hängt die Anzahl der abgelegten Eier vom Volumen der Wirtseier ab. Es wurden Untersuchungen zur Bestimmung der sensorischen Grundlagen dieses Verhaltens unternommen. Die Wespe weist sowohl im Tageslicht, als auch in totaler Dunkelheit jenen Manduca sexta (L.) (Lepidoptera: Sphingidae) Eiwirten (Diameter c. 1.3 mm) weniger eigene Eier zu, die teils (30%) in Substrat eingebettet sind, als solchen, die ganz herausragen. Da die Wirte einen identischen Radius und gleichen Oberflächengeruch haben, kann der Mechanismus der Volumenbestimmung weder chemisch noch optisch sein, sondern ist dem Wesen nach mechanosensorisch, und von der zugänglichen Oberfläche abhängig. Vor de Eiablage untersucht die Wespe die Wirtsoberfläche, indem sie darüber läuft, an den Ecken umkehrt und mit ihren Antennen trommelt. In Experimenten mit Glaskugelmodellen war die Anzahl der von der Wespe gemachten Wendungen an den Ecken (N_t) umgekehrt proportional zur Oberfläche, aber unabhängig vom Radius. Im Gegensatz dazu, war die Untersuchungszeit (T_e) direkt proportional zum Radius, aber unabhängig von der Oberfläche.Ein Mechanismus für die Volumenmessung wird vorgeschlagen. Gleich beim Besteigen des potentialen Wirtes wird die relative Kurve (und infolgedessen der Radius), wahrscheinlich durch den Antennenwinkel gemessen. Wenn diese Messung innerhalb einer bestimmten Spanne liegt, beginnt die Wespe ihren Trommelgang. Die Messung wird auch dazu benutzt, ein maximales Volumen (und folglich die Anzahl der Nachkommen) des Wirtes und eine maximale Untersuchungzeit (T_e) zu erzielen. Wird T_e konstantgehalten, hängt die Anzahl der Wendungen (N_t) von der zugänglichen Oberfläche ab, wobei die Wespe öfter bei kleineren Oberflächen wendet. Die Anzahl der Wendungen gestattet der Wespe, die ursprüngliche Einschätzung des Wirtsvolumens so zu verändern, daß eine angemessene Anzahl von Eiern in nicht kugelige Wirte gelegt wird. Die Anwendung von kurven-und oberflächenabhängigen Messungen ermöglicht eine optimale Zuweisung von Nachkommen für eine Anzahl von unterschiedlichen Wirtsgrössen und-formen. Die Oberflächenmessung gestattet eine bessere Volumenbestimmung von nicht-kugeligen Objekten. Die gezeigte Abhängigkeit von der Oberfläche ist wahrscheinlich auch eine Anpassung an Wirte, die ihre Eier in Gruppen ablegen. Unter solchen Bedingungen von grosser Wirtsdichte ist es günstiger für die Wespe, weniger Eier in jeden Wirt der Gruppe zu legen, weil diese Verteilung zu grösseren, fruchtbareren Nachkommen führt.

     

Flight initiation in the egg parasitoid Trichogramma minutum: Effects of ambient temperature, mates, food, and host eggs

Emergence, preening, and flight initiation were studied in laboratory-reared Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae). Male parasitoids emerged first and flew before females. When both sexes were present in flight cylinders, female parasitoids flew before males. Flight propensity in males was negatively related to the number of emerging females, while flight propensity in females was independent of the number of males present. Ambient temperature significantly affected the propensity and timing of flight; between 70–80% of the parasitoids flew at 25 and 30 °C while less than 4%, mostly males, flew at 20 °C. No flights were observed at 15 °C. The presence of fresh host eggs caused a reduction in the proportion of female parasitoids that flew and a delay in the time to flight for the females that did fly. The presence of food increased the flight propensity of female parasitoids, but did not affect the timing of flight. The relationship between flight behaviour and the efficiency of mass-reared Trichogramma is discussed in terms of its importance for inundative release programmes.     

Parasitism of non-target lepidoptera by mass released Trichogramma brassicae and its implication for the larval parasitoid Lydella thompsoni

The release of high numbers of the eggparasitoid Trichogramma brassicae Bezd.(Hym. Trichogrammatidae) to control theEuropean corn borer (ECB), Ostrinianubilalis Hb. (Lep.: Crambidae) in maize hasraised concerns about potential negativeeffects on native natural enemies. The nativelarval parasitoid Lydella thompsoniHerting (Dipt.: Tachinidae) is the mostfrequent and important ECB parasitoid insouthern Switzerland and can achieve highparasitism rates. Its first generation emergestoo early to find ECB larvae and must rely onalternative hosts living in natural habitatsclose to maize fields. Inundative releases ofT. brassicae coincide with theoviposition period of the alternative hosts ofthe tachinid. T. brassicae moving out ofrelease fields may attack and diminish thepopulation of these hosts, creating abottleneck situation for L. thompsoni inthe subsequent spring. Laboratory hostspecificity tests showed that the tachinid'stwo most abundant spring hosts Archanarageminipuncta Haworth (1809) (Lep.: Noctuidae)and Chilo phragmitellus Hübner (1805)(Lep.: Crambidae) are successfully parasitisedby T. brassicae females in no-choicesituations. Our extensive field surveys,however, showed that the two tested springhosts escape parasitism since their eggs arewell hidden or not attractive. Negativeeffects of inundative releases of T.brassicae on the native tachinid fly L. thompsoni, such as population densityreduction, displacement, or local extinction,are very unlikely.     

Host examination walk and oviposition site selection ofTrichogramma minutum: Studies on spherical hosts

The host examination behavior of the parasitoid wasp Trichogramma minutumon spherical Manduca sextahosts is described. The mean walking speed during host examination was 0.64 ± 0.03 mm s^–1and is independent of wasp size. The paths taken by the wasps are not evenly distributed over the host surface; the majority of time is spent below 45 ° latitude. The distribution of oviposition sites is also nonrandom with respect to latitude and is not influenced by phototaxic and geotaxic responses. The initial transit made by the wasps over the host surface is a straight path which frequently passes across the highest point on the host. The length and duration of the initial transit are independent of wasp body length. However, stride length and stepping rates are proportional to wasp body length, and small wasps take more steps to complete their initial transit. The roles of the examination walk in host recognition and host volume measurement by Trichogrammaare also discussed.     

Quality and environmental factors affecting Trichogramma brassicae efficiency under field conditions

Field parasitism of the egg parasitoid Trichogramma brassicae Bezdenko (Hymenoptera, Trichogrammatidae) (synonymous to T. maidis Pint. et Voegele) on Ostrinia nubilalis Hübner was compared to four single quality parameters (walking speed, fecundity on the factitious host Ephestia keuhniella Zeller, fecundity on the natural host O. nubilalis, and life span) previously measured in the laboratory and, a quality index calculated from three of the measured parameters. A single quality parameter (fecundity on E. keuhniella) and the calculated quality index showed a correlation to T. brassicae field parasitism, for different T. brassicae populations. The number of female Trichogramma released clearly influenced parasitism but not in the same proportion for all populations tested, reflecting that numbers released can only to some extent compensate for low quality in Trichogramma. Variation indegree hours above 18°C also influenced field parasitism. When incorporating degree hours to the quality index a significant correlation R²=0.56 (P=0.01) is obtained. In this work fecundity of T. brassicae on E. keuhniella can be as good an indicator for the potential field performance of T. brassicae, as the quality index which takes into account three quality parameters. Since environmental factors can obscure the potential performance of a population, i.e., one which under optimal laboratory conditions performs well, the quality index gives only a partial indication of how the released strains will perform in the field.     

Effect of host availability on reproduction and survival of the parasitoid wasp Trichogramma minutum

Abstract.

• 1 We tested the hypothesis that females of the egg parasitoid, Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), could adjust their fecundity schedule according to host availability and that there was a negative correlation between reproduction and survival in these wasps.
• 2 Newly-emerged females were provided with an unlimited or limited number of hosts in the first trial and with either unlimited, limited or zero hosts in the second trial.
• 3 When hosts were unlimited, wasps had the highest rate of reproduction in the first day, which decreased dramatically thereafter. When hosts were limited, wasps from the two trials differed in their response. In Trial I, females with limited hosts had lower first-day fecundity than, and the same subsequent-day fecundity as, those with unlimited hosts. However, in Trial II, females with limited host had a lower first-day but a higher subsequent-day fecundity than those with unlimited hosts. This indicates variation in Trichogramma's ability to shift its fecundity schedule in response to host availability.
• 4 There was a positive (rather than a negative) correlation between reproduction and survival. Wasps that oviposited (in host-unlimited treatment) had greater longevity than those that could not (in host-unavailable treatment).
• 5 The sex ratio of the progeny produced by wasps in both host-unlimited and limited treatments shifted gradually from a female to a male bias as the wasps aged.
• 6 We consider the ability of parasitoids to adjust their fecundity schedule as an adaptation to changing host resources and discuss our findings with regard to theories of life history evolution.

     

Influence of leaf surfaces on movements by the hymenopterous parasitoid Trichogramma exiguum

Walking speeds of female Trichogramma exiguum Pinto & Platner were fastest on maize and soybean (12 cm/min), intermediate on tomato (8 cm/min), and slowest on woolly mullein, Verbascum thapsus (3 cm/min). Similarly, rates of turning along the paths of walking T. exiguum were smallest on maize (median angle=0°±15°), intermediate on soybean and tomato, and greatest on V. thapsus (median angle=30°±15°). Leaf trichome density and morphology influenced walking behavior. Walking was slowed and flight initiation delayed for T. exiguum walking on Amaranthus hybridus leaves compared to either maize or filter paper. When inundative releases are conducted, the effects of plant surfaces on searching rates and arrestment should be considered in determining release rates of Trichogramma spp.

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Résumé La vitesse de marche a été déterminée en lâchant des individus sur chaque substrat végétal et en traçant leur parcours sur une plaque de verre placée à 9 mm au dessus du sujet. Les déviations angulaires de portions successives des tracés, longues de 1 mm, ont été utilisées pour mesurer les taux de changement de direction. Dans une deuxième expérience, des individus ont été lâchés au milieu d'une cercle de 40 mm de diamètre sur chaque substrat et les temps écoulés avant l'envol ou pour atteindre le bord du cercle à la marche ont servi à évaluer respectivement la propension au vol et la vitesse de déambulation.Les vitesses de marche ont été plus rapides sur maïs et soja (12 cm/min), moyennes sur tomate (8 cm/min) et les plus lentes sur Verbascum thapsus (Scrophulariaceae) (3 cm/min). De le même façon, les angles de changement de direction au cours des marches effectuées par T. exiguum ont été plus petits sur maïs, moyens sur soja et tomate, et plus grands sur V. thapsus. Chez T. exiguum marchant sur des feuilles d'Amaranthus hybridus L. (Amaranthaceae), la marche a été plus lente et l'envol plus tardif que sur maîs ou papier filtre.Lors de lâchers inondatifs, les effets des surfaces végétales sur les vitesses d'exploration et d'arrêt devraient être pris en compte pour déterminer les vitesses de lâcher des Trichogramma spp.

     

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.

     

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.     

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.

     

Effect of host diet on acceptance of eastern spruce budworm eggs byTrichogramma minutum

The acceptance behaviour of the parasitic wasp, Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), was studied on two types of egg masses of eastern spruce budworm (SBW), Choristoneura fumiferana Clemens. Host eggs were obtained from moths that had been fed balsam fir or artificial diet during their larval stages. Parasitoids were reared from both types of hosts. Host acceptance assays were conducted where females were provided with a single egg mass and with a choice between egg masses. Regardless of the parasitoid's rearing host, females parasitised hosts reared from artificial diet more often than those from balsam fir. Response was consistent in both no-choice and choice assays. Rejection of hosts tended to occur during the examining stage of behaviour, suggesting that acceptability is judged externally. If a host was accepted, the same number of eggs were laid per egg mass regardless of treatment.     

Suppression ofOstrinia nubilalis byTrichogramma nubilale in sweet corn

We conducted inundative release experiments withTrichogramma nubilale (Hymenoptera: Trichogrammatidae) to suppressOstrinia nubilalis (Lepidoptera: Pyralidae) in sweet corn (Zea mays): two experiment duringO. nubilalis first generation and three experiments during second generation. Five measurements of ear and stalk damage were used to assess.O. nubilalis control in treated and untreated plots within each experimental field. In one experiment during second generation, naturalO. nubilalis populations were sufficiently high to demonstrate that the parasitoids (three releases totaling 4.4 million parasitoids per ha) parasitized an estimated 57.4% of the placedO. nubilalis egg masses and reduced the mean number ofO. nubilalis larvae per ear by 97.4% the number of tunnels per stalk by 92.9%, and the number of larvae per stalk by 94.3% in the release plot. Ear damage in this experiment was suppressed to meet acceptable standards for use in cut-corn commercial processing. Larval mortality was apparently density independent, which implies that density-dependent larval loss would not compensate for egg parasitism byT. nubilale.     

Host-stage selection by Trybliographa rapae, a parasitoid of the cabbage root fly Delia radicum

Host-stage selection by Trybliographa rapae Westwood (Hymenoptera: Figitidae) was studied in choice and no-choice experiments in the laboratory. The parasitoid was able to reproduce in first, second, and third instars of the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae), but oviposition occurred more frequently in third instars when all three developmental stages were offered simultaneously. Oviposition in third instars increased the rate of development of offspring and their body size, but did not alter sex ratio. Results are discussed in the light of predictions made by the theory of optimal host acceptance.     

The effect of temperature on progeny allocation and short interval timing in a parasitoid wasp

Abstract The parasitoid wasp Trichogramma dendrolimi Matsumura sets the number of progeny allocated to its insect egg hosts according to the duration of its initial transit walk across the host surface. Although cooling the wasp reduces its walking speed, reduced temperature does not affect progeny allocation. By locally heating and cooling the wasps, the initial transit can be thermally uncoupled from the subsequent oviposition. Using this technique we show that the timing of initial transit duration is temperature-dependent. These findings suggest that short interval timing by Trichogramma differs physiologically from the temperature compensated clocks that have been described for other insects.     

Role of kairomones from host accessory gland secretion in host recognition by Telenomus remus and Trichogramma pretiosum, with partial characterization

Accessory gland materials from Spodoptera frugiperda (J. E. Smith) and Heliothis zea (Boddie) (Lepidoptera: Noctuidae) contained kairomones that influenced the host recognition or acceptance behavior of Telenomus remus Nixon (Hymenoptera: Scelonidae) and Trichogramma pretiosum Riley (Hymeoptera: Trichogrammatidae), respectively. Material from S. frugiperda accessory glands stimulated ovipositor probing and drilling by female T. remus but not T. pretiosum. Accessory gland material from H. zea stimulated ovipositor probing and drilling by female T. pretiosum but not T. remus. An active material for T. remus is found in the 700 K protein fraction of accessory gland material from S. frugiperda. The oviposition behavior of the parasitoids is also discussed.

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Résumé Cette étude a utilisé une technique voisine de celle employée par Strand & Vinson (1982), pour examiner les réactions de parasitoïdes à des billes de verre enduites de substances, afin de déterminer l'influence des sécrétions des glandes annexes (AGM) de Spodoptera frugiperda et Heliothis zea dans le repérage des hôtes par Telenomus remus et Trichogramma pretiosum.Les femelles de T. remus réagissaient plus aux billes enduites d'AGM de S. frugiperda qu'aux billes témois non enduites; celles de T. pretiosum réagissaient plus aux billes enduites d'AGM de H. zea qu'aux témoins. T. remus n'a pas réagi aux billes enduites d'AGM de H. zea, ni T. pretiosum à celles enduites d'AGM de S. frugiperda.Les poids moléculaires des protéines des glandes annexes de H. zea sont 1.100 K et 50 K, et ceux de S. frugiperda de 700 K et 30 K (Strand & Vinson, 1983a). T. remus a mieux réagi aux billes enduites de la fraction 700 K de S. frugiperda qu'a celles enduites de la fraction 30 K, aux deux fractions de H. zea et aux billes témoins. Les billes enduites de la fraction 1.100 K de H. zea ont provoqué aussi une réaction significative de T. remus. Les sécrétions des glandes annexes de S. frugiperda et de H. zea contiennent des kairomones qui interviennent dans le repérage des hôtes respectifs de T. remus et de T. pretiosum. Mais, il semble peu probable que ces substances soient seules responsables de la reconnaissance de l'hôte.

     

Host Preferences of Trichogramma pretiosum and the Influence of Prior Ovipositional Experience on the Parasitism of Plutella xylostella and Pseudoplusia includens Eggs

Trichogramma wasps are generalist egg parasitoids used in biological control efforts. In a multi host situation they may preferentially parasitize a non-target host species to the detriment of the control program. Plutella xylostella Linnaeus (Lepidoptera: Plutellidae) is a very serious pest of cabbage, but is only one in a number of species in the ‘cabbageworm’ complex. We investigated the host preferences of Trichogramma pretiosum Riley (Hymenoptera: Chalcidoidea) when offered the eggs of Plutella xylostella and the eggs of Pseudoplusia includens Hübner (Lepidoptera: Noctuidae), another species in the ‘cabbageworm’ complex. Trichogramma pretiosum reared on the eggs of the factitious host Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) parasitized both Plutella xylostella and Pseudoplusia includens eggs under laboratory conditions. For both choice and no-choice experiments, T. pretiosum parasitized significantly more P. xylostella eggs than P. includens eggs. Prior ovipositional experience with one or other of the two host species had no effect on the subsequent parasitism levels of the two host species. The preference for P. xylostella eggs was also not affected by this prior ovipositional experience.     

Influence of epicuticular waxes from white spruce and balsam fir on feeding behavior of the eastern spruce budworm

Fourth-instar larvae of the eastern spruce bud-worm Choristoneura fumiferanaClem. (Lepidoptera: Tortricidae) were shown in two-choice feeding tests to respond differently to epicuticular waxes from different white spruce and balsam fir trees. Larvae also showed different preferences for various wax fractions obtained by separation on silicic acid.Deceased.     

Assessment ofTrichogramma species for biological control of forest lepidopteran defoliators

In a laboratory study, we determined the potential of threeTrichogramma (Hymenoptera: Trichogrammatidae) species,T. brassicae Bezdenko,T. minutum Riley andT. nr.sibiricum Sorokina, for biological control against six species of forest lepidopteran pests, black army cutworm, hemlock looper, eastern spruce budworm, western spruce budworm, white-marked tussock moth, and gypsy moth. Females of each parasitoid species were offered eggs from each of the six host species. Parasitization and the effect of the host species on the emerging progeny were examined and recorded.Trichogramma minutum had the broadest host range and successfully parasitized four host species out of the six offered.Trichogramma nr.sibiricum had the narrowest host range and parasitized only two species of hosts. Of the six host species, black army cutworm was the most preferred by all threeTrichogramma species; white-marked tussock moth and gypsy moth were not parasitized by any parasitoids. There was a positive correlation between the size of female offspring and their corresponding egg complement in all three parasitoid species. The developmental time of parasitoids from egg to adult was influenced by both the parasitoid and host species. Our results suggest thatT. minutum has the greatest potential for biological control against various forest lepidopteran pests and that the black army cutworm may be the best target candidate for further study.