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.

     

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.

     

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.     

Influence of environmental conditions and parasitoid quality on field performance of Trichogramma minutum

The effects of weather conditions and two parasitoid quality attributes, realized fecundity and host acceptance, were assessed on the field efficacy of mass-released Trichogramma minutum. Temperature was the most important single variable, explaining up to 75% of the variation in field parasitism. There were significant positive relationships between both the sum of the maximum temperatures and the number of degree-hours above a 15 °C threshold, accumulated in the three days following the release, and parasitism in the field. There was a significant negative relationship between the mean relative humidity and the odds of parasitism in the field. Quality parameters based on parasitoid biology were not effective for predicting field efficacy if poor weather conditions persisted after a release. If weather conditions were good (i.e. accumulated maximum temperatures above 62 °C, in the 3 days following the release), then parameters such as release rate and fecundity in the lab were useful predictors of field performance. There was no relationship between host acceptance measured in the lab and field parasitism. Given the importance of field temperatures for field performance, selection for cold tolerance of T. minutum would be desirable.     

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.     

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.

     

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.

     

The effect of host nutrition on growth and development of the parasitoid waspVenturia canescens

We investigated the effect of host (Plodia interpunctella; Lepidoptera: Pyralidae) nutritional status on development of the solitary endoparasitoid,Venturia canescens (Hymenoptera: Ichneumonidae). Parasitoids from 3rd (L3) instars reared on a deficient diet during early parasitism took longer to develop and suffered higher mortality than those reared from hosts fedad libitum although there was not a significant difference in the size of eclosing wasps from the two groups. L5 hosts reared at high density produced smaller parasitoids, which developed more rapidly than those reared from hosts from low density containers, although mortality was higher in the latter. In a separate experiment we starved groups of 10–20 hosts (parasitized as L3) daily beginning on the 4th day after parasitism, to determine the host developmental stage required for successful parasitoid development to eclosion. Parasitoid survivorship increased with length of host access to food, while the egg-to-adult parasitoid development time increased throughout the experiment. Parasitoid size decreased with increasing periods of host starvation. The successful emergence ofVenturia depends uponPlodia reaching the size normally attained in the mid-5th instar, or 50–70% of the mass of healthy late 5th instars. Our results show that when earlier instars are parasitized, host growth is essential for successful parasitoid development to eclosion. Furthermore, they suggest that, for many koinobionts, host suitability may be greatly influenced by feeding rate and food quality.     

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.     

The origin of a selfish B chromosome triggering paternal sex ratio in the parasitoid wasp Trichogramma kaykai

This study uses molecular and cytogenetic methods to determine the origin of a B chromosome in some males of the wasp Trichogramma kaykai. This so-called paternal sex ratio (PSR) chromosome transmits only through sperm and shortly after fertilization triggers degeneration of the paternal genome, while keeping itself intact. The resulting embryos develop into haploid B-chromosome-carrying males. Another PSR chromosome with a very similar mode of action is found in the distantly related wasp Nasonia vitripennis and its origin was traced by transposon similarity to the genus Trichomalopsis, which is closely related to Nasonia. To determine whether both PSR chromosomes have a similar origin we aimed to reveal the origin of the Trichogramma PSR chromosome. Using fluorescent in situ hybridization, we discovered a major satellite repeat on the PSR chromosome, the 45S ribosomal DNA. Analysis of the internal transcribed spacer 2 (ITS2) of this repeat showed the presence of multiple ITS2 sequences on the PSR chromosome resembling either the ITS2 of T. oleae or of T. kaykai. We therefore conclude that the Trichogramma PSR chromosome originates from T. oleae or a T. oleae-like species. Our results are consistent with different origins for the PSR chromosomes in Trichogramma and Nasonia.     

Overwintering of the egg parasitoid Trichogramma brassicae in Northern Switzerland

As part of a general risk assessment study weinvestigated the overwintering abilities ofTrichogramma brassicae Bezd. (Hym.: Trichogrammatidae) in Northern Switzerland. Eggs of six host species parasitized by T. brassicae were exposed under outdoor conditions every two weeksbetween 26 September and 7 November 1998. Wefound that T. brassicae is able tooverwinter successfully on six lepidopteranspecies in the families Tortricidae, Noctuidae,Plutellidae, Pyralidae and Crambidae. Between75 and 100% emergence was observed in thefollowing spring for all of the six tested hostspecies exposed on 26 September. On laterexposure dates, spring emergence decreasedsignificantly and no development of T. brassicae offspring occurred from host eggsparasitized on 7 November.Emergence of T. brassicae from eggs ofEphestia kuehniella Zeller parasitized atweekly intervals in autumn 1999 occurred in thesame year until the end of October if the eggswere parasitized by 10 September. All offspringfrom eggs that were parasitized from 17September onwards went into diapause. Springemergence of T. brassicae adults wasobserved between end of April and beginning ofMay both in 1999 and 2000. Fecundity of T.brassicae females that overwintered on E.kuehniella was not significantly different fromthe fecundity of control females that werereared without diapause under optimalconditions at 25 °C. Our resultsdemonstrate that the egg parasitoid T.brassicae is able to overwinter successfullyin Northern Switzerland and has the potentialto establish in Switzerland. As a result,potential non-target effects are not locallyrestricted but may occur on a largegeographical scale.     

A comparison between the amino acid composition of an egg parasitoid wasp and some of its hosts

Amino acid compositions of the eggs of five lepidopteran hosts for Trichogramma minutum were compared with each other and with a non-host species, Rhodnius prolixus, in which T. minutum oviposits but does not develop. Host eggs are quite homogeneous, particularly when compared according to groupings of potentially interconvertible amino acids. Combined mole percent values for glycine, serine and alanine were higher in hosts (27.5–29.2 mole%) than in R. prolixus eggs (21.5 mole%), in bovine serum albumin (14.9%), which has been used as a protein source in artificial diets for T. minutum, or in many of the mixtures used in published diets for this species. Since these three amino acids make up 26.3 mole% of the adult amino acid content of T. minutum, their deficiency in diets could require metabolic compensation detrimental to development.Adult T. minutum arising from eggs of Manduca sexta, Choristoneura fumiferana, and Sitotroga cerealella are similar in amino acid composition to each other and, in general, to their hosts. Variability appears greater in hosts than in adult wasp composition, suggesting some interconversion of host amino acids to accommodate inflexible nutritional requirements of T. minutum.In the three host species tested, free amino acids constituted 15.8–19.3% by weight of the amino acid in egg contents. In M. sexta eggs, glycine, serine and alanine together make up 28.4% by weight of the total free amino acid, a much higher proportion than in many published diets. The four free amino acids (isoleucine, leucine, phenylalanine and histidine) reported to be oviposition stimulants in experiments on encapsulated diets are present in sufficient concentrations to induce oviposition in the host species tested and in R. prolixus. S. cerealella egg contents having approximately 1.8 g amino acid, yield one or rarely two adult T. minutum (1g amino acid/insect). In contrast, M. sexta eggs with 94 g amino acid each yield an average of 10–12 adults (8.2g amino acid/insect). This suggests that small hosts are allocated few eggs which can only develop into small adults because of nutrient supply (parasitoid size in metabolically restricted), whereas much larger hosts are allocated proportionately fewer eggs than the former resulting in larger, and presumably more viable and fecund, adults (parasitoid size is established behaviourally).     

Influence of host developmental state on host suitability and reproductive biology of the parasitoid Eucelatoria bryani

Eucelatoria bryani Sabrosky (Diptera: Tachinidae) successfully parasitized 2nd through 5th instars and prepupal Heliothis zea (Boddie) (Lepidoptera: Noctuidae) in the laboratory. Percent successful parasitism (that which resulted in the production of adult parasitoid progeny) increased with host developmental state, reaching 95% in 5th instars, but fell to 63% in prepupae. While 25% of the exposed prepupae metamorphosed to pupae, none developed into adults. E. bryani maggots emerged from only 5% of H. zea pupae. The mean number of fly puparia and adults produced per successfully parasitized larva increased with host stadium, reaching 14.5±1.33 (SEM) and 10.6±1.02, respectively, in prepupae. The sex ratio of adult parasitoid progeny per host larval stadium was variable, and did not appear to follow a pattern. A formula for calculating a host favorability index is presented. This index allows a direct comparison of the overall impact and reproductive potential of E. bryani attacking hosts of varying developmental states.

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Résumé Les influences de l'état de développement de l'hôte sur le taux de parasitisme, la production de descendants, l'émergence des adultes et le taux sexuel de la techinaire E. bryani Sabrosky ont été examinés au laboratoire. Un parasitisme réussi,-qui entraîne la formation d'un parasitoïde adulte-, s'effectue dans les chenilles de stades 2 à 5 et dans les prénymphes d'Heliothis zea Boddie. II s'agit d'un élargissement considérable de la gamme connue des tailles convenables d'hôtes. Bien que le parasitisme n'ait jamais réussi avec les chenilles de premier stade, une très forte mortalité, 93% a été observée quand ces petites chenilles ont été exposées à la tachinaire. Pour permettre l'introduction par le larvipositeur des asticots dans la chenille, la mouche perfore la cuticule de l'hôte avec un sclérite modifié; ainsi, une grande partie de la mortalité est provoquée vraisemblablement par le traumatisme dû au parasitoïde. Le succès du parasitisme a augmenté en fonction du développement de l'hôte du second stade (30%) au cinquième stade (95%), pour tomber à 63% dans les prénymphes. Cependant, aucune des prénymphes exposées aux mouches n'a été capable de donner des papillons.Les production moyennes de pupes et de mouches par chenille effectivement parasitée ont augmenté avec le stade de développement de l'hôte pouratteindre dans les prénymphes respectivement 14,5±1,33 et 10,6±1,02. Le taux sexuel des mouches obtenues a été très variable pour les différents stades de développement des chenilles, sans qu'aucune corrélation ait pu être mise en évidence. Une formule permettant de calculer un indice d'adéquation de l'hôte est proposé. Cet indice permet une comparaison directe des impacts globaux et des potentiels reproducteurs des femelles de E. bryani attaquant des hôtes à différents stades.

     

Host discrimination and evidence for a host marking pheromone in the almond seed wasp, Eurytoma amygdali

Laboratory experiments and observations on the oviposition behaviour of the almond seed wasp Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) revealed that the females of this species deposit a host-marking pheromone, immediately after an oviposition, by dragging the tip of their abdomen on the fruit surface. This pheromone enables them to discriminate between the infested and uninfested fruit and to select for oviposition the latter. Its primary function is apparently the prevention of repeated ovipositions in already infested fruit, thus contributing to the optimal utilisation of the available resources for larval development. The responses of individual females to different treatments of almonds, in a series of two-choice tests, revealed that the pheromone can be perceived by the females on direct contact and, when at high concentrations, also olfactorily from a short distance. The pheromone was present inside the abdomen and thorax of females but not of males, and, although water soluble, could not be entirely removed from heavily infested almonds when rinsed with water. Direct observations revealed that after an average of 3.7 successive visits to pheromone-bearing almonds, females were induced to walk or, most often, fly away from the experimental set-up. This suggests that the pheromone may also contribute to the dispersion of the wasps.

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Résumé Des expériences de laboratoire et des observations concernant le comportement de ponte de Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) ont montré que, immédiatement après la ponte, les femelles déposent une phéromone de marquage de l'hôte en trainant le bout de leur abdomen sur la surface de l'amande. Cette phéromone les rends capable de distinguer les fruits infectés des non-infectés dt de sélectionner pour la ponte les derniers. Apparemment, la fonction principale de cette phéromone est la prévention de la répétition des ovipositions dans les fruits déjà infectés et la répartition uniforme des oeufs dans les amandes, contribuant ainsi à la meilleure utilisation des ressources disponsibles pour le développement des larves. Des expériences de deux choix entre des fruits de différents traitements ont montré que la phéromone pouvait être perçue par les femelles par le direct contact et, quand elle était à hautes concentrations, par olfaction d'une courte distance. La phéromone était présente dans l'abdomen et dans le thorax des femelles, et bien qu'elle soit soluble à l'eau, elle ne pouvait pas s'éloigner entièrement par lavage des amandes sérieusement infectées à l'eau. Les observations ont démontré qu'après un numéro de 3.7 visites successives sur des amandes portant de la phéromone les femelles s'éloignaient du lieu de ponte en marchant ou, le plus souvant, en s'envolant, ce qui suggère que la phéromone contribue à la dispersion des femelles.

     

Influence of host distribution on foraging behaviour in the hyperparasitoid wasp, Dendrocerus carpenteri

The foraging behaviour of Dendrocerus carpenteriCurtis (Hymenoptera: Megaspilidae), an ectophagous hyperparasitoid of aphidiine wasps inside mummified aphids, was examined in the laboratory with an experimental system consisting of broad bean, Vicia fabaL, the pea aphid, Acyrthosiphon pisumHarris, and a primary parasitoid, Ephedrus californicusBaker. Pea aphids parasitised by E. californicusoften disperse from their feeding sites (or off host plants) before dying and mummifying. Response of female hyperparasitoids to host distribution was evaluated at two spatial scales. At the first scale, behaviour of hyperparasitoids was examined on individual plants with different densities of hosts. At the second scale, habitat complexity and host location were manipulated in large foraging cages containing several plants. I show that patterns of density-dependent hyperparasitism can result from the foraging behaviour of D. carpenteri. However, dispersal of parasitised aphids may not reduce the incidence of hyperparasitism if hyperparasitoids systematically search the habitat.     

Isoenzyme patterns and biology ofTrichogramma minutum as influenced by rearing temperature and host

Females ofTrichogramma minutum Riley reared at temperatures below 17°C had lower percent emergence, lower fecundity on the first day of emergence, and a reduced rate of population increase (r_i) than those reared at temperatures above 17°C. Parasitoids were more fecund and lived longer when raised onChoristoneura fumiferana (Clemens) compared toEphestia kuehniella (Zeller). The parasitoids performed better when parasitizing the same species of host on which they had been reared. Isoenzyme patterns were not affected whenT. minutum was reared at 13 or 27°C but additional bands appeared when parasitoids were reared onSitotroga cerealella (Olivier) compared toC. fumiferana. No correlation could be made between biological and biochemical changes inT. minutum. Isoenzymes have the potential to assess the quality of parasitoids during mass production.

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Einfluss von Zuchttemperatur und Wirt auf Isoenzymmuster und Biologie vonTrichogramma minutum Riley

Zusammenfassung Weibchen vonTrichogramma minutum, welche bei 17°C gezüchtet wurden, hatten eine geringere Schlüpfrate, eine geringere Fruchtbarkeit am ersten Tag nach dem Schlüpfen sowie ein reduziertes Populationswachstum (r_i) im Vergleich zu Tieren welche sich bei Temperaturen über 17°C entwickelt hatten. Wurden die Parasitoiden aufChoristoneura fumiferana gezogen, so war ihre Fruchtbarkeitsrate wie auch Lebensdauer grösser als wennEphestia kuehniella als Wirt diente. Die parasitische Fähigkeit vonT. minutum war immer auf derjenigen Wirtsart am grössten auf der die Schlupfwespen vorher gezogen wurden. Aufzuchttemperaturen von 13°C oder 27°C hatten keinen Einfluss auf den Isoenzymbändern wenn die Tiere statt aufC. fumiferana aufSitotroga cerealella gezogen wurden. Es konnte aber kein Zusammenhang zwischen den biologischen und biochemischen Veränderungen gefunden werden. Hingegen besteht die Möglichkeit während der Massenproduktion die Qualität der Parasitoiden anhand von Isoenzymprofilen zu überwachen.

     

Interactions between host attributes and wasp size: a laboratory evaluation of Trichogramma platneri as an augmentative biological control agent for two avocado pests

We compared egg parasitization of two sporadic pests of avocado, Persea americana Miller, by Trichogramma platneri Nagarkatti reared on Sitotroga cerealella (Oliver) (=smaller parasitoids) with those reared on Trichoplusia ni (Hübner) (= larger parasitoids) in the laboratory. Large T. platneri females invested less time than smaller parasitoids in obtaining offspring from eggs of either Amorbia cuneana Walsingham or Sabulodes aegrotata (Gueneé). Furthermore, large T. platneri invested substantially more time (ca. 3-fold) in obtaining offspring from S. aegrotata than A. cuneana eggs. These investment times positively correlated with percentage egg-load retained by a female after it had ceased exploiting a host species. This occurred even though the offspring produced by a parasitoid from either host species manifested the same size and fecundity (= quality). Host species utilization in the laboratory mirrored that observed in the avocado groves of southern California following augmentative release of commercially produced T. platneri (Oatman & Platner, 1985). Explanations for these differences in host utilization are discussed along with their implications for augmentative biological control.     

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.

     

Host exploitation by parasitoids: local variations in foraging behaviour of females among populations of Trichogramma species

Using artificial patches of Ephestia kuehniella (Lepidoptera: Pyralidae) eggs, we compared two strains of Trichogramma cacoeciae Marchal and three strains of T. brassicae Bezdenko (Hymenoptera: Trichogrammatidae) for their strategies in colonizing hosts. The method by which females distribute their eggs among hosts was assessed either in terms of the average distance between parasitized hosts within a given patch, or of the average distance of attacks in discontinuous patches arranged in a circle.Within each strain, the traits increased as females aged; they vary among strains, their range of variations within T. cacoeciae falling within that of T. brassicae. Variations of both traits are correlated and strains can be easily classified according to their overall tendency to concentrate their attacks.The adaptive importance of the foraging behaviour of parasitoid insects suggests that differences reported here could reveal adaptive behavioural differentiations of natural populations in response to local variations in selective constraints, mainly those emanating from host diversity, abundance and distribution.

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Résumé Le mode de répartition des attaques parmi les hôtes est estimé par la distance moyenne entre les hôtes parasités dans une grande ooplaque homogène (500 hôtes) ou dans un système discontinu formé de six ooplaques de taille plus réduite (50 hôtes), régulièrement réparties sur un cercle.Pour chaque souche les deux caractères augmentent avec l'âge des femelles. Ils varient entre les souches, l'amplitude des variations au sein de l'espèce brassicae englobant celles observées chez cacoeciae.Les variations des deux caractères sont corrélées et les souches peuvent être classées suivant leur tendance plus ou moins marquée à concentrer leurs attaques.L'importance adaptative du comportement de colonisation des hôtes chez les insectes parasitoïdes laisse supposer que les différences constatées ici traduisent des différenciations locales des populations naturelles en réponse aux variations des contraintes sélectives, plus particulièrement celles qui sont liées à la diversité des hôtes, à leur abondance et à leur répartition.