拟寄生蜂搜索产卵过程中对寄主的竞争

综述拟寄生蜂搜索产卵过程中对寄主竞争的最新研究进展.这类竞争具有四种方式,即标记寄主、杀卵和杀幼、守护寄主和捕食寄主.（1）标记寄主常涉及寄主标记信息素,这是由雌蜂在产卵前、产卵时或产卵后分泌的化学物质.寄主标记信息素常介导拟寄生蜂对已寄生和健康寄主的辨别、减少过寄生和多寄生、减轻种内和种间竞争压力.（2）雌蜂遇到已寄生寄主时,很多种类杀死前一雌蜂遗留的卵和幼虫,再产下自己的卵.雌蜂使用三种方法杀卵和杀幼,即产卵器穿刺、取食和使用有毒物质.通过杀卵和杀幼,产卵雌蜂清除了前一雌蜂遗留的后代,主动改善了寄主品质,从而有利于自身后代的生存.（3）守护寄主在肿腿蜂科、缘腹细蜂科、金小蜂科、缨小蜂科和茧蜂科中均有报道,守护者驱逐入侵者以保护后代及健康寄主.（4）捕食寄主不仅减少了健康寄主数量,且直接导致已寄生寄主中拟寄生蜂卵和幼虫的死亡.雌蜂一般在体内成熟卵量较少时捕食寄主.讨论了研究拟寄生蜂搜索产卵过程中竞争寄主的理论意义和实际应用价值.     

To mark the host or the patch: Decisions of a parasitoid searching for concealed host larvae

We found evidence for patch marking in the parasitic wasp Halticoptera laevigata (Hymenoptera: Pteromalidae) foraging for concealed hosts. Wasps attack larvae of the fruit fly Myoleja lucida (Diptera: Tephritidae) in fruits of honeysuckle. A special feature of this host-parasitoid system is the limited food supply of a patch (i.e. a fruit of honeysuckle), which allows the successful development of only a single host fly larva. Females of the parasitoid H. laevigata were found to mark the host patch with a pheromone and to abandon the patch following oviposition into a single host larva. Field data revealed that eggs of the parasitoid were spread out evenly among infested patches, with several larvae of the host fly left unparasitized in those patches that contained more than one host. Since many parasitic insects mark the parasitized host after oviposition, we assumed host marking to be the ancestral character state and studied the patch-marking behaviour of H. laevigata as a derived character state as an alternative foraging strategy. We used stochastic dynamic modelling to investigate under what conditions mutant (patch) markers would be able to invade a population of normal (larval) markers. The models suggested that, under a variety of conditions, wasps marking the patch obtained higher fitness than wasps only marking the larva. Consequently, the results from our model predict the evolution of the patch-marking behaviour found in the empirical investigation. Finally, we discuss alternative pathways to the evolution of patch marking and point out under what circumstances the evolution of a patch-marking behaviour can generally be expected.     

拟寄生蜂的寄主标记研究进展

拟寄生蜂常借助寄主标记信息素 (host marking pheromone, HMP) 来辨别已寄生寄主和健康寄主,避免过寄生和多寄生,减少后代的种内和种间竞争。寄主标记有外部标记、内部标记和“容器”及区域标记3种方式。HMP来源于拟寄生蜂的杜氏腺、毒腺、输卵管、卵巢、卵或咽侧体,由触角或产卵器感受。目前已鉴定出几种拟寄生蜂的HMP,多数为饱和与不饱和烃类的混合物,但卡氏盾痣细蜂Dendrocerus carpenteri的HMP是保幼激素。拟寄生蜂对寄主标记的反应受HMP持效期,拟寄生蜂内部状况如载卵量、年龄、经验和学习,及外部条件如拟寄生蜂和健康寄主种群密度以及寄主种类的影响。本文还讨论了HMP研究的理论和实际意义。     

Evidence for a host-marking pheromone in Chaetorellia australis

Experiments and observations on the oviposition behavior of Chaetorellia australis Hering (Diptera, Tephritidae) females, caged in the laboratory with their natural oviposition substrates, floral buds of the yellow starthistle (YST) Centaurea solstitialis L. and Centaurea cyanus L. (Asteraceae: Cardueae) revealed that females of that species deposit a contact host-marking pheromone immediately after oviposition by dragging their extended ovipositor over the floral bud surface. The function of this pheromone is the prevention of repeated oviposition in already infested oviposition sites. Apparently it also plays a role in sexual behavior by functioning as a male arrestant. The pheromone was found to be soluble in water but insoluble in methanol, ethanol, diethyl ether and acetone. Under laboratory conditions, its effectiveness persisted for at least 10 days. By preventing oviposition on already infested buds the pheromone apparently contributes to uniform egg dispersion among available resources for larval development. By functioning as a male arrestant, which causes aggregations of males on places visited frequently by females such as the floral buds of host plants, it also contributes to overall fertility by facilitating encounters of the sexes. The potential practical application for biocontrol of the YST through introducing and releasing C. australis flies into areas infested by the weed is discussed.

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Zusammenfassung Versuche und Beobachtungen über das Eiablageverhalten von im Labor mit ihren natürlichen Eiablagesubstraten, d.h. Blütenknospen von Centaurea solstitialis L. und Centaurea cyanus L. (Asteraceae: Cardueae) gehaltenen Chaetorellia australis Hering (Diptera: Tephritidae) zeigten, dass C. australis Weibchen sofort nach der Eiablage die Oberfläche der Blütenknospe mit einem eiablageverhindernden Pheromon markieren, indem sie ihren ausgestellten Ovipositor über die Oberfläche der Blütenknospe ziehen. Das Pheromon, das auch als Männchen-Arrestans wirkt, ist wasserlöslich, jedoch unlöslich in Methanol, Äthanol, Diethyläther und Aceton. Unter Laborbedingungen hält seine Wirkung mindestens 10 Tage an. Da das Pheromon die Eiablage an bereits infestierten Blütenknospen verhindert, trägt es zu einer gleichmässigen Eiverteilung in den für die Larvenentwicklung zur Verfügung stehenden Futtequellen bei. Als Männchen arretierendes Pheromon führt es zur Aggregation von Männchen an den Blütenknospen, die oft von Weibchen besucht werden und versichert somit eine hohe Fertilität, indem es das Zusammentreffen der Geschlechter erleichtert. Die praktische Anwendung der Ergebnisse für die biologische Bekämpfung von Centaurea solstitialis durch Einfuhr und Freilassung von C. australis in die mit dem Unkraut verseuchten Gebiete wird diskutiert.

     

Marking of host fruit by maleRhagoletis boycei Cresson flies (Diptera: Tephritidae) and its effect on egg-laying

MaleRhagoletis boycei flies (Cresson) have been shown previously to guard egg-laying punctures on host walnut (Juglans major) fruit. During their tenure,R. boycei males were observed to dip the posterior portion of their abdomen repeatedly toward the fruit surface. Closeup video analysis revealed that, during this dipping behavior, a clear viscous substance emanating from the cercus was deposited directly on the fruit. Laboratory assays indicated that the rate of abdomen-dipping was significantly higher on fruit bearing simulated egg-laying punctures than on control fruit and significantly higher in the area of the puncture than elsewhere on the fruit. In a final experiment, the occurrence of abdomen-dipping in the vicinity of a puncture was manipulated independently on separate punctures on the same fruit. On such fruit and in the absence of males, previously mated, reproductively mature females attempted oviposition significantly more often in or near a puncture exposed to males than in or near an unexposed control puncture. Possible functions of the putative male-marking behavior from the perspectives of both male and female are discussed.     

Functional shifts in the use of parasitized hosts by a tephritid fly: the role of host quality

Superparasitism, a phenomenon in which parasitic insects layeggs in already-exploited hosts, provides a useful context inwhich to examine the dynamics of parental investment. This studyexplored conditions under which female Mediterranean fruit flies(Ceratitis capitata) shift from avoiding superparasitism ofhost fruit to preferring it, even placing eggs direcdy intoexisting egg-laying cavities. An a priori hypothesis of costsand benefits was used to predict how use and avoidance of parasitizedfruit would change in -response to changes in fruit size andripeness. We predicted that avoidance would decrease with increasingfruit size, while use would increase with decreasing ripeness.Using a field-cage assay, ripeness was held constant and thesize of host coffee berries manipulated. Avoidance of parasitizedberries was significandy less pronounced on large berries thanon small ones. In a second experiment, size was held constantand ripeness manipulated. On unripe berries, females depositedthe majority of clutches directly into existing egg-laying cavities.On ripe berries, by contrast, the same females deposited mostclutches in previously unparasitized fruit. Parallel patternsin the frequency of female-female contests were observed, supportingthe notion that a fruit's value is determined by an interactionbetween fruit size or ripeness, on one hand, and the prior occurrenceof eggs, on the other. Laboratory assays suggested that useof existing sites had advantages in terms of time savings; femalebehavior thus constitutes a relatively uncommon example of adaptivesuperparasitism in which parasitized hosts are actually preferredover unparasitized ones.[Behav Ecol 7: 235-242 (1996)]