斑痣悬茧蜂的寄主辨别能力及其影响因素

为评价斑痣悬茧蜂Meteorus pulchricornis (Wesmael)的寄主辨别能力及其影响因素, 采用双选试验（斜纹夜蛾2龄寄主幼虫, 健康∶被寄生=5∶5）, 观察了寄生经历（无寄生经历、 有1次寄生经历、 有1次过寄生经历）和寄主被首次寄生后的间隔时间（1 - 7 d）对斑痣悬茧蜂在健康寄主和被寄生寄主之间的选择; 为探究斑痣悬茧蜂是否能够辨别寄主斑块质量, 观察了斑痣悬茧蜂连续3次访问不同质量寄主斑块（被寄生寄主∶健康寄主分别为 2∶8, 5∶5和 8∶2）时的产卵刺扎次数。对选择频次进行的分析表明, 寄主被首次寄生后的间隔时间和寄生蜂的寄生经历均对过寄生发生有显著影响(P<0.05), 过寄生概率随寄主首次被寄生后的间隔时间延长而降低; 有寄生经历的寄生蜂发生过寄生的概率低于无寄生经历的寄生蜂。用Cox比例风险模型对寄主辨别时间进行的分析表明, 发生过寄生的风险随寄主被初次寄生后间隔时间的延长而减小, 也因寄生蜂具有过寄生经历而减小。斑痣悬茧蜂在连续3次访问不同质量寄主斑块中, 产卵刺扎次数随寄主斑块的质量提高而显著增多。据此推论, 斑痣悬茧蜂不仅能够辨别被寄生寄主, 而且能够辨别含有被寄生寄主的寄主斑块。     

斑痣悬茧蜂的寄主种间选择性及其子代蜂发育适合度表现

为检验基于经典搜寻理论的最优膳食模型的预测——产卵雌蜂应该选择最适于子代蜂发育的寄主种进行产卵,本研究以斑痣悬茧蜂(Meteorus pulchricornis)及其寄主斜纹夜蛾(Spodoptera litura)和粘虫(Mythimna separata)的幼虫为材料,在控制寄主体型大小和龄期影响的基础上,分别在体型大小相近和日龄相同的3个水平上观察了斑痣悬茧蜂对2种寄主幼虫的选择偏好,并观察了子代蜂生长发育适应度表现。在观察期(1h)内,当2种寄主幼虫的体型相近或者龄期相同的情况下,斑痣悬茧蜂对粘虫的产卵器刺扎次数以及寄生率(用结茧率表示)均高于斜纹夜蛾;而在斜纹夜蛾幼虫体内发育出的子代蜂茧重和体型均大于粘虫,成蜂寿命无显著差异。最后,对斑痣悬茧蜂的寄主选择和子代发育表现不一致现象进行了讨论。     

螟蛉盘绒茧蜂的过寄生行为及其对子代生长发育的影响

【目的】过寄生现象普遍存在于寄生蜂寄生过程中。本研究旨在探究螟蛉盘绒茧蜂Cotesia ruficrus的过寄生对子代发育的影响及影响过寄生行为的主要因素。【方法】室内研究了螟蛉盘绒茧蜂在其寄主稻纵卷叶螟Cnaphalocrocis medinalis3龄幼虫上过寄生行为的发生,不同产卵次数对寄主存活及子代蜂生长发育的影响,研究了不同接蜂时间和不同接蜂密度对过寄生发生的影响。【结果】螟蛉盘绒茧蜂存在过寄生行为,无论雌蜂有无产卵寄生经历,均能在被自身寄生过和同种不同个体寄生过的寄主内产卵。寄生蜂茧量随着被产卵次数的增加而增加,被产卵3~5次的寄主体内死亡的寄生蜂幼虫数随着增大。寄主在育出蜂前的死亡率随着被产卵寄生次数的增加而增加,被产卵5次时,寄主育出蜂前死亡率达50%。过寄生使螟蛉盘绒茧蜂子代蜂卵-蛹的历期延长,羽化率和雌雄性比下降,雌蜂体型随寄生次数的增加显著变小。过寄生率随着接蜂密度及接蜂时间的增加而增加。【结论】在寄主上产卵2次对螟蛉盘绒茧蜂子代发育是最适合的,产卵3次及以上为过寄生。过寄生使蜂子代发育适合度降低,不利于子代生长发育。在室内大量繁殖螟蛉盘绒茧蜂时,应减少接蜂密度和接蜂时间,从而减少过寄生的发生。     

斑痣悬茧蜂在黏虫寄主体内与中红侧沟茧蜂竞争下的存活和发育表现

【目的】不同种寄生蜂在同一小生境中可能寄生同一共享寄主。本研究旨在为认识寄生蜂种间互作关系,并为科学评价寄生性天敌控害效应提供依据。【方法】在室内以单寄生性斑痣悬茧蜂Meteorus pulchricornis(Wesmael)和中红侧沟茧蜂Microplitis mediator(Halidy)及其寄主黏虫Mythimna separata(Walker)低龄幼虫为材料,以寄生顺序和间隔时间为因素进行种间寄生竞争效应实验,观察比较两种蜂的存活和斑痣悬茧蜂子代的发育历期、体型大小和寿命等发育参数。【结果】不同寄生顺序和间隔时间组合处理下斑痣悬茧蜂子代蜂存活率始终高于中红侧沟茧蜂,且先寄生的斑痣悬茧蜂子代存活率高于后寄生的蜂,提前24 h寄生的斑痣悬茧蜂子代存活率可高达96.7%。当斑痣悬茧蜂先于中红侧沟茧蜂寄生时,其子代幼虫发育历期比单独寄生(对照)时显著延长;当后于中红侧沟茧蜂寄生时,其子代幼虫发育历期与单独寄生时无显著差异。各竞争处理下,斑痣悬茧蜂子代成虫寿命均长于单独寄生的子代成虫;子代成虫后足胫节长度与单独寄生相比明显缩短。【结论】结果说明,斑痣悬茧蜂在黏虫低龄幼虫体内与中红侧沟茧蜂的竞争中占有优势,但竞争对斑痣悬茧蜂子代生长发育具有负面影响。     

斑痣悬茧蜂的飞行能力和野外扩散行为

为探究斑痣悬茧蜂在野外的寄主搜寻行为,首先利用飞行磨测定了最大理论飞行能力,然后在大豆田进行了两次放蜂试验,通过回收或采集斜纹夜蛾寄主幼虫统计寄生情况,分析了寄生蜂的有效搜寻及其影响因素.飞行磨吊飞测定结果表明,个体起飞次数为14.48次±0.82次,总飞行距离1283.00 m±45.60m,总飞行时间2374.79 s±89.91 s,平均飞行速度0.54 m·s-1±0.004 m·s-1,最长的单次飞行时间475.21 s±76.97 s.田间放蜂试验结果表明,在风速≤4.2 m·s-1的大豆田中,方位对寄生率没有显著影响;但寄生率随时间延长和离放蜂点的距离增加而显著降低.放蜂的次日寄生蜂可扩散到36 m处.最后,对斑痣悬茧蜂野外搜寻行为特征进行了讨论.     

斑痣悬茧蜂对斜纹夜蛾不同虫龄幼虫的选择及后代发育表现

为探究容性寄生蜂对不同龄期寄主幼虫的选择性及其子代蜂发育表现的关联,通过双选试验观察了斑痣悬茧蜂(Meteorus pulchricornis)对斜纹夜蛾(Spodoptera litura)不同虫龄幼虫的寄生选择,并观察了与子代蜂适合度相关的特性表现.结果表明:斑痣悬茧蜂在4龄与5龄之间未表现出偏好,在2龄和3龄、3龄和4龄之间显著偏好较低虫龄;结茧率在不同寄主虫龄间无显著差异,羽化率随寄主虫龄增大而减小,寄生2、3龄幼虫的子代蜂显著高于寄生5龄;寄生5龄幼虫的子代蜂死亡率比寄生2龄的高2.5倍,比寄生3龄的高5.4倍.寄生3龄幼虫的子代蜂发育历期最短(11.9 d),比寄生4龄幼虫的短6.8 d,比寄生2龄幼虫的短4.7 d;子代蜂体型大小在寄生的寄主虫龄间无显著差异.根据研究结果推测,斑痣悬茧蜂在寄生时可能不是根据寄主龄期来评价寄主品质,而是基于寄主体型大小进行评价.     

斑痣悬茧蜂寄主搜索中的学习行为

利用风洞技术研究了斑痣悬茧蜂不同阶段的经历对其寄主甜菜夜蛾搜索行为的影响.结果表明:寄生蜂幼期发育到羽化早期所经历的植物对其成虫搜索行为具有一定的影响,在供试的大豆、棉花和白菜等3种植物中,只有大豆-寄主幼虫复合体对无经验寄生蜂的行为反应具有显著影响.寄生蜂1次产卵经历对后续搜索行为的影响取决于饲养寄主幼虫的植物种类,只有在大豆和白菜上的产卵经历对后续搜索行为产生显著影响,即明显偏好经历过的植物;对于在不同植物上经历2次产卵的寄生蜂,后续搜索行为反应取决于植物种类及其顺序,若2次产卵经历发生在大豆与白菜之间,无论顺序先后,后续行为反应均偏好大豆-寄主幼虫复合体;但若2次产卵经历发生在大豆与棉花之间,只有当第2次为大豆时,后续行为反应显著偏好大豆-寄主幼虫复合体;若2次寄生经历发生在棉花与白菜之间,只有当第2次经历为白菜时,后续行为反应显著偏好白菜-寄主幼虫复合体.根据研究结果推断,斑痣悬茧蜂幼期发育、成虫早期羽化的经历、成虫产卵经历对后续搜索行为具有显著影响,而且这种影响与寄主幼虫的食料植物有关.     

聚寄生性蝶蛹金小蜂雌蜂体型大小对产卵策略的影响

在寄生蜂行为生态学研究中, 通常将寄主体型大小作为寄主品质的主要性状来探究寄生蜂的搜寻行为机理, 而忽略寄生蜂体型大小的意义。为揭示聚寄生蜂雌蜂体型大小对其产卵决策的影响, 在严格控制寄主菜粉蝶Pieris rapae蛹体型大小（体重）的情况下, 于室内观察了不同体型大小的蝶蛹金小蜂Pteromalus puparum雌蜂的产卵行为, 并调查了子代蜂数量（窝卵数）、 性比和体型大小的变化。结果表明： 雌蜂在寄主上的驻留时间随其自身体型增大而缩短, 但随寄主体重增大而延长。窝卵数和余卵量受到雌蜂体型大小的显著影响, 均随雌蜂体型增大而显著增加（P<0.05）; 但子代蜂性比不受雌蜂体型大小的显著影响 (P>0.05)。子代雌、 雄性体型大小均与雌蜂体型大小无关, 但子代雌蜂体型随寄主体重增大而增大。结果证实, 雌性蝶蛹金小蜂体型大小影响其部分产卵决策。因此, 在建立聚寄生蜂产卵决策模型中应考虑雌蜂体型大小这一重要变量因素。     

核型多角体病毒与侧沟茧蜂对斜纹夜蛾幼虫的协同作用

研究了斜纹夜蛾幼虫体内的斜纹夜蛾侧沟茧蜂存活率、发育历期、寄主感染病毒时间、病毒浓度之间的关系,并测定了斜纹夜蛾侧沟茧蜂的传毒效率．结果表明,病毒对寄主体内寄生蜂历期无明显影响,寄生在幼虫体内的寄生蜂能在寄主病死前完成发育,存活比例因寄主感染病毒的时间和浓度而异．斜纹夜蛾被寄生后接种病毒(SINPV),距离寄生时间越长,饲毒浓度越低,寄生蜂完成发育的比例越大,但饲毒时间是主要影响因素．从感病幼虫体内发育成的侧沟茧蜂或曾经在感病寄主上产过卵的寄生蜂,以及通过人工方式使产卵器被病毒污染的寄生蜂,均能携带一定数量的病毒．通过产卵活动,侧沟茧蜂成蜂能在寄主幼虫个体间传递病毒．当寄生蜂在感病的寄主幼虫上产卵带毒后,平均可传递病毒给2．14头幼虫;发育于感病幼虫体内的寄生蜂,平均可传递病毒给2．45头幼虫．通过用病毒液浸茧或用混有病毒的蜂蜜饲喂成蜂等方式使产卵器污染病毒的寄生蜂,传毒效率随饲毒浓度增加而提高,平均可传递病毒1．45头和0．94头幼虫     

甜菜夜蛾不同龄期幼虫被斑痣悬茧蜂寄生的风险分析

寄生蜂的寄主识别期是构建最优接受寄主模型的重要参数之一。本研究观察了斑痣悬茧蜂(Meteorus pulchricornis)对甜菜夜蛾(Spodoptera exigua)不同龄期幼虫的识别期,并首次用生存分析方法估计寄主被寄生的风险以及寄生经历对寄主识别期的影响。结果表明:在2.5h观察期内,除1龄寄主幼虫外,其他各龄(2~5龄)幼虫均被寄生;对首次寄主识别期的Cox回归模型拟合分析表明,与2龄寄主幼虫(参照虫龄)相比,3~5龄寄主幼虫被寄生的累计风险比率显著高于2龄寄主幼虫(2·6~4·0倍),而对再次寄主识别期的分析未发现3~5龄寄主幼虫与2龄幼虫之间存在显著差异。根据对首次与再次寄主识别期进行的生存曲线比较表明,对2~4龄幼虫的再次寄主识别期显著短于首次寄主识别期;说明寄生经历显著提高了寄生蜂的寄主识别期。由此推测,较高龄期甜菜夜蛾幼虫被无寄生经验斑痣悬茧蜂寄生的风险高于低龄幼虫。     

The impact of patch encounter rate on patch residence time of female parasitoids increases with patch quality

Abstract.  1. For animal species that forage on patchily distributed resources, patch time allocation is of prime importance to their reproductive success. According to Charnov's marginal value theorem (MVT), the rate of patch encounter should influence negatively the patch residence time: as the rate of patch encounter decreases, the patch residence time increases. Moreover, the MVT predicts that animals should stay longer in high quality patches.
2. Using the aphid parasitoid Aphidius rhopalosiphi (Hymenoptera: Aphidiinae), the effects of these two factors (patch encounter rate and host density) were combined in order to test if the increment in patch residence time for a given decrease in patch encounter rate was larger for high quality patches than for low quality patches.
3. The results show a significant effect of the interaction between the two factors. In high host density patches, parasitoids spent more time if they experienced a low patch encounter rate, while in low host density patches, patch encounter rate had no significant effect on the patch residence time. This suggests that the response of A. rhopalosiphi females to patch encounter rate varied with host density in the patch. Moreover, the same interaction effect was observed for the number of ovipositor contacts on aphids.
4. Parasitoid females can use patch encounter rate to estimate patch density in the habitat but the effect of this estimate on their patch residence time is modulated by patch quality. Staying longer in a patch when patches are rare is more advantageous when the fitness gained by doing so is large. In low quality patches, the expected fitness gain is small and the female may gain more by leaving and taking her chance at finding another patch.     

Nectar provisioning close to host patches increases parasitoid recruitment,retention and host parasitism

In the adult stage, many parasitoids require hosts for their offspring growth and plant-derived food for their survival and metabolic needs. In agricultural fields, nectar provisioning can enhance biological control by increasing the longevity and fecundity of many species of parasitoids. Provided in a host patch, nectar can also increase patch quality for parasitoids and affect their foraging decisions, patch time residence, patch preference or offspring allocation. The aim of this study was to investigate the impact of extrafloral nectar (EFN) provisioning close to hosts on parasitoid aggregation in patches. The aphid parasitoid Diaeretiella rapae (M’Intosh) was released inside or outside patches containing Brassica napus L. infested by Brevicoryne brassicae L. aphids and Vicia faba L. with or without EFN. When parasitoids were released outside patches, more parasitoids were observed in patches with EFN than in patches deprived of EFN. This higher recruitment could be linked to a higher attraction of a combination of host and food stimuli or a learning process. A release–recapture experiment of labeled parasitoids released within patches showed the higher retention of parasitoids in patches providing EFN and hosts, suggesting that food close to the host patch affects patch residence time. Both attractiveness and patch retention could be involved in the higher number of parasitoids foraging in host patches surrounded by nectar and for the higher parasitism recorded. Nectar provisioning in host patches also affected female offspring allocation inside the patch.     

Foraging Behavior and Patch Time Allocation by Fopius arisanus (Hymenoptera: Braconidae), an Egg-Larval Parasitoid of Tephritid Fruit Flies

This study quantitatively describes the host-searching behavior of Fopius arisanus (Sonan) (Hymenoptera: Braconidae), an important egg-larval parasitoid of tephritid fruit fly pests, on coffee berries infested with host eggs of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). We also investigate the parasitoid's response to local variation in host patch quality. The temporal pattern of behavioral organisation was examined by constructing an ethogram. The parasitoid spent over 90% of its foraging time in detecting and locating hosts after arriving on a host-infested fruit, and displayed a relatively fixed behavioral pattern leading to oviposition. Patch residence time increased in the presence of host-associated cues, following successful ovipositions, and with increasing size of host clutches per fruit, but decreased with each successive visit to the same host patch and with increasing availability of alternative host patches. The parasitoid females discriminated against previously parasitized hosts and spent significantly less time and searching effort on patches previously exploited by herself or by conspecific females. The effective host-searching behavior, perfect host discrimination ability, and success-motivated searching strategy shown by F. arisanus ensured a thorough exploitation of host resources by this parasitoid.     

X-ray computerized tomography as a new method for monitoring Amegilla holmesi nest structures, nesting behaviour, and adult female activity

Optimal foraging theory, specifically the marginal value theorem, predicts quicker leaving (shorter residence time) from poorer patches. One proximal mechanism for achieving the leaving is that exposure to lower-quality resources may trigger increased restlessness (proportion of time in locomotion). Which aspects of host quality, if any, affect restlessness was examined in females of the parasitoid wasp Nasonia vitripennis Walker (Hymenoptera: Pteromalidae). Females were individually exposed to a single host. Restlessness was greater both during and after exposure to a host, when the host was externally damaged vs. intact. Other aspects of host quality that affected restlessness were whether the host was parasitized and whether it was dead and unsuitable for offspring development. In contrast, the host's age and stage did not affect restlessness. Increased restlessness did not make females more willing to launch themselves across an inhospitable environment using their wings.     

Parasitoid aggregation and interference in host–parasitoid dynamics

1. Parasitoids do not distribute themselves evenly among available patches, which has an important bearing on the dynamics of host–parasitoid interactions. This study examined the density‐dependent nature of aggregation of the parasitoid Dirhinus giffardii Silvestri on the oriental fruit fly host, Bactrocera dorsalis (Hendel) distributed among discrete patches. 2. Four artificial patches were created in a cage, and the number of hosts in each patch was manipulated. Parasitoids were released into the cage, and whether parasitoid density and host density influence the degree of parasitoid aggregation was examined. 3. Parasitoid aggregation became stronger (e.g. uneven distribution among patches) as the parasitoid density decreased and also as the host density increased. The index of parasitoid aggregation was not influenced by the distribution of hosts among patches. 4. The empirically characterised aggregation pattern was incorporated in a host–parasitoid model that consists of one host and one parasitoid species. The analysis of the model shows that an unstable system (i.e. the coexistence of the host and parasitoid is impossible) can be stabilised (i.e. coexistence is possible) when the parasitoid aggregates in a way that is consistent with the pattern found in the experiment.     

Emergence of global behaviour in a host–parasitoid model with density-dependent dispersal in a chain of patches

We present a time discrete spatial host–parasitoid model. The environment is a chain of patches connected by dispersal events. Dispersal of parasitoids is host-density dependent. When the host density is small (resp. high), the proportion of migrant parasitoids is close to unity (resp. to zero). We assume fast patch to patch dispersal with respect to local interactions. Local host–parasitoid interactions are described by the classical Nicholson–Bailey model. By using time scales separation methods (or aggregation methods), we obtain a reduced model that governs the total host and parasitoid densities (obtained by addition over all patches). The aggregated model describes the time evolution of the total number of hosts and parasitoids of the system of patches. This global model is useful to make predictions of emerging behaviour regarding the dynamics of the complete system. We study the effects of number of patches and host density-dependent parasitoid dispersal on the overall stability of the host–parasitoid system. We finally compare our stability results with the CV² > 1 rule.     

Factors affecting the rate of attack by Cotesia rubecula (Hymenoptera: Braconidae)

Abstract. 1. The relationship between responses of the insect parasitoid Cotesia rubecula (Marshall) to kairomones produced by the feeding activities of its host, Pieris rapae (L.), and patterns of parasitism were investigated under field conditions.
2. Parasitoid adults aggregated in patches with the highest densities of host larvae but there was no commensurate increase in the rate of attack in these patches.
3. The rate of attack was not limited by the availability of eggs.
4. The rate of parasitoid attack was highest where feeding damage by the host was highest, irrespective of current host density.
5. The rate of parasitoid attack was further influenced by host age distribution. Late instar larvae were less susceptible to parasitism than were early instar larvae. The rate of attack on early instar larvae occupying the same plants as late instar larvae was reduced. This reduction in rate of attack was due to limitations on parasitoid search time imposed by the increased feeding damage associated with large host larvae and by the increased time the parasitoid required to recover from an attack on these large host larvae.     

Habitat assessment by parasitoids: mechanisms for patch use behavior

Animals foraging for patchily distributed resources may optimizetheir foraging decisions concerning the patches they encounter,provided that they base these decisions on reliable informationabout the profitability of the habitat as a whole. Females ofthe parasitoid Lysiphlebus testaceipes exploit aphid hosts,which typically aggregate in discrete colonies. We show herehow between-colony travel time and the number of aphids in previouslyvisited colonies affect parasitoid foraging behavior. We firstassumed that parasitoids use travel time and previous colonysize to estimate a mean rate of fitness gain in the habitatand derived quantitative predictions concerning the effect ofthese two variables on patch residence time and patch-leavingrate of attack. We then tested these theoretical predictionsin laboratory experiments in which female parasitoids were allowedto visit two successive colonies. As predicted, the observedresidence time in the second colony increased with increasingtravel time and decreasing size of the first colony. Patch-leavingrate of attack decreased with increasing travel time but wasnot affected by previous colony size. These results suggestthat parasitoids use these two variables to assess habitat quality.However, discrepancies between the data obtained and quantitativepredictions show that the effect of travel time on patch usemay be more complex than assumed in our model.     

Effects of asymmetric dispersal and environmental gradients on the stability of host–parasitoid systems

We consider host–parasitoid systems spatially distributed on a row of patches connected by dispersal. We analyze the effects of dispersal frequency, dispersal asymmetry, number of patches and environmental gradients on the stability of the host–parasitoid interactions. To take into account dispersal frequency, the hosts and parasitoids are allowed to move from one patch to a neighboring patch a certain number of times within a generation. When this number is high, aggregation methods can be used to simplify the proposed initial model into an aggregated model describing the dynamics of both the total host and parasitoid populations. We show that as the number of patches increases less asymmetric parasitoid dispersal rates are required for stability. We found that the 'CV²>1 rule' is a valid approximation for stability if host growth rate is low, otherwise the general condition of stability we establish should be preferred. Environmental variability along the row of patches is introduced as gradients on host growth rate and parasitoid searching efficiency. We show that stability is more likely when parasitoids move preferentially towards patches where they have high searching efficiency or when hosts go mainly to patches where they have a low growth rate.     

Plant patch structure modifies parasitoid assemblage richness of a specialist herbivore

1. The spatial structure of plant patches has been shown to affect host–parasitoid interactions, but its influence on parasitoid diversity remains largely ignored. Here we tested the prediction that parasitoid species richness of the specialist leafminer Liriomyza commelinae increases in larger and less isolated patches of its host plant Commelina erecta. We also explored whether parasitoid abundance and body size affected the occurrence of parasitoid species in local assemblages. 2. A total of 893 naturally established C. erecta patches were sampled on 18 sites around Córdoba city (Argentina). Also, two experiments were performed by creating patches differing in the number of plants and the distance from a parasitoid source. For these tests, plants were infected with the miner in the laboratory prior to placement in the field. 3. Plant patch size, independently of host abundance, positively affected the number of parasitoid species in both survey observations and experimental data. However, plant patch isolation did not influence parasitoid species richness. 4. The probability of finding rare parasitoid species increased with patch size, whereas occupation of isolated patches was independent of dispersal abilities (body size) of parasitoid species. 5. Overall, the results highlight the importance of considering spatial aspects such as the size of plant patches in the study of parasitoid communities.