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。     

管氏肿腿蜂对黄粉甲的寄生行为

在人工扩繁条件下观察管氏肿腿蜂Scleroderma guani对中间寄主黄粉甲Tenebrio molitor的寄生行为,其过程可分为：搜寻-识别-螯刺-清理和取食-产卵-护幼。雌蜂在寄生黄粉甲蛹时上蜂速度和螯刺率变异大,上蜂前时间为1.4～7.0天, 螯刺率为8.7%～91.7%,影响两者的因素包括： 雌蜂的交配状况、寄主状况及繁育温度等。蜂毒螯入寄主后可使其组织不腐烂而适于被寄生,被螯刺后的蛹可表现为 “保鲜”状态,平均“保鲜”时间为12.6天。蜂在蛹体的平均产卵量为30粒,最多为91粒。蜂卵在蛹体上分布合理,有利于幼虫有效利用寄主营养并避免拥挤和取食竞争。母蜂对幼虫期的发育具保护行为,包括护卵和保护幼虫正常发育并能搬运老熟幼虫集中结茧化蛹。     

白蜡吉丁柄腹茧蜂的寄生和繁殖生物学

白蜡吉丁柄腹茧蜂Spathius agrili Yang是我国天津地区寄生白蜡窄吉丁幼虫的一种重要天敌。本研究通过林间调查并结合室内观察,测定了该寄生蜂在不同营养条件下的存活时间和不同时期的产卵量,调查了不同寄主大小对寄生蜂后代生长发育的影响,以及寄生蜂对不同深度树皮下的寄主利用情况。结果表明：不同营养条件下白蜡吉丁柄腹茧蜂的存活时间差异很大,补充20％蜂蜜水的成蜂存活时间明显延长,最长可达3个月。相同条件下雌蜂的存活时间比雄蜂略长。在补充营养的条件下,茧蜂的产卵时间最长可达8周,产卵的高蜂在羽化后第2周前后,平均每雌每周产卵量在整个产卵期内基本保持在8～12粒上下,没有大的波动。茧蜂低龄幼虫也可能分泌毒素主动攻击寄主,提高了寄生蜂后代的生存能力。寄主个体大,营养量也大,寄生其上的茧蜂幼虫成功完成发育的数量随之增加,表明寄主营养资源的多少直接影响到寄生蜂后代的生长发育和存活。白蜡吉丁柄腹茧蜂对1～6.5 mm树皮深度范围内的适龄寄主幼虫均能够寄生利用,约92％的被寄生白蜡窄吉丁幼虫个体分布于1～4 mm深度的树皮下,在3 mm厚的树皮下分布的数量最多。茧蜂数量与树皮深度大致上呈抛物线关系,理论上在树皮深度为3.97 mm处茧蜂分布数量最多。     

中红侧沟茧蜂寄生粘虫的生物学特性(英文)

报道了中红侧沟茧蜂寄生粘虫时的某些生物学特性。雌蜂的每次产卵动作约有３．１％的比率产出两粒卵,４．６％的比率没有卵产出。寄主龄期与寄生率呈负相关关系,雌蜂在寄主体内产两粒卵能提高寄生率。寄主龄期同后代寄生蜂的性比没有相关性,不是影响性比的主要因素。文中讨论了影响中红侧沟茧蜂性比的可能因素,并认为二龄粘虫是大规模繁殖该寄生蜂的最好寄主。     

寄主大小对蝇蛹俑小蜂Spalangia endius(Walker)产卵选择和发育的影响

【背景】对寄生蜂与寄主之间关系的研究是寄生蜂有效利用的基础。寄生蜂雌蜂通常选择最适于子代蜂发育的寄主进行产卵。【方法】在室内26℃条件下,研究了橘小实蝇蛹体型大小对蝇蛹俑小蜂产卵和生长发育的影响。采用饥饿方法处理3龄橘小实蝇幼虫以获得体型大小(用体质量表示)差异较大的寄主蛹,供寄生蜂选择寄生。【结果】蝇蛹俑小蜂显著偏好寄生体型中等的寄主蛹,然而,随着寄主体型的增大,后代雌性比率增大,且寄主蛹个体大小与后代雌蜂体型大小存在显著的正相关;橘小实蝇蛹个体大小对蝇蛹俑小蜂后代发育历期和寿命无显著影响。【结论与意义】蝇蛹俑小蜂雌蜂能够根据寄主蛹质量来调整后代数量和性比,以使后代适应度最大化。     

啮小蜂寄主接受行为的研究

啮小蜂Tetrastichus sp.是一种群聚性蛹的内寄生蜂,主要寄生于茶长卷叶蛾的蛹.在实验室中,它还能寄生蓖麻蚕、粘虫、亚洲玉米螟、棉褐带卷蛾和家蝇等寄主蛹.在小环境中,啮小蜂雌蜂搜寻大蛹(粘虫)花费的时间要明显少于寻找小蛹(棉褐带卷蛾)所花费的时间.雌蜂多数从寄主的腹部爬上蛹,其次为胸部和头部.88%的雌蜂产卵管插入蛹的位置与上蛹的部位一致.蜂龄(2-6天龄)对雌蜂接受寄主的行为无明显影响.雌蜂对寄主蛹龄有一定的辨别能力,在第7天龄粘虫蛹中产的子蜂数明显减少,对第9天龄的粘虫蛹雌蜂虽作刺探,但未见寄生.随着寄主密度增加,平均每雌寄生的蛹数和产的子蜂数均相应增加,而寄主蛹的寄生率却逐渐下降.随着雌蜂密度增加,寄主蛹的寄生率和平均每头寄生蛹中的子蜂数相应提高,但平均每雌产的子蜂数则逐渐减少.雌蜂在寄生(平均1.74+5.05分.秒)和未寄生(平均2.90±6.50分.秒)寄主中产卵管插入的时间无明显差异,但在寄生(平均每蛹为182.20±85.06头)和未寄生(平均为87.80±69.50头)寄主中产的子蜂数则有明显差异(P<0.02).表明啮小蜂雌蜂对已寄生寄主蛹无明显的辩别能力.     

营养改变对潜蝇姬小蜂寄生行为和寄主取食行为的影响

为了研究营养状态对卵育型寄生蜂潜蝇姬小蜂Diglyphus isaea (Walker)雌蜂的寄主取食行为和产卵寄生行为及其二者行为权衡的影响, 在培养皿条件下, 比较了饥饿、加蜂蜜水和不加蜂蜜水3种营养状态的潜蝇姬小蜂雌蜂对美洲斑潜蝇Liriomyza sativae Blanchard各龄幼虫（低、中、高）的寄生、取食及致死能力。结果表明: 非选择条件下, 3种营养状态寄生蜂对高龄幼虫均具有较高的寄生率, 对中龄幼虫具有较高的取食率, 致死率和致死量之间存在显著性差异。3种状态的寄生蜂对低龄幼虫均没有表现出致死能力。有选择条件下, 饥饿状态的寄生蜂对寄主的寄生率最低（5.0%±1.6%）, 取食率最高（16.0%±2.9%）, 特别是对高龄幼虫的取食率占到了整个寄主食物取食率的91.9%; 加蜂蜜水状态下, 寄生蜂对寄主有最低的取食率（8.3%±0.9%）和致死率（17.7%±1.1%）; 不加蜂蜜水状态下, 寄生蜂对寄主有最高的寄生率（13.3%±1.1%）和致死率（28.4%±1.8%）。综合分析发现, 取食寄主的雌蜂比取食蜂蜜水的雌蜂具有更强的致死能力和活动能力。     

异律寄生蜂的寄主关系及其进化

异律寄生蜂（Heteronomous parasitoids）是指雌蜂、雄蜂卵及幼期分别对应不同的寄主关系和发育方式的寄生蜂。依据雌蜂、雄蜂卵及幼期所寄生的位置、所利用寄主种类和寄生方式的差异,异律寄生蜂可划分为异位初寄生蜂、异主初寄生蜂和异律复寄生蜂。本文重点关注蚜小蜂Coccophaginae亚科的异律寄生蜂,依据其雌蜂、雄蜂卵及幼期所对应的寄主关系、寄主种类和发育方式进行梳理与归类,总结了异律寄生蜂的生殖方式和异律发育模式,结合其生活史特性探讨了异律寄生蜂的寄主选择和适应性进化,并辨析了异律寄生蜂在生物防治中的利弊与应用。     

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。     

Fitness consequences of ovicide in a parasitoid wasp

Ovicide, superparasitism and host rejection are alternative reproductive tactics facing female parasitoid wasps encountering an already-parasitized host. Superparasitism is simply the addition of an egg or a clutch of eggs by the secondary parasitoid, but under ovicide the primary clutch is removed or destroyed. Host rejection occurs if the wasp leaves without laying a clutch. The ectoparasitoid Laelius pedatus (Say) (Hymenoptera: Bethylidae) performs ovicide in this situation. Clutch manipulation experiments show that secondary clutches suffer high mortality in competition with primary clutches, which increases with increasing time delay between clutches. Primary clutches however suffer little in competition with secondary clutches, even if there is minimal time delay between clutches. These data suggest that the offspring of ovicidal females are substantially fitter than the offspring of superparasitizing females. Handling time and clutch size do not differ significantly between first (sole) parasitoids and second (ovicidal) parasitoids. The same is true for offspring survival and development time. However, offspring of second females are slightly smaller. This suggests that parasitized and unparasitized hosts are resources of similar quality when ovicide is performed. These data strongly support the predictions of evolutionary models of ovicide. They may also give some insight into the taxonomic distribution of ovicide in parasitoids.     

Environmental and genetic determinants of ovicide in parasitic wasp Bracon hebetor

Superparasitism occurs when a parasitoid lays a second clutchof eggs on a host previously parasitised by herself or a conspecific.Ovicide refers to a parasitoid destroying an existing clutchof eggs on a parasitized host before laying a second clutch.We investigated environmental and genetic determinants of ovicidein the parasitic wasp Bracon hebstor. Characterization of egglayingbehavior revealed that B. hebetor commits ovicide during thehost examination phase of oviposition. The temporal costs ofovicide were found to be relatively small for females that experiencedlow rates of host encounter, whereas the costs of ovicide increasedfor females that experienced a high rate of host encounter.Individual wasps committed ovicide on conspecifically parasitizedhosts more frequently than on self-parasitized hosts. Manipulationexperiments suggested that B. hebetor females learn about theirenvironment while foraging and commit ovicide on the basis ofthe travel time between successive hosts. Significant differenceswere also found in ovicidal behavior among laboratory and fieldpopulations of B. hebetor. The implications of our results forclutch size theory and the evolution of ovicide are discussed.     

Ovicide in the whitefly parasitoid, Encarsia formosa

The oviposition decisions made by insect parasitoids when encountering hosts of variable quality have been the subject of extensive theoretical and experimental investigation. For parasitoids that lay their eggs inside the host, the possible outcomes of encounters with parasitized hosts have been assumed to include only oviposition (superparasitism), rejection, or in some cases feeding on host haemolymph. We document another outcome in Encarsia formosa Gahan (Hymenoptera: Aphelinidae), a species that has been a model system for the study of oviposition behaviour. In E. formosa, females may kill eggs previously laid within the host by jabbing them with their ovipositor before ovipositing themselves. (1) Our observations indicated that jabbed eggs were indeed killed. (2) In experimental arenas in the laboratory, ovicide occurred in the majority of encounters with parasitized hosts and at highest frequency in encounters resulting in oviposition. (3) There was no significant difference in the handling time associated with oviposition+ovicide in parasitized hosts in comparison with oviposition alone, suggesting that there is no time cost to ovicide. (4) Ovicide did not appear to be incidental to normal probing within a host. Radial analysis of the direction of ovipositor movement with respect to the centre of the previously laid egg within the host showed that females engaged in ovicidal bouts probed most often in the direction of the egg. This is the first well-documented study of ovicide in an endoparasitoid. We suggest ovicide may be under-reported in other endoparasitoid species due to the difficulty of observing it. Copyright 2000 The Association for the Study of Animal Behaviour.     

Shifting preference between oviposition vs. host-feeding under changing host densities in two aphelinid parasitoids

Destructive host-feeding is common in hymenopteran parasitoids. Such feeding may be restricted to host stages not preferred for oviposition. However, whether this is a fixed strategy or can vary according to resource levels or parasitoid needs is less clear. We tested the trade-off between host feeding and oviposition on two whitefly parasitoids under varying host densities. Females of two aphelinid parasitoids, Eretmocerus hayati and Encarsia sophia were exposed to nine different densities of their whitefly host, Bemisia tabaci, in single-instar tests to identify their functional response. Mixed-instar host choice tests were also conducted by exposing whiteflies at four densities to the parasitoids. We hypothesized that the parasitoid females can detect different host densities, and decide on oviposition vs. host-feeding accordingly. The results showed that both Er. hayati and En. sophia females tended to increase both oviposition and host-feeding with increased host density within a certain range. Oviposition reached a plateau at lower host density than host-feeding in Er. hayati, while En. sophia reached its oviposition plateau at higher densities. At low densities, Er. hayati parasitized most on first and second (the optimal ones), and fed most on third nymphal instars (the suboptimal one) of the whitefly host as theory predicts, while at high densities, both parasitism and host-feeding occurred on first and second instars which are preferred for oviposition. En. sophia parasitized most on third and fourth (the optimal ones), while fed on first instars (the suboptimal one) at low densities, and utilized third and fourth instars for both at high densities. In conclusion, oviposition vs. host-feeding strategy of parasitoid females was found to vary at different host densities. The balance between reserving optimal hosts for oviposition or using them for host-feeding depended on parasitoid life history and the availability of host resources.     

Parasitism-induced Effects on Host Growth and Metabolic Efficiency in Tobacco Hornworm Larvae Parasitized by Cotesia congregata

Parasitism by the braconid wasp Cotesia congregata affects the growth of Manduca sexta larvae in a parasitoid 'dose-dependent' fashion. Following parasitization of fourth-instar larvae, more heavily parasitized larvae grew larger compared to those containing fewer parasitoids due to an increase in host dry weight. The differences in host mass appeared to arise after oviposition. A 'dose-dependent' enhancement of host dry weight would appear nutritionally beneficial for the parasitoids developing in more 'crowded' hosts. The efficiencies of conversion of ingested and digested food to body mass and the approximate digestibility of the diet ingested by the host caterpillar did not vary significantly with clutch size although parasitoids took slightly longer to develop in the more heavily parasitized hosts. Larval parasitoids developing in the presence of many competitors weighed up to 50% less than those developing in hosts with fewer endoparasitoids, although the weight of adult female parasitoids did not vary significantly with wasp clutch size. The maximum number of emerging wasps was 200 parasitoids, possibly representing the host's 'carrying capacity' for larvae parasitized in the fourth-instar. The ratio of emerging to non-emerging parasitoids decreased as parasitoid clutch size increased, with few or none emerging from very heavily parasitized hosts containing more than 400 parasitoids. Copyright 1997 Elsevier Science Ltd. All right reserved     

Consequences of infanticide for a gregarious ectoparasitoid of leafroller larvae

1. In this laboratory study, the clutch size and handling time of Goniozus jacintae were investigated, a comparison of its life‐history performance between primary and secondary (laid after infanticide events) broods was carried out, and the lipid and protein concentrations in the haemolymph of non‐parasitised and parasitised hosts were estimated. 2. It was found that G. jacintae temporarily paralysed its host larvae for 66 min and briefly guarded its brood for 66 min. The clutch size of G. jacintae increased from two to seven with increasing larval fresh weight of its host, and both ovicide and larvicide of primary clutches occurred in 81% of encounters. 3. Secondary clutches of G. jacintae were significantly larger than primary clutches in two of three ovicide treatments for the same host individuals. Secondary clutches also experienced greater brood survivorship than primary clutches. 4. Lipid concentrations were consistently higher in the haemolymph of parasitised hosts, and protein concentrations were initially higher (until egg hatch), but increased at a lower rate in parasitised hosts than in non‐parasitised hosts. 5. This study is the first to provide evidence that improved nutritional quality could be an important benefit of infanticide for an insect parasitoid, allowing for larger clutch size and improved brood survivorship among secondary broods.     

Nonsiblicidal behavior and the evolution of clutch size in bethylid wasps

Parent-offspring conflict over clutch size may lead to siblicidal behavior between juveniles. In parasitoid wasps, selection for siblicide in small broods is predicted to produce a dearth of gregarious broods with few eggs. Here we document the clutch size distribution in the Bethylidae, a large family of aculeate parasitoids. Small gregarious clutches are the most common. Further data suggest that the most common gregarious clutches in the parasitoid Hymenoptera as a whole contain only a few eggs. Across bethylid species, both clutch size and wasp size increase with host size. Within genera clutch size is more closely related to host size, but between genera or larger clades wasp size is more closely related to host size. The volume of the emerging wasp brood does not depend on whether a species lays single- or multiple-egg clutches once host size is taken into account. These data suggest that clutch size in bethylid wasps is best described by traditional optimality models and that siblicide plays little role in this and possibly other families. We propose several ecological reasons for the rarity of siblicide in bethylids: ectoparasitism, idiobiosis, and a suite of characteristics associated with high within-brood relatedness.     

Host discrimination modulates brood guarding behaviour and the adaptive superparasitism in the parasitoid wasp Trissolcus semistriatus (Hymenoptera: Scelionidae)

Because hosts utilized by parasitoids are vulnerable to further oviposition by conspecifics, host guarding benefits female wasps. The present study aims to test whether female adults regulate brood guarding behaviour by host discrimination in a solitary parasitoid Trissolcus semistriatus by presenting an intact or parasitized host egg mass to a female adult. Virgin females without oviposition experience have host discrimination ability, which enables them to adjust the number of eggs laid in the hosts. Mating experience increases superparasitism by female adults, whereas mated females achieve a higher discrimination ability as a result of oviposition experience and show a lower superparasitism rate. As expected, females exhibit brood guard after parasitizing an intact host egg mass, whereas those females visiting a previously parasitized host egg mass, do not. Because the survival of eggs in superparasitized hosts is relatively low, regulating brood guarding behaviour by host discrimination is adaptive for female wasps.     

Host Handling Time in a Polyembryonic Wasp is Affected both by Previous Experience and by Host State (Parasitized or Not)

Foraging behavior for hosts in parasitoids resembles that of predators with respect to finding, evaluating and manipulating of the prey. Host handling time may depend on the life history of the parasitoid and can be affected by oviposition experience. Additionally, handling time can be affected by host aggregation, species, size and state (parasitized or not). We studied handling times in the egg-larval parasitoid wasp Copidosoma koehleri. We allowed naïve female wasps to oviposit into three consecutive unparasitized hosts, and measured time until oviposition, and the duration of ovipositor insertion. We recorded the same data for naïve females ovipositing into already parasitized hosts. We found that both previous experience by females and previous parasitism of hosts reduced handling time. The results suggest that host handling durations reflect the interplay between host state and parasitoid internal state.