Postemergence learning in the insect parasitoid,Cotesia congregata (Say) (Hymenoptera: Braconidae)

Exposing newly emerged females of Cotesia congregata(Say) to wild cherry, an inherently unattractive plant, and their host larvae at 0–4 h after adult emergence induced a positive searching response to wild cherry and an inhibited response to cabbage, an attractive plant. Inherent responses were not affected when females were exposed to their hosts at 0–12 h and to cherry at 8–12 h after emergence. The induced response to cherry was constant until its disappearance at 6–7 days;inhibition of the response to cabbage was released at 4–5 days after emergence. Postemergence exposure to cherry and parasitoid cocoons induced similar but weaker searching responses. Induced searching responses exhibit features of associative learning and receptor modification. In addition to its presumed role in foraging, postemergence experience with plants may encourage assortative mating of C. congregatawithin suitable host habitats and, thus, facilitate local adaptations to specific plants.     

Behavioral responses to host foodplants of two populations of the insect parasitoid Cotesia congregata (Say)

To test the hypothesis that natural enemy populations differ in their behavioral responses to plants or to plant allelochemicals, we compared two populations of the gregarious larval endoparasitoid, Cotesia congregata (Say) (Hymenoptera: Braconidae) that differed in their historical and present exposure to tobacco. The major hosts for both populations were Manduca sexta L. and M. quinquemaculata (Haworth) (Lepidoptera: Sphingidae), but these hosts were typically encountered on tobacco by parasitoids in one population (Upper Marlboro) and on tomato by parasitoids in another population (Wye). Early in the season, Wye parasitoids preferred to oviposit in M. sexta on tomato rather than on tobacco and Upper Marlboro parasitoids showed no preference; neither population showed any preference later in the season. Neither of the strains originating from the two populations showed a landing preference for tobacco or tomato in flight chamber trials, but Upper Marlboro parasitoids searched longer on tobacco than on tomato, and Wye parasitoids searched longer on tomato. When nicotine solutions were applied to tobacco leaf, searching responses of Upper Marlboro parasitoids were enhanced by 0.001–1.0% nicotine, and searching responses of Wye parasitoids were decreased by 0.01–1.0% nicotine. We speculate that population differences in searching responses to tobacco and nicotine may explain the differential parasitism responses found early in the season.     

Characterization of microsatellite loci in the parasitoid,Cotesia congregata (Say) (Hymenoptera Braconidae)

Nine microsatellite loci were developed and assayed for polymorphism in four groups of the insect parasitoid, Cotesia congregata (Say) (Hymenoptera). One locus was monomorphic and eight were polymorphic in C. congregata. Allelic distributions differed significantly between groups of C. congregata collected from solanaceous and catalpa host food plants (P = 0). Loci were tested for cross‐reactivity to six congeners: C. acuminata (Reinhard), C. bignellii (Marshall), C. cynthiae (Nixon), C. euphydryidis (Musebeck), C. koebelei (Riley) and C. melitaearum (Wilkinson). In the congeners, eight loci were polymorphic, four of which yielded allele sizes that differed substantially from sizes of amplification products observed for C. congregata.     

Cold hardiness of Habrobracon hebetor (Say) (Hymenoptera: Braconidae), a parasitoid of pyralid moths

The ectoparasitoid Habrobracon hebetor (Say) attacks stored-product infesting pyralid moths that are able to overwinter under extremely cold conditions. The extent to which H. hebetor can withstand these conditions is not known, but has important implications for the ability of H. hebetor to provide long-term suppression of these pests in temperate climates. We investigated basic cold hardiness aspects of a mutant eye-color strain of H. hebetor. Feeding larvae and adults of H. hebetor had supercooling points (SCPs) at temperatures higher than those of eggs and pupae. Mean SCPs of females and males were equivalent, as were those of naked and silk-encased pupae. Feeding on honey prior to being subjected to low temperatures significantly increased the SCP of adult females by approximately 8 degrees C. Mortality of pupae and adults increased significantly whenever the temperature dropped below the mean SCP, indicating that H. hebetor does not tolerate freezing. For pupae and adults exposed to -12 and -5 degrees C, the hourly mortality rate increased with time of exposure. Pupae and adults exposed to -12 degrees C for different time intervals showed high mortality after only 1d of exposure. At -5 degrees C, none survived 12d of exposure. A better understanding of how well this parasitoid tolerates low temperatures will be useful in evaluating its potential as a biological control agent of stored-product moths in temperate regions.     

螟黄足盘绒茧蜂复合群支序系统学研究（膜翅目：茧蜂科：小腹茧蜂亚科）

对螟黄足盘绒茧蜂复合群Cotesia flavipes complex (膜翅目：茧蜂科：小腹茧蜂亚科)分布全世界的5个种和外群侧沟茧蜂Microplitis 及荻茧蜂Diolcogaster 的25个性状,以及复合群5个种和外群螟蛉盘绒茧蜂Cotesia ruficrus、粘虫盘绒茧蜂Cotesia kariyai、粉蝶盘绒茧蜂Cotesia glomerata 的24个性状,分别进行比较研究,并运用支序分析的方法探讨该复合群内5个种种间的系统发育关系。支序分析表明螟黄足盘绒茧蜂复合群是一单系群,二化螟盘绒茧蜂C. chilonis 和大螟盘绒茧蜂C. sesamiae 近缘,芦螟盘绒茧蜂C. chiloluteelli 和汉寿盘绒茧蜂C. hanshouensis 近缘,螟黄足盘绒茧蜂C. flavipes相对独立。 以上研究表明无论是以近缘属作外群还是以同属其它种作外群,所得结果基本上都能反映螟黄足盘绒茧蜂复合群各种之间的分类地位。     

Host plants of the herbivorous insect Mythimna separata (Lepidoptera: Noctuidae) affect its susceptibility to parasitism by the larval parasitoid Cotesia kariyai (Hymenoptera: Braconidae)

Previous studies have shown that the food plant species consumed by herbivorous insects affect their parasitism by parasitoid wasps or flies, but this phenomenon has only been observed in a limited number of systems. Here, we investigate how feeding on different plant species affects the survival of the rice armyworm Mythimna separata (Walker) (Lepidoptera: Noctuidae) and its successful parasitism by the larval endoparasitoid Cotesia kariyai (Watanabe) (Hymenoptera: Braconidae). Survival rate and pupal weight of unparasitised M. separata fed with daikon Raphanus sativus L. var. longipinnatus was lower than with maize Zea mays L. In addition, caterpillars fed with daikon were less susceptible to parasitism by C. kariyai than those fed with maize, and parasitised M. separata larvae fed with daikon had higher survival rates than those fed with maize. These results indicate that while daikon is not an optimal food for unparasitised caterpillars, it may protect the larvae from death by C. kariyai parasitism.     

Adult experience modifies attraction of the leafminer parasitoidOpius dissitus (Hymenoptera: Braconidae) to volatile semiochemicals

Oviposition-experienced females of Opius dissitus Muesebeck, a braconid parasitoid of Liriomyza sativaeBlanchard, preferentially landed on leafminer-infested rather than uninfested lima bean (Phaseolus lunatus L.) plants in a flight tunnel assay. Both naive and oviposition-experiencedparasitoids responded strongly to odors of infested lima bean plants in a four-arm olfactometer in comparison with odors of uninfested plants, suggesting that volatile semiochemicals are used in host location. Parasitoids with an oviposition experience on lima bean (lima-experienced) spent significantly more time in the infested odor than naive individuals, however, eggplant-experienced wasps did not spend significantly more time in the infested odor field than naive wasps. When parasitoids reared on leafminers in lima bean were provided a choice between the odor of infested lima bean and the odor of infested eggplant or cotton, naive and lima-experienced wasps preferred infested lima odor. An oviposition experience on the other plant species resulted in a dramatic shift in preference. It was concluded that the experience effect was due, at least in part, to associative learning, as has been reported for other parasitoids. The parasitoids may perceive unconditioned stimuli during host contact and oviposition on an infested leaf and may associate those stimuli with volatile semiochemicals emanating from the leaf or host. Subsequently, the volatiles associated with the presence of hosts are used in directing the search for hosts.     

When good bugs go bad: intraguild predation by Jalysus wickhami on the parasitoid, Cotesia congregata

We report a case of direct intraguild predation involving an insect predator and parasitoid in an agricultural system. The spined stilt bug, Jalysus wickhami Van Duzee, feeds on eggs of the tobacco hornworm, Manduca sexta L., and also on prepupal and pupal stages of the gregarious hornworm parasitoid, Cotesia congregata (Say). In two separate trials, mean mortality of attached parasitoids was significantly lower (66%, 73%) than that of their detached siblings (97%, 96%) after a 3 day exposure to stilt bugs, demonstrating that attachment to the host offered some protection against predation. In no-choice experiments, prepupal parasitoids suffered greater mortality (0 day-old=61%, 1 day-old=65%) than pupal parasitoids (2 day-old=50%, 3 day-old=14%). When offered in combination with 0 or 2 day-old hornworm eggs, respective mortality of 0, 1, 2 and 3-day-old pupal parasitoids showed a similar pattern (67%, 63%, 33% and 23%). In another experiment, mortality of 0-day-old pupal parasitoids (64%) was greater than that of 3 day-old pupal parasitoids (38%). Mortality of pupal parasitoids was not affected by the availability of hornworm eggs, a highly acceptable food. Younger pupal parasitoids (=prepupae) probably suffered greater mortality because they were more easily fed on by stily bugs than older (pupated) ones. Because C. congregata overwinters in the prepupal stage, it may be particularly vulnerable to attack late in the season when stilt bug populations are large and hornworm eggs are relatively uncommon.     

Effects of the polydnavirus of Cotesia congregata on the immune system and development of non-habitual hosts of the parasitoid

Polydnaviruses (PDV) are obligate mutualistic symbionts found in association with some groups of parasitic Hymenoptera. In these groups, they suppress the immune response of the parasitoid’s host and are required for successful parasitoid reproduction. Several PDV effects have been described in different experimental systems, but no clear picture of PDV mode of immunosuppression has emerged. No study to date has directly tested if PDV modes of action are evolutionarily conserved or divergent among parasitoid taxa within the Ichneumonoidea. We hypothesize the divergence in PDV mode of immunosuppression can be detected by identifying points of divergence in the immune response of different host species to PDV from one parasitoid species. This study tests the effects of purified PDV from Cotesia congregata on the immune response of three larval lepidopteran species that naturally are hosts of parasitoid species that differ in taxonomic relatedness to C. congregata. Here we demonstrate that despite associations with distantly related parasitoids (Ichneumonidae and Braconidae), Manduca sexta and Heliothis virescens showed similar patterns of increased glucose dehydrogenase (GLD) activity, suppressed cellular encapsulation in vitro, and increased time to pupation. In contrast, Lymantria dispar showed no response to C. congregata PDV across any of the parameters measured, even though it has an evolutionary association with several parasitoids closely related to C. congregata and within the Microgastrinae. The PDV immunosuppression in H. virescens and M. sexta does not correlate with host molecular phylogeny either. The suborganismal effects shown in M. sexta and H. virescens translated into significantly reduced pupation success in M. sexta only. Results demonstrate that while some PDV modes of immunosuppression in hosts may be divergent, others may be conserved across broad host groups.     

Courtship and intersexual signaling in the parasitic waspCotesia rubecula (Hymenoptera: Braconidae)

The courtship behavior of the parasitic wasp Cotesia rubeculawas studied in a flight tunnel using standard quantitative ethological techniques. Emission of a female sex pheromone induces searching and signaling behavior in males. Males combine wing-fanning, which produces low-frequency airborne sound, with pulsing behavior, which transmits a vibrational signal through the substrate to the female and induces her receptivity. Female receptivity is indicated by a stereotyped antennal position, which may provide a visual or tactile signal to courting males. Comparison of successful and unsuccessful courtships indicated that courtship success was dependent primarily on the effective production or reception of the male pulse signal. Overall, the sequence of courtship behavior was similar to that reported for other parasitic wasps.     

The role of volatiles from cruciferous plants and pre-flight experience in the foraging behaviour of the specialist parasitoid Cotesia plutellae

The braconid Cotesia plutellae is an important larval parasitoid of the diamondback moth (Plutella xylostella), a major pest of crucifers in the tropics and sub-tropics. The in-flight searching behaviour of C. plutellae was investigated in a wind tunnel and the close-range attack behaviour observed in cages. The relative importance of volatile stimuli emanating from the plant-host-complex, oilseed rape (Brassica napus) – P. xylostella, in the long-range attraction of C. plutellae was investigated. Plants that were mechanically damaged, or damaged by P. xylostella larvae, were attractive to the parasitoid. Host-damaged leaves remained attractive to the parasitoid after removal of the host larvae. These results indicate that C. plutellae predominantly uses plant derived stimuli in its in-flight searching behaviour. An oviposition experience or contact with a host-damaged leaf prior to the bioassay significantly increased the response to these volatile cues. The foraging behaviour of C. plutellae is compared with other braconid larval parasitoids attacking lepidopteran hosts on crucifers.     

Influence of dietary nicotine and colony source ofManduca sexta [Lepidoptera: Sphingidae] on its suitability as a host ofCotesia congregata [Hymenoptera: Braconidae]

Larval tobacco hornworms,Manduca sexta (L.), of 2 different colonies were exposed to parasitism by the gregarious endoparasitoid,Cotesia congregata (Say). A comparison was made of parasitoid larval, pre-pupal, and pupal mortality, female and male dry weight and larval development time. In general, “Maryland” hornworms were more suitable hosts than “North Carolina” hornworms. Although the presence of dietary nicotine increased parasitoid mortality in individuals reared from hornworms of both colonies, the effect was more severe among individuals parasitizing the North Carolina hornworms. Scientific contribution No. 8125, article No. A-5066 of the Maryland Agricultural Experiment Station, Department of Entomology.     

外源茉莉酸诱导植物反应对菜蛾绒茧蜂寄生选择行为的影响

茉莉酸是植物体内重要的伤信号分子,向植物施用外源茉莉酸后, 可诱导植物产生各种防卫反应, 如挥发物组成发生改变等, 进而影响植食性昆虫及其天敌。该文报道用不同浓度外源茉莉酸处理白菜和甘蓝后,诱导植物反应所产生的挥发物对菜蛾绒茧蜂搜索及寄生选择行为的影响。外源茉莉酸处理白菜和甘蓝后,处理植株的挥发物对菜蛾绒茧蜂的引诱力增强;与在对照植株上相比,该蜂对经茉莉酸处理后白菜植株上的小菜蛾幼虫的寄生数显著要高。表明茉莉酸处理白菜及甘蓝后,植物诱导反应导致其挥发物的作用发生变化,进而可提高该蜂的搜索和寄生效率。     

Foraging for solitarily and gregariously feeding caterpillars: A comparison of two related parasitoid species (Hymenoptera: Braconidae)

In the present study we apply a comparative approach, in combination with experimentation, to study behavior of two parasitoid species that attack caterpillar hosts with different feeding strategies (gregarious or solitary). In a semifield setup, consisting of clean cabbage plants and plants infested with one of two host species, the foraging behavior of the specialistCotesia rubecula, on obligate parasitoid of solitarily feedingPieris rapae larvae, was compared to that of the generalistCotesia glomerata, a polyphagous parasitoid of several Pieridae species (mainly the gregariously feedingPieris brassicae).Cotesia glomerata displayed equal propensity to search for and parasitize larvae of both host species. AlthoughC. glomerata exhibited a relatively plastic foraging behavior in that it searched differently under different host distribution conditions, its behavior seems more adapted to search for gregariously feeding hosts. Females exhibited a clear area-restricted search pattern and were more successful in finding the gregariously feeding caterpillars.Cotesia rubecula showed a higher propensity to search forP. rapae than forP. brassicae, i.e., females left the foraging setup significantly earlier when their natural hostP. rapae was not present.C. rubecula showed a more fixed foraging behavior, which seems adapted to foraging for solitarily feeding host larvae. In a setup with onlyP. rapae larvae, the foraging strategies of the two parasitoid species were quite similar. In a choice situationC. glomerata did not show a preference for one of the host species, whileCotesia rubecula showed a clear preference for its natural host species. The latter was shown by several behavioral parameters such as the number of first landings, allocation of search time, and percentage parasitization.     

中国并腹茧蜂属一新种记述（膜翅目：茧蜂科）

记述采自中国黑龙江的并腹茧蜂属Hylcalosia Fischer 1新种——沟腹并腹茧蜂H.ventisulcata sp.nov..该新种与海氏并腹茧蜂Hylcalosia hymaenei Belokobylskij最为相似,但新种的如下特征容易与后者相区别:1)下颚须相当长,可达后足基节处;2)腹部第2-3节背板具1条宽而浅的中纵沟,几乎纵贯2、3背板;3)前翅2-SR脉基部弯曲,后翅m-cu脉对叉式;4)腹部第1背板中长等于端宽。文中修订了该属全世界已知种的检索表。新种模式标本保存于福建农林大学益虫研究所。     

Effects of experience on food-searching behavior in the antFormica schaufussi (Hymenoptera: Formicidae)

Foragers of the ant Formica schaufussitend to return to and search at a site of a previous food find. The search tactic employed by a forager on its return trip is related to the type of food previously encountered: search is more persistent in response to carbohydrate than to protein food. Using different reinforcement schedules with carbohydrate and protein food rewards, we show that, on a short-term as well as on a long-term basis, the basic pattern of search observed in naive foragers is only slightly modified through foraging experience. Foragers do not increase their search effort or adjust their search pattern when either type of food is systematically renewed on a fixed reward schedule and thus do not seem to be able to learn to assess the food predictability. Collective responses that could compensate for this lack of individual flexibility and increase foraging efficiency at the colony level are discussed.     

菜蛾盘绒茧蜂卵携带的免疫抑制因子

抑制寄主昆虫的免疫反应是内寄生蜂存活的关键。菜蛾盘绒茧蜂Cotesia vestalis（Haliday）寄生小菜蛾Plutella xylostella （L.）幼虫后,蜂卵如何逃避和抑制寄主的免疫攻击,尚未得到全面揭示。本文采用电镜技术系统观察了菜蛾盘绒茧蜂卵表面的超微结构。结果显示：蜂卵表面覆盖有纤维层和絮状的类病毒样纤丝（VLFs）,同时携带了含多分DNA病毒粒子（PDV）的萼液。在寄生初期,包裹在蜂卵表面的纤维层和VLFs首先起到保护蜂卵不被小菜蛾血细胞包囊的被动防御作用。随后,PDV发挥主动的免疫抑制作用。通过假寄生手段,证明了菜蛾盘绒茧蜂PDV (CvBV) 具有较持久的克服寄主免疫攻击的能力,是主要的免疫抑制因子。在假寄生后连续8 d的观察时间内,菜蛾盘绒茧蜂的蜂卵均未被包囊。结果提示,在菜蛾盘绒茧蜂-小菜蛾寄生体系中,菜蛾盘绒茧蜂采取被动防御和主动攻击两种方式应对寄主小菜蛾的免疫攻击。     

Seasonal changes of resistance to fenvalerate and cypermethrin in the larval parasitoid Cotesia plutellae （Hymenoptera： Braconidae）

The seasonal changes of insecticide resistance and stability in hymenopteran Cotesia plutellae, collected from Jianxin, Fuzhou-City, and Shangjie, Minhou-County, Fujian, China, were assessed by using a dry residual film method. The resistance to two insecticides in the field populations of C. plutellae was not stable under insecticide-free conditions in the insectarium. Compared with susceptible F11 progeny of C. plutellae in the insectarium, the resistance ratios （RR） in F0 parents were 18.4 for fenvalerate and 11.4 for cypermethrin based on LC50 at 9 hours, and 32.8 for fenvalerate and 28.5 for cypermethrin based on LC50 at 24 hours when the parasitoids were left in contact with the insecticides for 1 hour and mortalities were recorded at 9 and 24 hours, respectively. However, the RR in a field population of C. plutellae were 9.2 for fenvalerate and 12.7 for cypermethrin, if the parasitoids were left in contact with the insecticides for 24 hours. The resistances to the two pyrethroids in other field populations collected from Jianxin and Shangjie from November 2000 and July 2004 were also determined. Significant seasonal variations of resistance to the two insecticides in the field populations of C. plutellae were found. The RR were 3.0-18.4 for fenvalerate and 4.8-20.6 for cypermethrin in Jianxin populations from November 2000 to April 2002 based on LC50 at 9 h, and 2.3-13.6 for fenvalerate and 3.6-16.0 for cypermethrin in Shangjie populations from May 2002 to July 2004 based on LC50 at 24 hours. The resistance levels were high in spring and autumn and decreased sharply in summer. In addition, significant recovery from the knocked-down caused by the insecticides was found in the F0 and field populations of C. plutellae which were resistant to fenvalerate and cypermethrin if the parasitoids were left in contact with the pyrethroids for 1 hour. However, no recovery was found in susceptible F11 progeny.     

Genetic Variation in Behavioral Response to Herbivore-Infested Plants in the Parasitic Wasp, Cotesia glomerata (L.) (Hymenoptera: Braconidae)

Female Cotesia glomerata (L.) relies on stimuli from herbivore-infested plants to select suitable hosts, but behavioral response to such stimuli is highly variable among individuals. This study investigates a genetic component of phenotypic variability in both short-range host-search and long-range host-location behaviors in the tritrophic system consisting of cabbage plants (Brassica oleracea L.), cabbage butterfly (Pieris brassicae L.) and the parasitoid, by comparing full-sib families established from a laboratory population and isofemale strains from a field population. Short-range host-search behaviors were examined within a Petri-dish test arena, and long-range host-location behaviors assessed in a wind tunnel. Significant differences among full-sib families were shown in the duration of walking on a plant-herbivore complex (i.e., a leaf section with two host caterpillars, their silk and feces) and searching off the complex, and the total time elapsed for wasps to locate a host larva after release into the test arena. Flight responses to and landing choices between the intact and the herbivore-infested plants were also significantly different among these families. Effects of families on both short-range host-search and long-range host-location behaviors were consistent, without significant influences of host larvae from which wasps emerged. The analysis of isofemale strains reveals that strains account for significant variation in the oriented flight response to herbivore-infested plants, and the isofemale heritability for this behavioral character is estimated as 0.447. The results suggest that genetic variation exists at different behavioral levels of the host-selection process in this parasitoid.