Phylogeny of the parasitic wasp subfamily Charipinae (Hymenoptera, Cynipoidea, Figitidae)

The Charipinae are a major group of hyperparasitoids of Hemiptera. Here, we present the first cladistic analysis of this subfamily's internal relationships, based on 96 morphological characters of adults. The data matrix was analysed using uniformly weighted parsimony. The effects of using alternative weighting schemes were explored by performing additional searches employing implied weights criteria. One of the caveats of implied weights analysis is that it lacks an objective criterion for selecting the value of the concavity function. In the present study, differential weighting was used to explore the sensitivity of our results to the alternative assumptions made in the analysis and to select one of the most parsimonious trees under equal weights, which we regard as being the hypothesis that minimizes the amount of ad hoc assumptions. The validity of the two existing tribes and the monophyly of all the genera of Charipinae were tested, in particular the cosmopolitan and highly species-rich Alloxysta and Phaenoglyphis , which appear repeatedly in ecological and biochemical studies of host–parasitoid associations. The evolution of several major characters and the relationships between genera are discussed. On the basis of the phylogenetic results, we discuss a number of taxonomic issues. A new classification of the subfamily is proposed in which no tribes are maintained, Carvercharips is synonymyzed with Alloxysta , and the creation of a new genus from Nepal is justified. Our analysis points to the need for a world revision of the basal genus Phaenoglyphis , which is shown as paraphyletic.     

Phylogeny of the genera of the parasitic wasps subfamily Doryctinae (Hymenoptera: Braconidae) based on morphological evidence

The evolutionary relationships among most (143 genera) of the currently recognized genera of the braconid wasp subfamily Doryctinae were investigated using maximum parsimony analysis, employing 100 characters from external morphology and four additional, less well‐studied character systems (male genitalia, ovipositor structure, venom apparatus and larval cephalic structure). We investigated the ‘performance’ of characters from external morphology and the other character systems and the effects of abundant missing entries by comparing the data decisiveness, retention and consistency indices of four different character partitions. The results indicate that the performances of the different partitions are not related to the proportions of missing entries, but instead are negatively correlated to their proportion of informative characters, suggesting that the morphological information in this group is subject to high levels of homoplasy. The external morphological partition is significantly incongruent with respect to a data set comprising the other character systems based on the ILD test. Analyses supported neither the monophyly of the large tribes Doryctini and Hecabolini, nor the monophyly of the Spathiini and Heterospilini. Relationships obtained from successive approximation weighting analysis for the complete data differ considerably from the currently accepted tribal and subtribal classifications. The only exceptions were the Ypsistocerini and the Ecphylini, whose recognized members were recovered in single clades. A close relationship between the Binaerini and Holcobraconini, and also Monarea, is consistently supported by venom apparatus and ovipositor structure characters but is not indicated by external morphological data. Low bootstrap values obtained for most of the recovered clades in all analyses do not allow us to propose a meaningful reclassification for the group at this time. A complete list of the recognized genera and their synonymies is given. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 369–404.     

Phylogenomics of the lepidopteran endoparasitoid wasp subfamily Rogadinae (Hymenoptera: Braconidae) and related subfamilies

Rogadinae are a cosmopolitan, species‐rich braconid wasp subfamily whose species are endoparasitoids that attack larvae of a number of lepidopteran families. Members of this subfamily are characterized by pupating within the mummified host larval skin. The subfamily contains six tribes whose relationships have only been partially clarified: Aleiodini, Betylobraconini, Clinocentrini, Rogadini, Stiropiini and Yeliconini. The limits and composition of the closely related subfamilies to the Rogadinae, Hormiinae and Lysiterminae, also remain unclear. Here, we generated ultraconserved element data to reconstruct an almost fully resolved phylogeny for the members of Rogadinae and related subfamilies. Based on our best estimate of phylogeny, we confirm the monophyly of Rogadinae including Betylobraconini, synonymize Xenolobus Fahringer and Bequartia Cameron within the species‐rich genus Aleiodes Wesmael ( syn.n. ) based on DNA, and synonymize Promesocentrus van Achterberg with Pilichremylus Belokobylskij ( syn.n. ) based on morphology. We also consistently recovered Hormiinae and Lysiterminae as not reciprocally monophyletic, and thus propose to unite their members under Hormiinae. The ancestral host preference for Rogadinae was probably attacking concealed lepidopteran larvae, with the occurrence of at least two main subsequent transitions to attack both concealed and exposed hosts, one within Rogadini and a second within Aleiodini. We highlight the importance of natural history collections as a source for conducting genomic‐based studies using techniques that allow to obtain a substantial amount of data from considerably old preserved insect specimens.     

Polymorphic microsatellite loci in the parasitic wasp Cotesia glomerata (Hymenoptera: Braconidae)

Five polymorphic microsatellite loci were isolated in a gregarious endoparasitoid, Cotesia glomerata, using an enrichment protocol. We detected two to 15 alleles per locus in a sample of 50 female wasps collected from a field population near Zürich, Switzerland. The observed heterozygosities ranged from 0.21 to 0.92. These markers are currently being used in our studies of sex determination in this species.     

Systematics and evolution of gall formation in the plant‐associated genera of the wasp subfamily Doryctinae (Hymenoptera: Braconidae)

Gall formation is a specialised form of phytophagy that consists of abnormal growth of host plant tissue induced by other organisms, principally insects and mites. In the mainly parasitoid wasp subfamily Doryctinae, gall association, represented by gall inducers, inquilines and their parasitoids, is known for species of seven genera. Previous molecular studies recovered few species of six of these genera as monophyletic despite their disparate morphologies. Here, we reconstructed the evolutionary relationships among 47 species belonging to six gall‐associated doryctine genera based on two mitochondrial and two nuclear gene markers. Most of the Bayesian analyses, performed with different levels of incomplete taxa and characters, supported the monophyly of gall‐associated doryctines, with Heterospilus (Heterospilini) as sister group. Percnobracon Kieffer and Jörgensen and Monitoriella Hedqvist were consistently recovered as monophyletic, and the validity of the monotypic Mononeuron was confirmed with respect to Allorhogas Gahan. A nonmonophyletic Allorhogas was recovered, although without significant support. The relationships obtained and the gathered morphological and biological information led us to erect three new genera originally assigned to Psenobolus: Ficobolus gen.n. (F. paniaguai sp.n. and F. jaliscoi sp.n. ), Plesiopsenobolus gen.n. (Pl. mesoamericanus sp.n. , Pl. plesiomorphus van Achterberg and Marsh comb.n. , and Pl. tico sp.n. ), and Sabinita gen.n. (S. mexicana sp.n. ). The origin of the gall‐associated doryctine clade was estimated to have occurred during the middle Miocene to early Oligocene, 16.33–30.55 Ma. Our results support the origin of true gall induction in the Doryctinae from parasitoidism of other gall‐forming insects. Moreover, adaptations to attack different gall‐forming taxa on various unrelated plant families probably triggered species diversification in the main Allorhogas clade and may also have promoted the independent origin of gall formation on at least three plant groups. Species diversification in the remaining doryctine taxa was probably a result of host shifts within a particular plant taxon and shifts to different plant organs. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:0021F253‐4ABA‐4EAA‐A7A9‐FC0AD1932EA3     

A new rod-shaped virus from parasitic wasp Diachasmimorpha longicaudata (Hymenoptera: Braconidae)

A new rod-shaped nucleocapsids (NCs) was found inadvertently in Diachasmimorpha longicaudata accessory gland filaments (AGFs). The NCs were 30 nm in diameter and nearly 900 nm in length. They replicated in a small cell type of the AGFs in D. longicaudata, and following oviposition, invaded and proliferated in the hemocytes of a parasitized host Bactrocera dorsalis Hendel. This finding of a completely new virus in the AGF indicate that different geographical populations (subspecies) of D. longicaudata may carry different sybionts. This is the first report showing that the same wasp species, but from a different geographical populations, can carry an entirely different virus.     

红脉穗螟寄生蜂寄主选择研究

[目的]优化红脉穗螟寄生蜂繁育技术,为该虫的生物防治提供参考。[方法]通过室内饲养研究了红脉穗螟幼虫寄生蜂褐带卷蛾茧蜂和蛹寄生蜂周氏啮小蜂(海南种)对4种昆虫的寄生效果,并对最佳接种比例进行筛选。[结果]褐带卷蛾茧蜂对大蜡螟幼虫寄生效果优于米蛾幼虫,且对黄粉虫和大麦虫幼虫不表现寄生特性,褐带卷蛾茧蜂和大蜡螟幼虫的最佳接种比例为1∶1和2∶1,其中在2∶1接种比例处理中的寄生率和单寄主产蜂量分别为76.67%和34.60头;4种昆虫蛹均可用于繁育周氏啮小蜂,从寄主的繁育成本和寄生效果分析,以黄粉虫蛹效果较最佳,接种蜂虫比以2∶1为宜,此时的单寄主产蜂量为148.60头。[结论]寄主和接种比例不同会影响寄生蜂的寄生效果,本研究中褐带卷蛾茧蜂适宜寄主为大蜡螟,最优蜂虫比为1∶1和2∶1,周氏啮小蜂(海南种)适宜寄主为黄粉虫,最优接种蜂虫比为2∶1。     

Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa

Zaldivar‐River ó n, A., Areekul, B., Shaw, M. R. & Quicke, D. L. J. (2004). Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa. —Zoologica Scripta, 33, 223–237. The morphology of the venom apparatus intima in representatives of 38 genera of the problematic braconid wasp subfamily Rogadinae and other cyclostome braconids was investigated and a preliminary phylogenetic analysis for the group was performed with the information obtained. Despite the limited number of characters, the data suggest several relationships at various taxonomic levels. The venom apparatus in the Clinocentrini and the Stiropiini is relatively unmodified and similar to that found in other genera previously placed within a broader concept of the Rogadinae (e.g. genera of Lysitermini, Pentatermini, Tetratermini, Hormiini) and also to that of the Betylobraconinae. The presence of a cone of filaments located inside the secondary venom duct near to its insertion on the venom reservoir/primary venom duct is proposed as a synapomorphy for the tribe Rogadini to the exclusion of Stiropiini, Clinocentrini and Yeliconini. Other features of the secondary venom duct and its insertion on the venom reservoir/primary venom duct support a number of relationships between the genera of the Rogadini and also within the large genus Aleiodes. A clade containing 15 Rogadini genera (Bathoteca, Bathotecoides, Bulborogas, Canalirogas, Colastomion, Conspinaria, Cystomastacoides, Macrostomion, Megarhogas, Myocron, Pholichora, Rectivena, Rogas, Spinaria and Triraphis) is supported by the presence of a thickened and short secondary venom duct, whereas the different members of Aleiodes (excluding members of the subgenus Heterogamus) and Cordylorhogas are distinguished by having a recessed secondary venom duct with well‐defined and numerous internal filaments. New World Rogas species exhibit a unique venom apparatus and may not be closely related to the Old World ones. Features of the venom apparatus of the enigmatic genus Telengaia and the exothecine genera Shawiana and Colastes suggest that the Telengainae and Exothecinae are both closely related to the Braconinae, Gnamptodontinae, and possibly to the Opiinae and Alysiinae. An unsculptured venom reservoir was found in one specimen of the type species of Avga, A. choaspes, which is consistent with it occupying either a very basal position within the cyclostome braconids or belonging to a recently recognized ‘Gondwanan’ clade that also includes the Aphidiinae.     

Phylogeny of bird-grasshopper subfamily Cyrtacanthacridinae (Orthoptera: Acrididae) and the evolution of locust phase polyphenism

Locust phase polyphenism is an extreme form of density-dependent phenotypic plasticity in which solitary and cryptic grasshoppers can transform into gregarious and conspicuous locusts in response to an increase in local population density. We investigated the evolution of this complex phenotypic plasticity in a phylogenetic framework using a morphological phylogeny of Cyrtacanthacridinae, which contains some of the most important locust species, and a comprehensive literature review on the biology and ecology of all known members of the subfamily. A phylogenetic analysis based on 71 morphological characters yielded a well-resolved tree and found that locust phase polyphenism evolved multiple times within the subfamily. The literature review demonstrated that many cyrtacanthacridine species, both locust and sedentary, are capable of expressing density-dependent color plasticity. When this color plasticity was divided into two smaller components, background coloration and development of black pigmentation, and when these plastic traits were optimized on to the phylogeny, we found that the physiological mechanisms underlying this plasticity were plesiomorphic for the subfamily. We also found that different locust species in Cyrtacanthacridinae express both similarities and differences in their locust phase polyphenism. Because locust phase polyphenism is a complex syndrome consisting of numerous plastic traits, we treat it as a composite character and dissected it into smaller components. The similarities among locust species could be attributed to shared ancestry and the differences could be attributed to the certain components of locust phase polyphenism evolving at different rates.
© The Willi Hennig Society 2007.     

与中红侧沟茧蜂生境与寄主定位相关的玉米及棉铃虫幼虫体表挥发性成分的提取与鉴定

【目的】鉴定玉米Zea mays L.和棉铃虫Helicoverpa armigera (Hübner)幼虫体表挥发性成分中对中红侧沟茧蜂 Microplitis mediator (Haliday)具有生境与寄主定位作用的信息化合物,从化学生态学的角度研究玉米-棉铃虫-中红侧沟茧蜂三重营养关系,解释中红侧沟茧蜂寻找寄主的过程中的信息识别机制,为害虫的综合防治的“推-拉”方法提供一定的理论基础。【方法】利用触角电位仪(EAG)、触角电位联用仪(GC-EAD)、气质联用仪(GC-MS)及“Y”型嗅觉仪确定玉米和棉铃虫幼虫体表提取物的信息化合物。在室内利用玉米以及棉铃虫幼虫体表挥发物标准品化合物以及模拟混合物,使用“Y”型嗅觉仪进行中红侧沟茧蜂成虫行为反应试验。【结果】玉米挥发物中有11种化学成分,棉铃虫幼虫体表挥发物中有6种化学成分对中红侧沟茧蜂的触角具有电生理活性,其中4种成分在两种挥发物中都存在。室内行为反应试验发现：与正己烷对照相比,玉米的模拟组分对雌、雄蜂均表现出显著（P<0.05）的诱引作用;棉铃虫1龄幼虫体表模拟组分对雌蜂具有极显著的诱引作用（P<0.01）,对雄蜂具有显著的诱引作用（P<0.05）;棉铃虫2龄幼虫体表模拟组分对雌蜂具有显著的诱引作用（P<0.05）。【结论】本研究证明了玉米以及棉铃虫幼虫体表挥发物中分别存在11种（庚醛、2-己醇、1-己醇、1-辛烯-3-醇、壬醛、葵醛、苯甲醛、反式-2-壬烯-1-醇、己酸、苯基乙醇、月桂醇）和6种（2-己醇、己酸乙酯、1-己醇、壬醛、辛酸乙酯、癸醛）中红侧沟茧蜂生境及寄主定位的化学信息物质。     

Effects of mating on the oviposition behaviour of the parasitic wasp,Apanteles glomeratus L. (Hymenoptera: Braconidae)

The oviposition behaviour of mated or virgin females of the parasitic wasp,Apanteles glomeratus L., was investigated. Virgin females laid fewer eggs in a shorter time than did mated females (P<0.01), though the attack ratio and attack time of these 2 female types were the same. Progeny sex ratios of mated females suggested the occurrence of sperm shortage, because old mated females produced clutches of a high male proportion, sometimes consisting of males only. Recopulation of females was also observed, but actual insemination did not occur, and therefore,A. glomeratus females are considered to be fundamentally monogamous.     

Body temperature of the parasitic wasp Pimpla turionellae (Hymenoptera) during host location by vibrational sounding

Abstract.  The pupal parasitoid Pimpla turionellae (L.) uses self-produced vibrations transmitted on the plant substrate, so-called vibrational sounding, to locate immobile concealed pupal hosts. The wasps are able to use vibrational sounding reliably over a broad range of ambient temperatures and even show an increased signal frequency and intensity at low temperatures. The present study investigates how control of body temperature in the wasps by endothermic mechanisms may facilitate host location under changing thermal environments. Insect body temperature is measured with real-time IR thermography on plant-stem models at temperature treatments of 10, 18, 26 and 30 °C, whereas behaviour is recorded with respect to vibrational host location. The results reveal a low-level endothermy that likely interferes with vibrational sound production because it occurs only in nonsearching females. At the lowest temperature of 10 °C, the thoracic temperature is 1.15 °C warmer than the ambient surface temperature whereas, at the high temperatures of 26 and 30 ° C, the wasps cool down their thorax by 0.29 and 0.47 °C, respectively, and their head by 0.45 and 0.61 °C below ambient surface temperature. By contrast, regardless of ambient temperature, searching females always have a slightly elevated body temperature of at most 0.30 °C above the ambient surface temperature. Behavioural observations indicate that searching females interrupt host location more frequently at suboptimal temperatures, presumably due to the requirements of thermoregulation. It is assumed that both mechanisms, producing vibrations for host location and low-level endothermy, are located in the thorax. Endothermy by thoracic muscle work probably disturbs signal structure of vibrational sounding, so the processes cannot be used at the same time.     

Hagen's glands of the parasitic wasp Diachasmimorpha longicaudata (Hymenoptera: Braconidae): ultrastructure and the detection of entomopoxvirus and parasitism-specific proteins

Hagen's glands of males of the parasitic wasp Diachasmimorpha longicaudata (Hymenoptera: Braconidae) secrete compounds that are involved in courtship and defense. Like the poison glands of female wasps, the Hagen's glands are secretory, membranous, and of ectodermal origin. The poison glands contain the symbiotic entomopoxvirus, DlEPV and the parasitism-specific protein, PSP 24. DlEPV proteins were detected in homogenates of male wasps. Our goal was to describe the ultrastructure of the Hagen's glands and determine whether they contain DlEPV virions and/or proteins as well as PSP 24. The Hagen's glands are bilateral and each consists of 12-16 tubules arranged in two clusters. In cross-section, a tubule has three zones that enclose a central cuticle-lined lumen. The outermost zone consists of aggregates ('islands') of small vesicles, interconnected by narrow ductules that lead to large cuticle-lined ducts, which transport electron-dense material to the lumen prior to its release from the gland. Large vesicles in Zone 2 and a thick layer of ribosomes and rough endoplasmic reticula in Zone 3 are the likely sites of storage and protein synthesis, respectively. While DlEPV virions were not seen in the Hagen's gland, DlEPV and PSP 24 proteins were present.