Relationships among the tiphiid wasp subfamilies (Hymenoptera)

Abstract. The phylogenetic relationships among the subfamilies of Tiphiidae are discussed, with detailed analysis of characters, and presentation of a phylogenetic tree. The tribe Diammini Turner is elevated to subfamily status. A key to the subfamilies is included, and a brief synopsis of each subfamily is given.     

Functional morphology of the sting apparatus of the spider wasp Cryptocheilus versicolor (Scopoli, 1763) (Hymenoptera: Pompilidae)

The females of the spider wasps (Hymenoptera: Pompilidae) hunt spiders to provision their larvae. The genital structures of pompilid females are modified in a sting that is used for paralyzing the prey (spiders) and defense. The skeleto‐muscular structure of the sting apparatus of a typical representative of the family (Cryptocheilus versicolor) is examined. The shape of sclerites, their relative positions and articulations are described. Some morphological adaptations are described for the first time. The wide anal arc of the tergum IX provides a stiff support for the muscles that move the valvulae. The resilin structures in the areas of articulation support the work of muscles and in some cases replace them. The 1st valvulae form a venom duct along their entire length, which provides the delivery of the venom to a specific point. An unpaired flap in the venom duct provides a dose of venom in the sting. This mechanism probably enhances the speed and accuracy of the wasp's sting movements. Functions of muscles and interactions of the structures of the sting apparatus of C. versicolor are discussed.     

Behavioural notes on the Neotropical parasocial spider wasp Ageniella (Lissagenia) flavipennis (Banks) (Hymenoptera: Pompilidae), with host association

1. Ageniella is a species‐rich group of spider wasps restricted to the New World. Knowledge regarding the behaviour of this genus is based mainly on the Nearctic species, which have been reported to nest solitarily in the soil. This study reports for the first time the nesting behaviour, with host association of Ageniella (Lissagenia) flavipennis Banks. 2. Behavioural aspects on the nesting of A. flavipennis were observed from a study of six nests found in an Atlantic Forest conservation area in São Paulo State, Brazil. Host specimens were collected from a nest, as well as while being carried by an A. flavipennis individual. 3. The present study reports the A. flavipennis females cohabiting or nesting solitarily in mud nests, indicating that this spider wasp shows some lower level of parasociality. In addition, the spider Enoploctenus cyclothorax (Bertkau) was reported for the first time as host. As has been observed for other Ageniellini, females of A. flavipennis amputate the host's legs and transport the spider to the nest, flying or walking forward. 4. Communal behaviour has been reported for species of different genera of Pompilidae, such as Macromeris Lepeletier, Paragenia Bingham and Auplopus Spinola. Recent phylogenetic analyses suggest that the Lissagenia species are more closely related to the other Ageniellini genera than to the other Ageniella species. The present information on nesting and prey could contribute towards a more conclusive phylogenetic position of Lissagenia.     

New records of spider wasps (Hymenoptera,Pompilidae) from Colombia

New records of genera and species of spider wasps (Hymenoptera: Pompilidae) from Colombia are provided. Agenioideus, Cryptocheilus, Evagetes, Mystacagenia, and Xerochares are newly recorded genera from Colombia. Nineteen species are first recorded from Colombia: Aimatocare vitrea (Fox); Ageniella azteca (Cameron); Ageniella curtipinus (Cameron); Ageniella fallax (Arlé); Ageniella hirsuta Banks; Ageniella pilifrons (Cameron); Ageniella pretiosa Banks; Ageniella sanguinolenta (Smith); Ageniella zeteki (Banks); Agenioideus birkmanni (Banks); Aporus (Aporus) cuzco Evans; Aporus (Cosmiaporus) diverticulus (Fox); Aporus (Notoplaniceps) canescens Smith; Euplaniceps exilis (Banks); Euplaniceps herbertii (Fox); Irenangelus clarus Evans; Mystacagenia bellula Evans; Phanochilus nobilitatus (Smith) and Xerochares expulsus Schulz. The following species and genera have their occurence ranges expanded for South America: Ageniella azteca (Cameron); Ageniella zeteki (Banks); Agenioideus birkmanni (Banks); and Xerochares expulsus Schulz; Cryptocheilus Panzer; and Xerochares Evans.     

A phylogenetic analysis of higher-level gall wasp relationships (Hymenoptera: Cynipidae)

We present the most comprehensive analysis of higher-level relationships in gall wasps conducted thus far. The analysis was based on detailed study of the skeletal morphology of adults, resulting in 164 phylogenetically informative characters, complemented with a few biological characters. Thirty-seven cynipid species from thirty-one genera, including four genera of the apparently monophyletic Cynipini and almost all of the genera in the other tribes, were examined. The outgroup included exemplar species from three successively more distant cynipoid families: Figitidae (the sister group of the Cynipidae), Liopteridae and Ibaliidae. There was considerable homoplasy in the data, but many groupings in the shortest tree were nonetheless well supported, as indicated by bootstrap proportions and decay indices. Partitioning of the data suggested that the high level of homoplasy is characteristic of the Cynipidae and not the result of the amount of available phylogenetically conservative characters being exhausted. The analysis supported the monophyly of the Cynipini (oak gall wasps) which, together with the Rhoditini (the rose gall wasps), Eschatocerini and Pediaspidini formed a larger monophyletic group of gall inducers restricted to woody representatives of the eudicot subclass Rosidae. The inquilines (Synergini) were indicated to be monophyletic, whereas the Aylacini, primarily herb gall inducers, appeared as a paraphyletic assemblage of basal cynipid groups. The shortest tree suggests that the Cynipidae can be divided into three major lineages: one including the inquilines, the Aylacini genera associated with Rosaceae, and Liposthenes ; one consisting entirely of Aylacini genera, among them Aulacidea , Isocolus and Neaylax ; and one comprising the woody rosid gallers (the oak and rose gall wasps and allies), the Phanacis-Timaspis complex and the Aylacini genera associated with Papaveraceae.     

Intraspecific parasitism and nest defence in the solitary pompilid wasp Anoplius viaticus (Hymenoptera: Pompilidae)

The nesting behaviour of individually marked female pompilid wasps, Anoplius viaticus , was observed at a Breckland heath site with particular emphasis on intraspecific parasitism and nest defence. Prey was stolen from conspecifics while it was being carried to the nest site, and while it was left unattended during nest construction. Females also appeared to brood parasitize each other's completed nests. Parasitism appeared to be opportunistic. Brood parasitism may be a tactic by which time-limited females can increase their fecundity. By placing prey in vegetation tufts during nest construction, females may reduce the risk of prey theft. An individual female's successive unicellular nests were clustered and therefore easier to defend, in many ways resembling a multicellular nest. Females defended their clusters vigorously, visiting them every few minutes during foraging and expelling conspecifics from the vicinity. This type of nest defence may be costly, and has rarely been observed in solitary wasps.     

Spider hawk in sand dunes: Anoplius bicinctus (Hymenoptera: Pompilidae), a parasitoid wasp of the sex-role reversed spider Allocosa brasiliensis (Araneae: Lycosidae)

In general pompilid wasps parasitize female spiders due to their larger size and/or higher fat reserves compared to males. We describe parasitoid behavior of the pompilid wasp Anoplius bicinctus on Allocosa brasiliensis and investigate the frequencies of parasitoidism according to spider developmental stage. We performed field observations and recorded frequency and duration of parasitoidism behaviors, stage and sex of the paralyzed spider (n?=?29). We found two distinct behaviors of carrying the spider (forward transport or backward transport), that were dependent of wasp size and were maintained during the transport process. Juveniles and females of A. brasiliensis showed higher frequencies of parasitoidism compared to males, although males are larger and have higher fat content, suggesting more efficient defenses against wasp attacks in this sex.     

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.     

Preliminary evolutionary relationships within the parasitoid wasp genus Cotesia (Hymenoptera: Braconidae: Microgastrinae): combined analysis of four genes

Abstract. The braconid wasp genus Cotesia Cameron (Braconidae: Microgastrinae) is one of the largest genera of parasitoid wasps, and its species are employed frequently as biological control agents against pest insects. Several Cotesia species are also used as model organisms in physiology, ecology and population genetics studies. The genus thus has considerable importance from both applied and basic science perspectives. We investigated the phylogenetic relationships of twenty‐five species of Cotesia commonly used in field and laboratory research, using the genes 16S, ND1, 28S and LW opsin and employing a range of phylogeny estimation methods including maximum parsimony, maximum likelihood, minimum evolution and Bayesian inference. Phylogenetic estimates differed little among methods, especially for the combined data analysis. The combined analysis of four genes identified four well‐supported clades within Cotesia: the melanoscela group (containing C. melanoscela, the C. flavipes species complex and probably also C. ruficrus), the kariyai group (containing C. kariyai, C. kazak, C. cyaniridis, C. flaviconchae and probably also C. anisotae and C. griffini), the rubecula group (containing C. congregata, C.electrae, C.euchaetis, C. marginiventris, C. obsuricornis and C. schizurae), and the glomerata group (consisting of C. glomerata, C. melitaearum and C. plutellae), plus a basal unresolved complex including C. hyphantriae, C. diacrisiae and C. empretiae. These groups correspond poorly with previous broad subgroups of Cotesia defined by Papp based on morphology. The current work constitutes the first real framework for comparative studies in systematics, ecology, physiology and population genetics of Cotesia. A preliminary analysis of the evolution of gregarious development from solitary is presented, in which it is apparent that solitary development is ancestral, and gregariousness has arisen several times independently within separate groups.     

Prey selection by a spider wasp,Batozonellus lacerticida (Hymenoptera: Pompilidae): Effects of seasonal variation in prey species,size and density

Prey selection by a spider wasp,Batozonellus lacerticida, was investigated at the riverside of the Toyohira River in Misumai, Sapporo, Hokkaido over 7 years from 1981 to 1987. Seventeen species of araneid spiders were found in the study area and six of them were hunted by the wasps. Most (97.3%) of the 223 prey records obtained were of three species,Araneus macacus, A. marmoreus andA. pinguis. Nesting activity of the wasps continued until mid-August in 1981, 1982 and 1986, but stopped in July in 1984, 1985 and 1987. The length of nesting period was related to whether the wasps successfully switched prey species fromA. macacus toA. marmoreus andA. pinguis andA. pinguis or not Switching seemed to occur at the phase when density ofA. macacus remained high, depending on the density of largerA. marmorus andA. pinguis. This switching had large effects on species composition and size distribution of actual prey. Chesson's index α calculated at every 10 day period revealed that the wasps preferredA. macacus to the two other species and large prey to medium and small ones. The effect of prey density on preference depended on whetherA. macacus was present or absent. In the presence of it there were only slight differences in preference among various conditions of prey density. However, whenA. macacus was absent, densities of bothA. marmoreus andA. pinguis had some complementary effects on preference. The importance of prey size selection by pompilid wasps, and implications of density effects on preference are discussed.     

中国蛛蜂科十新记录种

记述采自中国河北省小五台山、宁夏同族自治区六盘山的蛛蜂科 Pompilidae 2亚科(Pepsinae,Pompilinae)、5 属(Agenioideus,Arachnospila,Cryptocheilus,Evagetes,Priocnemis)、10中国新记录种.研究标本分别保存在河北大学和云南农业大学昆虫...     

Nesting behavior of the semi-aquatic spider wasp,Anoplius eous,which transports its prey on the surface film of water (Hymenoptera,Pompilidae)

Anoplius eous Yasumatsu (Hymenoptera, Pompilidae) exhibits some outstanding nesting behavior. Females excavate a unicellular or multicellular nest in wet ground, or dig a single-celled nest in rotten wood, or build clustered mud cells in narrow spaces between walls of such substances as wood or vinyl sheets. The latter nest-type has been unknown within the subfamily Pompilinae. Females prepare a nest-cell before hunting, and construct it further after hunting, leaving the prey near the nest (behavior-formula: IVPTIOC). They transport their prey backward on the ground, grasping it in their mandibles by any part of the legs, and forward on the surface film of water or on the ground, grasping it by the middle part of the 1st or 2nd legs. Their only prey is the adult female semi-aquatic spider,Pardosa pseudoannulata (Lycosidae). These nesting and provisioning behavior patterns are compared with those of other pompilids. The position of the present species in the behavioral evolution of the Pompilidae is suggested.     

Systematic studies on the Pompilidae occurring in Japan: Genus Platydialepis Haupt (Hymenoptera: Pompilidae: Pepsinae)

Generic characters of Platydialepis Haupt, 1941 are presented. A new species of the genus, P. amamiensis sp. nov. from the Amami Islands, the Ryukyus, Japan, is described, and another species of the genus, P. ryoheii Ishikawa, 1957, distributed in Honshu, Shikoku and Kyushu, Japan, is redescribed. Their biological notes are also given. A new combination is proposed: Platydialepis taiwaniana (Tsuneki, 1989) (=Malloscelis taiwaniana Tsuneki).     

Systematic study of the genus Cyphononyx (Hymenoptera: Pompilidae) occurring in Japan

Generic characteristics and phylogenetic relationships of the genus Cyphononyx are presented, together with a brief review of the biology of the genus. It is shown that only one species of the genus, C. fulvognathus (Rohwer), occurs in Japan. This species is distributed also in Taiwan and south China, and has hitherto been recognized under the name of C. dorsalis. In addition, two new combinations are presented: Cyphononyx hirtus (Haupt, 1935) (= Cryptochilus hirtus Haupt) and Cyphononyx plebejus (de Saussure, 1867) (= Agenia plebeja de Saussure), both of which were recorded from the Oriental Region.     

Behavior during artifically induced swarm emigration in an old world polistine wasp,Ropalidia romandi (Hymenoptera: Vespidae)

Absconding swarms of the Australian swarm-founding wasp,Ropalidia romandi, were artificially induced. Workers may initially search for new nesting sites and once a new nest site is decided, they recruit their nest mates including multiple queens there. During the swarm emigration, workers often land on prominent objects, such as tree leaves, and drag the metasoma; those that do not drag usually antennate the object surfaces. These findings suggest thatR. romandi workers use a scent trail during swarm emigration. Artificial transfer of colonies resulted in clustering of wasps from different colonies, indicating thatR. romandi may not act exclusively against non-nestmates. In the formation of tentative aggregation of wasps prior to commencing emigration, and also in recognizing a new nesting site by following swarm members that have been guided by a scent trail, a visual cue (aerial swarm formed around an aggregation of wasps) seems to play an important role.     

Functional analysis of the myrmecophilous relationships between ants (Hymenoptera: Formicidae) and lycaenids (Lepidoptera: Lycaenidae)

Summary Mature larvae of the myrmecophilous lycaenid Polyommatus coridon produce an average of 30.9 droplets of a honeydew-like secretion per hour. They occur in population densities of about 20/m². The volume of secretion over the whole larval period is estimated to be 22–44 l with an energy content of 55–110 J. Thus, P. coridon larvae produce carbohydrate secretions with an energy equivalent of 1.1–2.2 kJ/m². Using data from the literature on ant metabolism, it is shown that these carbohydrate secretions may contribute significantly to the nutrition of attending ants. The myrmecophilous relationship between the larvae of P. coridon and ants should therefore be regarded as a mutualistic symbiosis.     

Proteomic analysis of the venom of the social wasp Apoica pallens (Hymenoptera: Vespidae)

Wasps are a diverse group of insects that possess a sting apparatus associated with a venom gland, which is used for predation and colony defense. The biochemistry of Hymenoptera venom has been evaluated in relation to allergy and immunology, and proteomics has been shown to be a powerful tool for the identification of compounds with pharmacological potential. Data on wasps venom the of genus Apoica are scarce, so the objective of the present work was to identify the venom proteins of the eusocial wasp Apoica pallens, as a first step towards further investigation of applied uses of the venom and its protein constituents. The venom proteins were separated by two-dimensional gel electrophoresis, followed by MALDI-TOF/TOF mass spectrometry. A total of 259 spots were detected, with molecular weights from 4.9 to 141 kDa. Thirty of these proteins were identified and classified into eight functional categories: allergen, enzyme, metabolism, structural, environmental response, proteoglycan, active in DNA and RNA, and unknown function. Due to the few available proteomic data for wasp venom, many proteins could not be identified, which makes studies with proteomic analysis of Hymenoptera venom even more important.