The phylogeny and evolution of Figitidae (Hymenoptera: Cynipoidea)

A phylogeny of the Figitidae (Hymenoptera: Cynipoidea) is presented based on combined analysis of molecular (28S‐D2 and D3, COI and 18S‐E17‐35), morphological and life‐history data. Data are analyzed by parsimony and Bayesian inference methods. Taxon sampling was held at a premium, and the resulting matrix contained 168 terminal taxa representing eight of nine subfamilies (Pycnostigminae not included) and all major subgroups of each subfamily. Alignment of the 28S D2 + D3 gene fragment based on a structural model resulted in the most defendable and least conflicting alignment tested. Melanips, previously classified in Figitinae, was consistently found to be the sister group of the Aspicerinae; Euceroptres, historically classified in Thrasorinae, frequently rendered that subfamily paraphyletic in these analyses. The general evolutionary trend is for early figitids to be parasitoids of gall inducing insects, with later host shifts occurring to exposed hosts associated with aphids. © The Willi Hennig Society 2007.     

Phylogeny of the Eucoilinae (Hymenoptera: Cynipoidea: Figitidae)

The Eucoilinae are a diverse and important group of parasitoids of Diptera, particularly in the tropics, but they are poorly known systematically and their generic classification is partly chaotic. Here, we present the first comprehensive cladistic analysis of higher eucoiline relationships. The analysis is based on 148 skeletal characters of adults documented in more than 1100 digital images available in an Internet-accessible database. The characters were coded for 45 taxa representing 35 eucoiline genera, spanning the entire diversity of the group, and 7 outgroup genera. Relationships were partly difficult to resolve and parsimony analysis under implied weights performed considerably better than analysis under uniform weights. The results support the monophyly of the Eucoilinae and show that eucoilines are most closely related to the figitid subfamilies Emargininae and Pycnostigminae, but are ambiguous concerning the exact relationships among these three lineages. Of the 6 eucoiline genus groups recognized by Nordlander in 1982 ( Entomol. Scand. 13, 269–292), only 2 are supported as monophyletic: the Trybliographa and Kleidotoma groups. The Gronotoma group is a paraphyletic assemblage of two different basal clades of eucoilines. The Rhoptromeris group is unnatural and only the 2 core genera, Rhoptromeris and Trichoplasta , form a monophyletic lineage. The data are ambiguous concerning the Ganaspis group, which appears to be paraphyletic, and the Chrestosema group, which may be a good clade. Based on the results we propose a modified system of informal genus groups in the Eucoilinae and discuss putative synapomorphies supporting each genus group. The proposed relationships imply that the first eucoilines were parasitoids of leaf-mining agromyzids. The earliest split in the group was apparently between an Afrotropical and a Neotropical lineage, and much of the early radiation of the group occurred in these regions, particularly in the Neotropics.     

A new subfamily of Figitidae (Hymenoptera, Cynipoidea)

Larvae of the parasitic wasp family Figitidae develop as internal parasitoids of other endopterygote insect larvae. The hosts are typically dipteran larvae living in other microhabitats but the earliest figitids probably attacked gall-inhabiting hymenopteran larvae. Here, we formally describe a new genus (Parnips) and subfamily (Parnipinae) for a species that is likely to be a surviving representative of these early gall-associated figitids. The species, P. nigripes , has been reared repeatedly from galls inside the seed capsules of annual poppies (Papaver dubium and P. rhoeas) in the Mediterranean region together with the gall inducer Barbotinia oraniensis belonging to the Cynipidae, the sister group of Figitidae. Parnips nigripes is strikingly cynipid-like and was first assumed to be a cynipid gall inducer of the genus Aulacidea. Phylogenetic analyses have since indicated that the similarity with the Cynipidae is symplesiomorphic and that P. nigripes belongs to the Figitidae, where it forms the sister group of all other extant figitids. Recently, it has also been shown that P. nigripes is a parasitoid of the gall-inducing Barbotinia oraniensis , consistent with its proposed phylogenetic position. Parnips nigripes shares several unusual morphological traits with its host. We speculate that many of these similarities are homologous even though the lineages separated at least 83 million years ago.     

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 and early evolution of the Cynipoidea (Hymenoptera)

Based on several structural and biological characteristics, the Cynipoidea can be divided into two groups, 'macrocynipoids' and 'microcynipoids'. The macrocynipoids (i.e. the family Liopteridae and the genera Austrocynips, Eileenella, Heteribalia and Ibalia ) are generally large insects that parasitize wood- or cone-boring insect larvae. The microcynipoids are smaller insects that are either phytophagous gall inducers and inquilines (Cynipidae) or parasitoids of larvae of Hymenoptera, Neuroptera or Diptera (Figitidae sensu lato , including the families Eucoilidae, Charipidae and Anacharitidae). The phylogenetic relationships among genera of macrocynipoids and between these and a sample of four genera representing the Figitidae and Cynipidae were examined by parsimony analysis of 110 external morphological characters of adults. Within the macrocynipoids, three monophyletic lineages emerged, classified here as different families: the Austrocynipidae, with a single species, Austrocynips mirabilis , the only cynipoid with a true pterostigma; the Ibaliidae, including the genera Eileenella, Ibalia and Heteribalia ; and the Liopteridae, comprising the remaining genera of macrocynipoids. The analysis further supported the monophyly of the microcynipoids and indicated that the macrocynipoids form a paraphyletic group relative to the microcynipoids, with the shortest tree suggesting the relationship (Austrocynipidae, (Ibaliidae, (Liopteridae, microcynipoids))). These results imply that cynipoids were originally parasitoids of wood-boring insect larvae and that the other modes of life evolved secondarily within the group.     

Afrotropical Cynipoidea (Hymenoptera)

The Afrotropical Cynipoidea are represented by 306 described species and 54 genera in four families: Cynipidae, Figitidae, Liopteridae and Ibaliidae, the latter represented by a single introduced species. Seven of these genera are only represented by undescribed species in the region. Seven new genus-level synonymies, one genus resurrected from synonymy, 54 new combinations, one combination reinstated, and one new replacement name are presented. We provide identification keys to the families, subfamilies and genera of cynipoid wasps occurring in the Afrotropical region (Africa south of the Sahara, including Madagascar and southern Arabian Peninsula). Online interactive Lucid Phoenix and Lucid matrix keys are available at: http://www.waspweb.org/Cynipoidea/Keys/index.htm. An overview of the biology and checklists of species for each genus are provided. This paper constitutes the first contributory chapter to the book on Afrotropical Hymenoptera.     

Description of a New Genus and Three New Species of Figitinae (Hymenoptera: Cynipoidea: Figitidae) from Colombia

A new genus and three new species of Figitinae (Hymenoptera: Figitidae) are described from Colombia: Ferpereira Pujade-Villar n. gen., Ferpereira fiorellae Pujade-Villar n. sp., Neralsia levis Pujade-Villar & Petersen-Silva n. sp., and Figites colombiensis Pujade-Villar & Paretas-Martínez n. sp., which is the first species of Figites Latreille from Colombia and the second from South America. The diagnostic characters of these new taxa are illustrated.     

Redescription of Acanthaegilips Ashmead, 1897, with characterization of the Anacharitinae and Aspiceratinae (Hymenoptera: Cynipoidea: Figitidae)

We redescribe the South American genus Acanthaegilips Ashmead, 1897 and discuss its phylogenetic position within the Figitidae (sensu lata). The genus was originally placed in the Anacharitinae but shows affinities with both the Anacharitinae and the Aspiceratinae in the characters used currendy to separate figitid subfamilies. In a recent revision of the higher‐level classification of cynipoids, Acanthaegilips was separated from the remainder of the Anacharitinae and placed in a monotypic higher‐level taxon. We analyse the morphological differences between die Anacharitinae and Aspiceratinae and their bearing on the monophyly of the two subfamilies and the placement of Acanthaegilips. We conclude that, after removal of Seitneria and Paraegilips from the Anacharitinae, both subfamilies are well defined monophyletic groups and that Acanthaegilips belongs to the Anacharitinae, within which it forms a monophyletic group together with the South American genera Calofigites Kieffer, 1909 and Solenofigites Diaz, 1979.     

Description of nine South American species of the genus Neralsia cameron, with low scutellar carina (Hymenoptera: Cynipoidea: Figitidae)

In this study a total of nine new species of the genus Neralsia are described for South America: N. desantisi Jiménez & Pujade-Villar n. sp., N. equilatera Jiménez & Pujade-Villar n. sp., N. hermafrodita Jiménez & Pujade-Villar n. sp., N. marioi Jiménez & Pujade-Villar n. sp., N. moisesi Jiménez & Pujade-Villar n. sp., N. obelix Jiménez & Pujade-Villar n. sp., N. parafossulata Jiménez & Pujade-Villar n. sp., N. pseudoneralsia Jiménez & Pujade-Villar n. sp. and N. rauli Jiménez & Pujade-Villar n. sp., all of them have in common the low carina separating the scutellar foveae. Other six known Neralsia species, N. albipennis (Kieffer), N. bogotensis (Kieffer), N. flavidipennis (Kieffer), N. fossulata (Kieffer), N. pilosa (Borgmeier) and N. striaticeps (Kieffer) have the same scutellar character. The morphological characters to differentiate all mentioned species are presented.     

Two new genera of Eucoilidae (Hymenoptera: Cynipoidea)

Psichacra marshalli Cameron is redescribed and assigned to the new genus Pseudopsichacra. ‘New Genus B’ of Weld is redescribed as Bewelda gen.n. and striata sp.n. is designated as the type–species. An emendation of Weld's key is included to show the positions of both genera.     

Review of Afrotropical Figitinae (Figitidae,Cynipoidea, Hymenoptera) with the first records of Neralsia and Lonchidia for the region

The cynipoid subfamily Figitinae is poorly represented in the Afrotropical region with two genera (Figites Latreille and Xyalophora Kieffer) and six species currently known. Here we record an additional two genera (Neralsia Cameron and Lonchidia Thomson) for the region and describe three new species: Neralsia haddocki sp. n.; Xyalophora tedjoansi sp. n.; Xyalophora tintini sp. n. Benoit’s species described in 1956 are synonymized under Figites aciculatus (Benoit, 1956): Figites effossus syn. n.; Figites favonius syn. n.; Figites furvus syn. n.; Figites fraudator syn. n. Identification keys to the figitine genera and species occurring in the Afrotropical region are provided. Online interactive Lucid Phoenix and Lucid matrix keys are available at: http://www.waspweb.org/Cynipoidea/Keys/index.htm     

Taxonomy and phylogeny of Leptopilina species (Hymenoptera: Cynipoidea: Figitidae) attacking frugivorous drosophilid flies in Japan,with description of three new species

Despite the intensive use of the Leptopilina genus and its drosophilid hosts as model systems in the study of host–parasitoid interactions, the diversity and distribution of the species occurring in the Asian region remain elusive. Here we report the phylogeny of Japanese Leptopilina species attacking frugivorous drosophilid flies, based on COI, ITS1 and ITS2 sequences. Consistent with molecular data, hybridization experiments and morphological examination, five species were recorded in Japan: Leptopilina heterotoma, L. victoriae and three new species, two occurring in the Ryukyu archipelago, L. ryukyuensis and L. pacifica, and another species, L. japonica, distributed in Honshu and Hokkaido. Leptopilina japonica is further divided into two subspecies, L. j. japonica occurring in Japan, and L. j. formosana occurring in Taiwan. According to these results, we discuss the evolution, speciation and colonization history of Japanese Leptopilina species.     

Systematics of Australian Thrasorinae (Hymenoptera, Cynipoidea, Figitidae) with descriptions of Mikeiinae, new subfamily, two new genera, and three new species

The Australian Thrasorinae are revised and Mikeius is transferred to Mikeiinae Paretas-Martínez & Pujade-Villar, subfam. n., and Mikeius clavatus Pujade-Villar & Restrepo-Ortiz, sp. n., is described. Two new genera of Thrasorinae are erected: Cicatrix Paretas-Martínez, gen. n., including Cicatrix pilosiscutum(Girault), comb. n. from Amblynotus, Cicatrix schauffi (Buffington), comb. n. from Mikeius, and Cicatrix neumannoides Paretas-Martínez & Restrepo-Ortiz, sp. n.; and Palmiriella Pujade-Villar & Paretas-Martínez, gen. n., including Palmiriella neumanni (Buffington), comb. n. from Mikeius, Thrasorus rieki Paretas-Martínez & Pujade-Villar, sp. n., is also described. A phylogenetic analysis of 176 morphological and biological characters, including all these new taxa and all genera previously included in Thrasorinae, was conducted. All subfamilies were recovered as monophyletic, with the following relationships: Parnipinae (Euceroptrinae (Mikeiinae (Plectocynipinae (Thrasorinae)))). A worldwide key to the subfamilies of Figitidae is provided that includes the new subfamily, as well as a key to genera Thrasorinae.     

Quasimodoana, a new Holarctic genus of eucoiline wasps (Hymenoptera, Cynipoidea, Figitidae), with a phylogenetic analysis of related genera

Abstract.  A new Holarctic genus, Quasimodoana gen.n. is described for the Palearctic eucoiline wasp Eucoila decipiens Förster, 1869, as Quasimodoana decipiens comb.n. A lectotype is designated for this species. A new North American species is described here as Quasimodoana gibba sp.n. As the new genus has similarities with several eucoiline genera, belonging to two related but distinct lineages, a phylogenetic analysis was carried out, based on 83 morphological characters. The two lineages included in the analysis are referred to here under the tribal names of Eucoilini and Trichoplastini stat.nov. We discuss reasons for adopting available tribal names (rather than 'informal genus groups'), and Trichoplastini is removed from synonymy with Eucoilini. The phylogenetic analysis places Quasimodoana unambiguously within Eucoilini, as a sistergroup to the Trybliographa complex. However, there is only weak support for the monophyly of Eucoilini and for the hypothesized sistergroup relationship between Eucoilini and Trichoplastini.     

Diversity and function of male antennal glands in Cynipoidea (Hymenoptera)

The functional anatomy of antennal glands located either on the 3rd or on the 3rd and 4th antennomeres in males of several species of cynipoids was investigated. SEM observations revealed variously modified antennomeres with elevated plates, tyloids and excavated areas. In all the cases, the antennomeres are equipped with cuticular pores, corresponding internally to cuticular ducts. TEM studies showed the presence of type III integumentary glands, as classified by Noirot & Quennedey. Each glandular unit is made up of an innermost secretory cell, producing the secretion, and an outermost canal cell, producing the evacuating duct. The secretion passes through the duct and reaches the cuticular pores, concentrated in a ventro-lateral portion of the antennomere called the 'release and spread structure'. Both in Cynipidae and in Eucoilinae (Figitidae), the courtship behaviour includes a pre-copulatory phase characterized by intense antennal stroking. Bioassays in the eucoilins Leptopilina boulardi and L. heterotoma showed that these glands are the production site of a contact sex recognition pheromone, necessary for the female to accept the male.     

Morphological and molecular phylogenetics in the genus Leptopilina (Hymenoptera: Cynipoidea: Eucoilidae)

We have conducted cladistic analyses of the genus Leptopilina , a group of Drosophila parasitoids studied intensively by (behavioural) ecologists. Twenty-three morphological characters were scored in ten Leptopilina and two outgroup species. At the same time, DNA sequences for the second ribosomal internal transcribed spacer (ITS2) were gathered for eight Leptopilina and one outgroup species. Both data sets yielded phylogenetic trees which were largely compatible. A 'total evidence' analysis resulted in a single tree that provides a relatively robust phylogenetic reconstruction of the group, which may serve as a basis for historically interpreting the distribution of ecological and behavioural traits.