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Capturing phylogenetic signal from a massive radiation can be daunting. The superfamily Chalcidoidea is an excellent example of a hyperdiverse group that has remained recalcitrant to phylogenetic resolution. Chalcidoidea are mostly parasitoid wasps that until now included 27 families, 87 subfamilies and as many as 500,000 estimated species. We comb...
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... In order to reach their hosts and permit greater control over egg placement, several parasitoid wasps are able actively to bend and rotate their terebra in any direction relative to their body axis [41][42][43][44], despite the lack of intrinsic terebral musculature. Such terebra movements have also been reported in chalcidoid wasps of the family of Pteromalidae [40,[45][46][47], a polyphyletic group sensu lato [6,10,48,49] (over 3500 species described [8]). However, little is known about the actuation of the various ovipositor movements, with the mechanisms involved in terebra steering (i.e. ...
... The structure of the terebra of Chalcidoidea, featuring a longitudinally split 2nd valvula with overlapping, asymmetric halves, is strongly consistent in structure within families and basically similar across families (with the exception of the primitive Mymaridae [18,39], which recently have been identified as a sister group to all remaining Chalcidoidea [10,49]), but is unique among other superfamilies of parasitoid Hymenoptera [39]. The similar structure of the terebra of chalcidoid taxa might indicate similar underlying working mechanisms, since form and function are strongly connected [113,114]. ...
... [127]) and presumably have led to rapid adaptive radiations in Chalcidoidea [10] (for speciation process in L. distinguendus see [128][129][130]). Thus, the ability actively to steer the terebra potentially has been a central factor in the evolution of the parasitoid life history strategy and the diversification of chalcidoid wasps, resulting in the evolutionary success of this group with its tremendous extant species richness [10,49]. The Chalcidoidea are the most diverse group of parasitoid hymenopterans, with estimations of more than 500,000 chalcidoid species, the vast majority of them being parasitoids, out of a total of 680,000 parasitoid hymenopteran species [6,7]. ...
Various chalcidoid wasps can actively steer their terebra (= ovipositor shaft) in diverse directions, despite the lack of terebral intrinsic musculature. To investigate the mechanisms of these bending and rotational movements, we combined microscopical and microtomographical techniques, together with videography, to analyse the musculoskeletal ovipositor system of the ectoparasitoid pteromalid wasp Lariophagus distinguendus (Förster, 1841) and the employment of its terebra during oviposition. The ovipositor consists of three pairs of valvulae, two pairs of valvifers and the female T9 (9th abdominal tergum). The paired 1st and the 2nd valvulae are interlocked via the olistheter system, which allows the three parts to slide longitudinally relative to each other, and form the terebra. The various ovipositor movements are actuated by a set of nine paired muscles, three of which (i.e. 1st valvifer-genital membrane muscle, ventral 2nd valvifer-venom gland reservoir muscle, T9-genital membrane muscle) are described here for the first time in chalcidoids. The anterior and posterior 2nd valvifer-2nd valvula muscles are adapted in function. (1) In the active probing position, they enable the wasps to pull the base of each of the longitudinally split and asymmetrically overlapping halves of the 2nd valvula that are fused at the apex dorsally, thus enabling lateral bending of the terebra. Concurrently, the 1st valvulae can be pro- and retracted regardless of this bending. (2) These muscles can also rotate the 2nd valvula and therefore the whole terebra at the basal articulation, allowing bending in various directions. The position of the terebra is anchored at the puncture site in hard substrates (in which drilling is extremely energy- and time-consuming). A freely steerable terebra increases the chance of contacting a potential host within a concealed cavity. The evolution of the ability actively to steer the terebra can be considered a key innovation that has putatively contributed to the acquisition of new hosts to a parasitoid’s host range. Such shifts in host exploitation, each followed by rapid radiations, have probably aided the evolutionary success of Chalcidoidea (with more than 500,000 species estimated).
... In previous classification, Walkerella microcarpae and Micranisa ralianga belonged to Otitesellinae and Apocrypta bakeri, Philotrypesis tridentata, Philotrypesis pilosa, and Philotrypesis sp. to Sycoryctinae. Based our results, Otitesellinae were recovered as nested within Sycoryctinae, which made Sycoryctinae paraphyletic, a result consistent with Zhao et al. [40] and Cruaud et al. [78]. The clade comprising Otitesellinae and Sycoryctinae appeared sister to other Pteromalinae included in our analysis, a result consistent with Rasplus et al. [23]. ...
... In addition, Sycoryctinae were not recovered as monophyletic, a result consistent with Munro et al. [18]. However, according to the classification system recently proposed by Burks et al. [68] and based on the analyses by Cruaud et al. [78], our results also corroborated that members of the previous Otitesellinae and Sycoryctinae belong in fact to Pteromalinae and represent at most a tribe of this subfamily. All other pteromalid subfamilies were recovered as monophyletic. ...
... All other pteromalid subfamilies were recovered as monophyletic. Therefore, our study supports the classification proposed by Burks et al. [68] based on nuclear Ultra-Conserved Elements and exons [78]. Our result shows the power of mitogenomes to reconstruct family-level phylogenies in Chalcidoidea, only for three of the eight pteromalid subfamilies and 11 of the 500+ recognized genera. ...
The mitochondrial genomes of Muscidifurax similadanacus, M. sinesensilla, Nasonia vitripennis, and Pachycrepoideus vindemmiae were sequenced to better understand the structural evolution of Pteromalidae mitogenomes. These newly sequenced mitogenomes all contained 37 genes. Nucleotide composition was AT-biased and the majority of the protein-coding genes exhibited a negative AT skew. All 13 protein-coding genes (PCGs) initiated with the standard start codon of ATN, excepted for nad1 of N. vitripennis, which started with TTG, and terminated with a typical stop codon TAA/TAG or an incomplete stop codon T. All transfer RNA (tRNA) genes were predicted to fold into the typical clover-leaf secondary structures, except for trnS1, which lacks the DHU arm in all species. In P. vindemmiae, trnR and trnQ lack the DHU arm and TΨC arm, respectively. Although most genes evolved under a strong purifying selection, the Ka/Ks value of the atp8 gene of P. vindemmiae was greater than 1, indicating putative positive selection. A novel transposition of trnR in P. vindemmiae was revealed, which was the first of this kind to be reported in Pteromalidae. Two kinds of datasets (PCG12 and AA) and two inference methods (maximum likelihood and Bayesian inference) were used to reconstruct a phylogenetic hypothesis for the newly sequenced mitogenomes of Pteromalidae and those deposited in GenBank. The topologies obtained recovered the monophyly of the three subfamilies included. Pachyneurinae and Pteromalinae were recovered as sister families, and both appeared sister to Sycophaginae. The pairwise breakpoint distances of mitogenome rearrangements were estimated to infer phylogeny among pteromalid species. The topology obtained was not totally congruent with those reconstructed using the ML and BI methods.
... According to the modern molecular and intricate combined analyses of Munro et al. [31] and Cruad et al. [32] (and references therein), the evolutionary history of Encyrtidae began over 100 million years ago during the Cretaceous, when Chalcidoidea underwent ...
... According to the modern molecular and intricate combined analyses of Munro et al. [31] and Cruad et al. [32] (and references therein), the evolutionary history of Encyrtidae began over 100 million years ago during the Cretaceous, when Chalcidoidea underwent a rapid radiation. Along with several other families of "soft bodied" chalcidoids of the "Tiny Wasp clade", Encyrtidae diverged soon after. ...
... Along with several other families of "soft bodied" chalcidoids of the "Tiny Wasp clade", Encyrtidae diverged soon after. The first lineages to diverge (Mymaridae, Baeomorphidae (Rotoitidae), and "Tiny Wasp clade") were likely first oophagous and later associated mostly with hemipteran hosts [32]. ...
Balticalcarus archibaldi Simutnik, gen. et sp. n., is described and illustrated based on a female specimen from late Eocene Baltic amber. The new genus is characterized by the absence of a filum spinosum, a “boat”-shaped hypopygium enclosing the ovipositor, reaching far past the apex of the syntergum, the presence of a line of long setae along the entire costal cell of the hind wing, and a transverse line of thickened setae alongside the hyaline spur vein. Moreover, like most previously described Eocene Encyrtidae, the new taxon differs from the majority of the extant ones by a number of morphological features. The new fossil differs from most extant and all known fossil Encyrtidae by its unusually small, thin, smooth (without microsetae) mesotibial spur.
In this study, we explore the biological resources of Eunotidae, Herbertiidae, Pteromalidae
and Eulophidae in the Altun Mountain Nature Reserve, Xinjiang, China. Sixty-one species
are listed and we described ten new species, including Eunotus caeruleus Kang & Hu,
sp. nov. and Eunotus argenteus Kang & Hu, sp. nov. of Eunotidae, Herbertia altunensis
Kang & Hu, sp. nov. of Herbertiidae, Thinodytes splendens Kang & Hu, sp. nov., Erdoesina
maculata Kang & Hu, sp. nov., Homoporus flavus Kang & Hu, sp. nov. and Stenomalina
viridis Kang & Hu, sp. nov. of Pteromalidae and two new species Diaulinopsis altunensis
Kang & Hu, sp. nov. and Hyssopus altunensis Kang & Hu, sp. nov. of Eulophidae. Detailed
illustrations of all new species are included to support identification and further study.
The earliest representatives of Chalcidoidea are described from Barremian age Early Cretaceous Lebanese amber and classified in Protoitidae Ulmer & Krogmann, fam. nov. (Hymenoptera: Chalcidoidea). Protoitidae exhibits a high morphological diversity of the terminal metasomal tergum which may indicate a broad spectrum of oviposition capabilities and the ability to occupy a diverse range of ecological niches. Protoitidae comprises two genera, Protoita Ulmer & Krogmann, gen. nov. , and Cretaxenomerus Nel & Azar, 2005 based on C. jankotejai Nel & Azar, 2005, which is transferred from Scelionidae (Hymenoptera: Platygastroidea) to Protoitidae. Together, 10 new species, all by Ulmer and Krogmann, are described in the two included genera– Protoita bidentata , P. istvani , P. noyesi , P. petersi , Cretaxenomerus brevis , C. curvus , C. deangelis , C. mirari , C. tenuipenna , and C. vitreus . Keys to the genera and species of Protoitidae are provided. In addition, we examine the postulated plesiomorphies and apomorphies within Chalcidoidea with respect to the fossil record, and provide additional hypotheses on their biogeographic origins.
Leptoomidae Gibson fam. nov. (Chalcidoidea) is described for the Eocene Baltic amber fossil genera Leptoomus Gibson,
type genus, reassigned from Tanaostigmatidae, and Neanaperiallus Gibson, reassigned from Neanastatinae (Chalcidoidea: Eupelmidae) sensu Gibson (2009). One new species of Neanaperiallus, N. defunctus Fusu sp. nov., is described. The new family is differentiated from other families of Chalcidoidea that are partly characterized by a greatly enlarged acropleuron. In species of Leptoomidae the prepectus is anteriorly rounded to angulate and extends to or slightly over the posterolateral margin of the pronotum, with the dorsal prepectal margin intersecting the base of the tegula distinctly anterior to and forming an almost right-angle with the posterior margin of prepectus, and the posterior margin truncate along the anterior margin of the acropleuron. This prepectal structure is similar to that in Tanaostigmatidae and Cynipencyrtidae, except the prepectus is elongated anteriorly exterior to the pronotum in Tanaostigmatidae and interior to the lateral surface of the pronotum in Cynipencyrtidae. A difference in prepectal structure also indicates that an anteriorly elongated mesoscutal process internal to the pronotum in Encyrtidae is convergent to that of Cynipencyrtidae, and similarity in shape of the prepectus among Encyrtidae, Eopelma Gibson and Neanastatus Girault might be functionally correlated with an anterior elongation of the mesoscutal process. New or corrected morphological data are provided for the two included genera. Of other Eocene fossil genera originally classified in Neanastatinae, Brevivulva Gibson and Propelma Trjapitzin, are
assigned to Neanastatidae sensu Burks et al. (2022) based on similar mesoscutellar structures. Possible relationships of Aspidopleura Gibson, a taxon with a puzzling combination of features, are discussed. Because Aspidopleura cannot be placed with confidence in any extinct or extant higher taxon it is treated as incertae sedis at family level within Chalcidoidea.
Parasitoid wasps are the most successful group of insect parasitoids, comprising more than half the known diversity of Hymenoptera and probably most of the unknown diversity. This life-style has enabled them to be used as pest control agents conferring substantial economic benefits to global agriculture. Major lineages of parasitoid wasps include Ichneumonoidea, Ceraphronoidea, Proctotrupomorpha and a number of aculeate families. The parasitoid life-style arose only once among basal Hymenoptera, in the common ancestor of the Orussidae and Apocrita some 200+ Ma ago. The ancestral parasitoid wasp was probably an idiobiont on wood-living beetle larvae. From this comparatively simple biology, Hymenoptera radiated into an incredible diversity of hosts and parasitoid lifestyles, including hyperparasitoidism, kleptoparasitoidism, egg-parasitoidism and polyembryony, in several instances co-opting viruses to subdue their hosts. Many lineages evolved beyond the parasitoid niche, becoming secondarily herbivorous or predatory nest provisioners and eventually giving rise to most instances of insect societies.
Anucleate animal cells are a peculiar evolutionary phenomenon and a useful model for studying cellular mechanisms. Anucleate neurons were recently found in one genus of miniature parasitic wasps of the family Trichogrammatidae, but it remained unclear how widespread this phenomenon is among other insects or even among different tissues of the same insect species. We studied the anatomy of miniature representatives of another parasitic wasp family (Hymenoptera: Mymaridae) using array tomography and found two more species with nearly anucleate brains at the adult stage. Thus, the lysis of the cell bodies and nuclei of neurons appears to be a more widespread means of saving space during extreme miniaturization, which independently evolved at least twice during miniaturization in different groups of insects. These results are important for understanding the evolution of the brain during miniaturization and open new areas of studying the functioning of anucleate neurons.
The family Pteromalidae (Hymenoptera: Chalcidoidea) is reviewed with the goal of providing nomenclatural changes and morphological diagnoses in preparation for a new molecular phylogeny and a book on world fauna that will contain keys to identification. Most subfamilies and some tribes of Pteromalidae are elevated to family level or transferred elsewhere in the superfamily. The resulting classification is a compromise, with the aim of preserving the validity and diagnosability of other, well-established families of Chalcidoidea. The following former subfamilies and tribes of Pteromalidae are elevated to family rank: Boucekiidae, Ceidae, Cerocephalidae, Chalcedectidae, Cleonymidae, Coelocybidae, Diparidae, Epichrysomallidae, Eunotidae, Herbertiidae, Hetreulophidae, Heydeniidae, Idioporidae, Lyciscidae, Macromesidae, Melanosomellidae, Moranilidae, Neodiparidae, Ooderidae, Pelecinellidae (senior synonym of Leptofoeninae), Pirenidae, Spalangiidae, and Systasidae. The following subfamilies are transferred from Pteromalidae: Chromeurytominae and Keiraninae to Megastigmidae, Elatoidinae to Neodiparidae, Nefoeninae to Pelecinellidae, and Erotolepsiinae to Spalangiidae. The subfamily Sycophaginae is transferred to Pteromalidae. The formerly incertae sedis tribe Lieparini is abolished and its single genus Liepara is transferred to Coelocybidae. The former tribe Tomocerodini is transferred to Moranilidae and elevated to subfamily status. The former synonym Tridyminae (Pirenidae) is treated as valid. The following former Pteromalidae are removed from the family and, due to phylogenetic uncertainty, placed as incertae sedis subfamilies or genera within Chalcidoidea: Austrosystasinae, Ditropinotellinae, Keryinae, Louriciinae, Micradelinae, Parasaphodinae, Rivasia , and Storeyinae. Within the remaining Pteromalidae, Miscogastrinae and Ormocerinae are confirmed as separate from Pteromalinae, the former tribe Trigonoderini is elevated to subfamily status, the former synonym Pachyneurinae is recognized as a distinct subfamily, and as the senior synonym of Austroterobiinae. The tribe Termolampini is synonymized under Pteromalini, and the tribe Uzkini is synonymized under Colotrechnini. Most former Otitesellinae, Sycoecinae, and Sycoryctinae are retained in the tribe Otitesellini, which is transferred to Pteromalinae, and all other genera of Pteromalinae are treated as Pteromalini. Eriaporidae is synonymized with Pirenidae, with Eriaporinae and Euryischiinae retained as subfamilies. Other nomenclatural acts performed here outside of Pteromalidae are as follows: Calesidae: elevation to family rank. Eulophidae: transfer of Boucekelimini and Platytetracampini to Opheliminae, and abolishment of the tribes Elasmini and Gyrolasomyiini. Baeomorphidae is recognized as the senior synonym of Rotoitidae. Khutelchalcididae is formally excluded from Chalcidoidea and placed as incertae sedis within Apocrita. Metapelmatidae and Neanastatidae are removed from Eupelmidae and treated as distinct families. Eopelma is removed from Eupelmidae and treated as an incertae sedis genus in Chalcidoidea. The following subfamilies and tribes are described as new: Cecidellinae (in Pirenidae), Enoggerinae ( incertae sedis in Chalcidoidea), Erixestinae (in Pteromalidae), Eusandalinae (in Eupelmidae), Neapterolelapinae ( incertae sedis in Chalcidoidea), Solenurinae (in Lyciscidae), Trisecodinae (in Systasidae), Diconocarini (in Pteromalidae: Miscogastrinae), and Trigonoderopsini (in Pteromalidae: Colotrechninae). A complete generic classification for discussed taxa is provided.
