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The trictenotomid genus Autocrates Thomson, 1860 is remarkable for its large and robust adults, but its larval morphology and bionomics have been unknown over the last 160 years. Here, we describe and illustrate in detail the eggs, and also the first and last instar larva and the pupa of Autocrates maqueti Drumont, 2006, based on specimens reared from identified adults collected in South Korea. The first instar larva is very similar to the known trictenotomid larvae of the genus Trictenotoma Gray, 1832, sharing the following features: distinctly flattened and parallel-sided, well-sclerotized head and largely unpigmented postcephalic body, lyre-shaped frontal arms, very short coronal suture, five pairs of stemmata, asymmetrical and tridentate mandibles with mola, simple and apically upturned urogomphi, and absence of longitudinal ridges on the thorax and abdomen. Differences occur in the primary chaetotaxy. The last instar larva is almost identical with Trictenotoma in general appearance, but the longitudinal tergal ridges of Autocrates are much denser and more widely distributed than those of Trictenotoma. Our preliminary assessment of features of immature stages confirms a close relationship of Trictenotomidae with the “salpingid group”, i.e. Salpingidae, Boridae, Pyrochroidae and Pythidae. A sister group relationship with Pythidae is likely. The putative synapomorphy is the subdivision of the ventral element of tergites IX, even though this condition is not visible in the first instars of Autocrates, and quite indistinct (but recognizable) in the last larval stage. The complete mitochondrial genome of A. maqueti is provided. A preliminary phylogenetic analysis of trictenotomid species is presented using two mitochondrial genes (16S and COI). The morphology of eggs and biological information on feeding and oviposition behaviors are also provided with photographs of adults and eggs.

The Umenocoleidae is among the most perplexing fossil insect taxa, widespread at lower and middle latitudes during the Early Cretaceous and probably becoming extinct after the mid-Cretaceous. This problematic taxon was first described from the Lower Cretaceous of China and has been variously assigned to stem-group Coleoptera, Protelytroptera, Blattaria, stem-group Dictyoptera, or Mantodea. The systematic position of the enigmatic family — especially its type species Umenocoleus sinuatus Chen et T’an, 1973 — has remained highly controversial owing to the lack of preserved phylogenetically informative or diagnostic characters. Here, we describe a new specimen of the type species from the type horizon of the type locality (Lower Cretaceous Zhonggou Formation at Jiuquan, Yumen City, China), and additionally two forewings of Ponopterix. We found that the Umenocoleidae is similar to stem-group Coleoptera in its forewing venation but differs in its multi-segmented antennae, orthognathous orientation of the head, subdivision of the pronotum by a transverse furrow, ultrastructure of the forewing punctures, and unfolded hindwings. Our phylogenetic analysis, using an extended matrix of 72 characters and 37 taxa, indicates that the Umenocoleidae is likely a specialized taxon of Dictyoptera, sister to Alienoptera. The beetle-like appearance is interpreted to be a result of convergent evolution.

Conversion of forewings into hardened covers, elytra, was a ground-breaking morphological adaptation that has contributed to the extraordinary evolutionary success of beetles. Nevertheless, the knowledge of the functional aspects of these structures is still fragmentary and scattered across a large number of studies. Here, we have synthesized the presently available information on the evolution, development, modifications and biological functions of this crucial evolutionary novelty. The formation of elytra took place in the earliest evolution of Coleoptera, very likely already in the Carboniferous, and was achieved through the gradual process of progressive forewing sclerotization and the formation of inward directed epipleura and a secluded sub-elytral space. In many lineages of modern beetles, the elytra have been distinctly modified. This includes multiple surface modifications, a rigid connection or fusion of the elytra, or partial or complete reduction. Beetle elytra can be involved in a very broad spectrum of functions: mechanical protection of hind wings and body, anti-predator strategies, thermoregulation and water saving, water harvesting, flight, hind wing folding, diving and swimming, self-cleaning and burrow cleaning, phoresy of symbiotic organisms, mating and courtship, and acoustic communication. We postulate that the potential of the elytra to take over multiple tasks has enormously contributed to the unparalleled diversification of beetles.

Ants are highly polyphenic Hymenoptera, with at least three distinct adult forms in the vast majority of species. Their sexual dimorphism, however, is overlooked to the point of being a nearly forgotten phenomenon. Using a multimodal approach, we interrogate the near total head microanatomy of the male of Dorylus helvolus, the "sausagefly," and compare it with the conspecific or near-conspecific female castes, the "driver ants." We found that no specific features were shared uniquely between the workers and males to the exclusion of the queens, indicating independence of male and worker development; males and queens, however, uniquely shared several features. Certain previous generalizations about ant sexual dimorphism are confirmed, while we also discover discrete muscular presences and absences, for which reason we provide a coarse characterization of functional morphology. Based on the unexpected retention of a medial carinate line on the structurally simplified mandible of the male, we postulate a series of developmental processes to explain the patterning of ant mandibles. We invoke functional and anatomical principles to classify sensilla. Critically, we observe an inversion of the expected pattern of male-queen mandible development: male Dorylus mandibles are extremely large while queen mandibles are poorly developed. To explain this, we posit that the reproductive-limited mandible phenotype is canalized in Dorylus, thus partially decoupling the queen and worker castes. We discuss alternative hypotheses and provide further comparisons to understand mandibular evolution in army ants. Furthermore, we hypothesize that the expression of the falcate phenotype in the queen is coincidental, that is, a "spandrel," and that the form of male mandibles is also generally coincidental across the ants. We conclude that the theory of ant development and evolution is incomplete without consideration of the male system, and we call for focused study of male anatomy and morphogenesis, and of trait limitation across all castes.

Platypsyllus castoris is closely associated with beavers and displays a unique set of structural specializations. We document the morphology of adults with modern techniques, and interpret evolutionary changes linked with the specific life style. The small subfamily Platypsyllinae has evolved an entire suite of features correlated with a more or less close association with mammals, for instance a flattened body, a dorsal cephalic shield, flightlessness, eye reduction, and depigmentation. Within this small group, Platypsyllus displays numerous autapomorphic features, correlated with a close association with the beaver. Essential is a combination of mechanical stabilization and firm anchorage on the host, and efficient forward movement in the fur. Exo- and endoskeletal structures of the head and thorax are reinforced by vertical cuticular columns and by an array of internal ridges. The antennae are shortened and strongly modified, the mandibles distinctly reduced and flattened, unsuitable for cutting, scraping or grinding. The musculature of the mouthparts is simplified, whereas an enhanced set of prepharyngeal and pharyngeal dilators forms an efficient sucking pump. The prothoracic musculature is strongly developed. In contrast, the pterothoracic muscle system is distinctly simplified, even though leg muscles are strongly developed. Using the legs, the flattened beetles move sideways through the dense fur of the beaver, using posteriorly directed groups of setae and ctenidia to prevent being pushed backwards by the densely arranged hairs. In contrast to the anterior body, the cuticle of the abdomen is thin, and the entire tagma flexible, with thin layers of segmental muscles. The hind gut is not connected with the mid gut. The beetles probably consume liquid, possibly with emulgated minute skin debris. As the morphology of the mouthparts excludes damage to the skin of the host, the association should not be addressed as ectoparasitic but as commensalism.

A new genus and species of the extinct beetle-like dictyopteran family †Umenocoleidae, †Laticephalana liuyani gen. et sp. nov., is described based on a well-preserved adult female from mid-Cretaceous Kachin (Myanmar/Burmese) amber. The new genus is characterized by two conspicuous autapomorphies, a triangular posterior extension of the head capsule and four large protuberances of the pronotum. Both structures can be interpreted as mechanic protective devices, but alternative/additional interpretations are possible. The large female subgenital plate is an autapomorphy of Dictyoptera. The transverse pronotal furrow suggests a placement in a clade with †Alienopteridae and Mantodea. A close relationship between †Laticephalana and †Enervipraela is tentatively supported by a very narrow pronotum and relatively short antennae. The very broad head of †Laticephalana with widely separated large compound eyes indicates predaceous habits, and the slender body and well-developed hindwings good flying abilities. The specializations of the distal legs, very similar to the conditions found in †Alienopteridae, suggest a preference for the foliage of trees and shrubs.

External and internal structures of the thorax of the myrmecophile beetle Claviger testaceus (Clavigeritae, Pselaphinae) were examined and documented with state-of-the-art visualization techniques. Following a general trend in the omaliine lineage (Staphylinidae), the skeletal elements of the pro- and pterothorax in Claviger reach a maximum degree of compactness, with largely reduced inter- and intrasegmental sutures and skeletal elements linked with the flight apparatus. The musculature, especially metathoracic direct and indirect flight muscles, also shows a high degree of reduction. Two forms of wings were found among individuals of C. testaceus, both non-functional and representing an advanced stage of reduction. However, that wing vestiges are still present and the metanotum, only slightly reduced, suggests that loss of flight in this species is likely the result of a young evolutionary process. Several structures are linked with myrmecophilous habits: small body size facilitates transportation of beetles by ant workers and makes it easier to move inside nest tunnels; the remarkably compact body and mechanically robust appendages make the beetles less vulnerable to attacks by ant mandibles; the improved elytral interlocking mechanism and unusually expanded epipleura enhance the protection of vulnerable dorsal parts of the pterothorax and anterior abdomen; and glands associated with trichomes on the posterolateral elytral angle produce secretions attractive for ants. Various modifications of the thorax and anterior abdomen lead to an optimization of intimate associations with ants. The morphological syndrome enabling these beetles to cope with life in ant colonies evolved in several steps. This is suggested by an increasing solidification of the thoracic skeleton in related non-myrmecophilous groups and also by less modified related clavigerites;for instance, ant-associated tropical species are still able to fly. Introduction Materials and methods - Studied species - Light microscopy - Micro-computed tomography (μCT) - Scanning electron microscopy (SEM) - Terminology Results - Prothorax - Pterothorax - Anterior abdominal segments Discussion - Phylogenetic background - Musculature - Elytral locking system and flightlessness - Myrmecophilous habits Conclusions

Clavigeritae is a group of obligate myrmecophiles of the rove beetle subfamily Pselaphinae (Staphylinidae). Some are blind and wingless, and all are believed to depend on ant hosts through feeding by trophallaxis. Phylogenetic hypotheses suggest that their ancestors, as are most pselaphines today, were free-living predators. Morphological alterations required to transform such beetles into extreme myrmecophiles were poorly understood. By studying the cephalic morphology of Claviger testaceus, we demonstrate that profound changes in all mouthpart components took place during this process, with a highly unusual connection of the maxillae to the hypopharynx, and formation of a uniquely transformed labium with a vestigial prementum. The primary sensory function of the modified maxillary and labial palps is reduced, and the ventral mouthparts transformed into a licking/'sponging' device. Many muscles have been reduced, in relation to the coleopteran groundplan or other staphylinoids. The head capsule contains voluminous glands whose appeasement secretions are crucial for the beetle survival in ant colonies. The brain, in turn, has been shifted into the neck region. The prepharyngeal dilator is composed of an entire series of bundles. However, the pharynx does not show any peculiar adaptations to taking up liquid food. We demonstrate that far-reaching cephalic modifications characterize C. testaceus, and that the development of appeasement glands and adaptation of the mouthparts to trophallaxis determine the head architecture of this extreme myrmecophile.

A newly discovered adult beetle †Lepiceratus ankylosaurus gen. & sp. nov. (mid-Cretaceous Myanmar amber) is the first extinct member of the suborder Myxophaga to reveal fine morphological structure...

The Entomophthoromycotina, a subphylum close to the root of terrestrial fungi with a bias toward insects as their primary hosts, has been notoriously difficult to categorize taxonomically for decades. Here, we reassess the phylogeny of this group based on conserved genes encoding ribosomal RNA and RNA polymerase II subunits, confirming their general monophyly, but challenging previously assumed taxonomic relationships within and between particular clades. Furthermore, for the prominent, partially human-pathogenic taxon Conidiobolus, a new type species C. coronatus is proposed in order to compensate for the unclear, presumably lost previous type species C. utriculosus Brefeld 1884. We also performed an exhaustive survey of the broad host spectrum of the Entomophthoromycotina, which is not restricted to insects alone, and investigated potential patterns of co-evolution across their megadiverse host range. Our results suggest multiple independent origins of parasitism within this subphylum and no apparent co-evolutionary events with any particular host lineage. However, Pterygota (i.e., winged insects) clearly constitute the most dominantly parasitized superordinate host group. This appears to be in accordance with an increased dispersal capacity mediated by the radiation of the Pterygota during insect evolution, which has likely greatly facilitated the spread, infection opportunities, and evolutionary divergence of the Entomophthoromycotina as well.

One hundred seven external and internal characters of larval and adult representatives of 28 genera of the coleopteran suborders were analyzed cladistically. Four groups of Neuropterida were introduced as outgroup. The analysis yielded 18 trees with a minimum of 194 steps (CI 0.691). All trees support the monophyly of all four suborders and a branching pattern (Archostemata + (Adephaga + (Myxophaga + Polyphaga))). The presence of elytra with meso- and metathoracic locking devices, the specific hind-wing folding, the close connection of exposed sclerites, the absence of the mera, the absence of eight thoracic muscles, the reduced abdominal sternite I, and the invagination of terminal segments are autapomorphies of Coleoptera. The monophyly of Coleoptera excl. Archostemata is supported by further transformations of the thoracic sclerites such as absence of the mesothoracic discriminal line and katepisternal joint, by an internalized or absent metathoracic trochantin, by the presence of a bending zone in the hind-wing, and by eight further muscle losses. Fusion of tibia and tarsus and presence of a single claw are larval synapomorphies of Myxophaga and Polyphaga. Adults are characterized by fusion of protrochantin and propleura and by the rigid connection of the meso- and metathoracic ventrites. The eucinetoid lineage of Polyphaga is characterized by the secondary absence of the bending zone of the alae. This results in a distinctly simplified wing folding mechanism. The monophyly of Cucujiformia (+ Bostrichoidea) is supported by the presence of cryptonephric Malpighian tubules. Transformations of fore-and hind-wings, reinforcement and simplification of the thoracic exoskeleton, and an efficient use of a distinctly reduced set of thoracic muscles play an important role in the early evolution of Coleoptera. Many different larval character transformations take place in the earlier Mesozoic within the suborders.

Thoracic structures of Tetraphalerus bruchi are described in detail. The results were compared with features found in other representatives of Archostemata and other coleopteran suborders. Differences between thoracic structures of Tetraphalerus and members of other archostematan subgroups are discussed. External and internal characters of larval and adult representatives of 37 genera of the coleopteran suborders are outlined, coded and analysed cladistically, with four groups of Neuropterida as outgroup taxa. The results strongly suggest the branching pattern Archostemata + [Adephaga + (Myxophaga + Polyphaga)]. Coleoptera excluding Archostemata are supported with a high Bremer support. Important evolutionary changes linked with this branching event are simplifications of the thoracic skeleton resulting in reduced degrees of freedom (i.e. a restricted movability, especially at the leg bases), and a distinct simplification of the muscle system. This development culminates in Polyphaga, which are also strongly supported as a clade. Internalization of the partly reduced propleura, further muscle losses, and the fusion of the mesoventrites and metaventrites—with reversal in Scirtoidea and Derodontidae—are autapomorphies of Polyphaga. Archostemata is a small relict group in contrast to highly successful xylobiontic groups of Polyphaga. The less efficient thoracic locomotor apparatus, the lack of cryptonephric Malpighian tubules, and the rise of angiosperms with beetle groups primarily adjusted to them may have contributed to the decline of Archostemata. © The Willi Hennig Society 2008.

An adult male of a newly discovered strepsipteran species from Brazil—Bahiaxenos relictus—is described. A new family Bahiaxenidae is suggested based on cladistic analyses of comprehensive morphological data sets with a broad taxon sampling including the stem group. It is unambiguously placed as the sister group of all other extant families of Strepsiptera. Bahiaxenos relictus is the only species of basal, i.e. non-stylopidian, Strepsiptera occurring in the New World. It appears to be a relict taxon that has survived in the fossil sand dunes of the Sao Francisco River (Bahia State). The loss of the 8th antennomere and the greatly reduced labrum are autapomorphies of Strepsiptera s.s. excluding Bahiaxenidae. The sister group relationship between†Protoxenidae and the remaining Strepsiptera, and between †Cretostylops and a clade comprising †Mengea and Strepsiptera s.s., is confirmed, as is the monophyly of Stylopidia and Stylopiformia.

We present the largest morphological character set ever compiled for Holometabola. This was made possible through an optimized acquisition of data. Based on our analyses and recently published hypotheses based on molecular data, we discuss higherlevel phylogeny and evolutionary changes. We comment on the information content of different character systems and discuss the role of morphology in the age of phylogenomics. Microcomputer tomography in combination with other techniques proved highly efficient for acquiring and documenting morphological data. Detailed anatomical information (356 characters) is now available for 30 representatives of all holometabolan orders. A combination of traditional and novel techniques complemented each other and rapidly provided reliable data. In addition, our approach facilitates documenting the anatomy of model organisms. Our results show little congruence with studies based on rRNA, but confirm most clades retrieved in a recent study based on nuclear genes: Holometabola excluding Hymenoptera, Coleopterida (= Strepsiptera + Coleoptera), Neuropterida excl. Neuroptera, and Mecoptera. Mecopterida (= Antliophora + Amphiesmenoptera) was retrieved only in Bayesian analyses. All orders except Megaloptera are monophyletic. Problems in the analyses are caused by taxa with numerous autapomorphies and ⁄ or inapplicable character states due to the loss of major structures (such as wings). Different factors have contributed to the evolutionary success of various holometabolan lineages. It is likely that good flying performance, the ability to occupy different habitats as larvae and adults, parasitism, liquid feeding, and co-evolution with flowering plants have played important roles. We argue that even in the ‘‘age of phylogenomics’’, comparative morphology will still play a vital role. In addition, morphology is essential for reconstructing major evolutionary transformations at the phenotypic level, for testing evolutionary scenarios, and for placing fossil taxa. � The Willi Hennig Society 2010.

The earliest branching event in winged insects, one of the core problems regarding early insect evolution, was addressed using characters of the head. The head is arguably one of the most complex body regions in insects and the phylogenetic information content of its features has been demonstrated. In contrast, the wings and other body parts related to the flight apparatus and sperm transmission are not useful in the context of this problem, as the outgroups (silverfish and bristletails) are wingless and transmit spermatophores externally. Therefore, they show profound differences in the organization of the postcephalic body, and assessment of homology and subsequent phylogenetic analysis of features of these body regions is extremely difficult. The core of this study is the investigation of head structures of representatives of the major clades of dragonflies. A detailed description of the head of Lestes virens is presented and was used as a starting point for the compilation of a character set and a character state matrix for the entire Dicondylia (winged insects + silverfish), with a main focus on the placement of dragonflies and consequently the basal branching event within winged insects. Our results indicate a sister-group relationship between a clade Palaeoptera (dragonflies + mayflies) and the megadiverse monophyletic lineage Neoptera. We show that despite considerable structural similarity between the odonate and neopteran mandible, the muscle equipment in dragonflies is more plesiomorphic with respect to Dicondylia than previously known. Odonata and Ephemeroptera also share presumably derived features of the antenna, maxilla, and labial musculature. Parsimony analyses of the head data unambiguously support a clade Palaeoptera. � The Willi Hennig Society 2012.

The systematic position of the Cretaceous genus †Umenocoleus Chen & Tan is re-evaluated. The re-assignment in 2014 back to Coleoptera is rejected based on numerous morphological features incompatible with such a placement. This includes an orthognathous head, multi-segmented antennae, the lack of a cephalic neck region, the exposed cervical membrane, a dorsal surface with cup-shaped punctures but without tubercles, moderately sclerotized forewings with rounded apices and lacking window punctures, the lack of an elevated mesoscutellar shield, and unfolded hind wings with numerous veins. Supposedly ancestral coleopteran features are implausible considering the comparatively young age (Early Cretaceous). As previously suggested, the genus is phylogenetically closely related to “roachoids” like †Ponopterix Vršanský & Grimaldi and †Petropterix Vršanský (†Umenocoleoidea Chen & Tan; see Vršanský, 2003) and probably also to †Alienopteridae (†Alienoptera).

Structural features and life habits of described species of the extinct †Alienoptera are evaluated based on previously published studies on the group. Head structures and feedings habits are addressed, as are the locomotor organs, especially the wings and adhesive devices. Suggested pollen feeding habits and the possible role as pollinators are discussed, as well as hypothesized ant and wasp mimicry and myrmecophily. Species of †Alienoptera were likely predators, in the case of †Caputoraptor elegans Bai, Beutel et Wipfler, 2018 with a unique cephalo-prothoracic prey grasping mechanism. They were likely strong fliers with anatomical dipterism with functional hind wings. Wing joints protected by scale-like sclerotized fore wings probably allowed them to move very efficiently in dense foliage of trees or shrubs and to prey upon smaller insects. Ant mimicry, myrmecophily and “weevil mimicry” are rejected. †Meilia Vršanský et Wang, 2018 is a possible case of wasp mimicry but more evidence is required. Other suggested cases of mimicking wasps are unfounded.

The Carboniferous insect taxon †Stephanastus polinae was assigned to a monotypic new order †Skleroptera, which was postulated as the sister group of the clade Strepsiptera + Coleoptera (= Coleopterida). The placement of †Stephanastus in Holometabola and the proposed relationship with Coleopterida are very likely unjustified. Suggested shared derived features of Skleroptera + Coleopterida are vague, insufficiently documented (characters of proximal leg segments) or likely based on misinterpretation (wing veins). Based on the wing venation we suggest a position in the extinct hemimetabolous order †Protelytroptera.

The morphology of mature larvae of Sisyra nigra was studied and documented with a broad spectrum of techniques. Special emphasis is on the cephalic anatomy and on the digestive tract. Cephalic structures are highly modified, with numerous autapomorphic conditions, including a globular head capsule, an extended area with large cornea lenses, a massive tentorium, a strongly developed prepharyngeal pumping apparatus with a horizontal arrangement of dilators, a sharp bend between the prepharynx and pharynx, and an unusual filter apparatus at the entrance of the large crop. The thoracic and abdominal muscle sets, and the legs are largely unmodified. Postcephalic apomorphies are conspicuous tergal setiferous tubercles, trifid setiferous pleural projections, single pretarsal claws, zigzag-shaped abdominal tracheal gills, and a dense vestiture of setae on the terminal abdominal segments. Mandibulo-maxillary stylets curved outwards are an unusual apomorphy also found in the semiaquatic larvae of Osmylidae. Semiaquatic or aquatic habits and secondarily multisegmented antennae are potential synapomorphies of these two groups and Nevrorthidae (Osmyloidea). A sistergroup relationship between Sisyridae and Nevrorthidae suggests that fully aquatic habits of larvae may be a synapomorphy of both families. A specialized terminal antennal seta is a potential groundplan apomorphy of Neuroptera, with secondary loss in Nevrorthidae and Ithonidae + Myrmeleontiformia, respectively. A trumpet-shaped empodium is likely an apomorphy of Neuroptera excluding Coniopterygidae and Osmyloidea, and the secondary loss an apomorphy of Ithonidae on one hand, and Myrmeleontiformia excl. Psychopsidae on the other.

†Tunguskagyrus, a Late Permian beetle fossil, was assigned to the adephagan family Gyrinidae in the original description. This placement was challenged in a recent study, where the genus was considered as part of a broad new concept of the extinct family †Triaplidae and transferred from Adephaga to Archostemata. Even though some structural features deviate from the typical pattern found in extant Gyrinidae, we maintain the placement of †Tunguskagyrus in the stemgroup of this family as the most plausibe interpretation. The suggested taxonomic changes concerning †Triaplidae (and †Schizophoroidea) are invalidated by an insufficient phylogenetic approach and several misinterpretations of the fossil imprints. The arguments are based on 1) general habitual similarity, 2) ubiquitous characters, 3) plesiomorphies, or 4) apomorphic conditions (compared to the Coleopteran groundplan) occurring in many beetle groups, strongly affected by homoplasy. The taxonomic shift of †Tunguskagyrus and the new concept of †Schizophoroidea and †Triaplidae are discarded due to the lack of a defined phylogenetic methodology and synapomorphies.

Mengenillidae is a small, basal family of Strepsiptera, mainly characterized by free-living females in contrast to the endoparasitic females of Stylopidia. Here, we describe external and internal structures of the female abdomen of Eoxenos laboulbenei (Mengenillidae). The external morphology was examined and documented using microphotography. Internal structures were reconstructed three-dimensionally using a μCT-data set. The morphologically simplified abdomen comprises 10 segments. The integument is weakly sclerotized and flexible. Spiracles are present dorsolaterally on segments I-VII. Segment VII bears the posteroventral birth opening and the small abdominal segment X carries the anus at its apex. Numerous eggs float freely in the hemolymph. The musculature of segments I-IV is composed of ventral and dorsal longitudinal muscle bundles, strongly developed paramedial dorsoventral muscles and a complex meshwork of small pleural muscles, with minimal differences between the segments. Segments V-X contain more than 50 individual muscles, even though the musculature as a whole is weakly developed. Even though it is not involved in processing food, the digestive tract is well-developed. Its postabdominal section comprises a part of the midgut and the short hindgut. The midgut fills a large part of the postabdominal lumen. The lumina of the midgut and hindgut are not connected. Five or six nodular Malpighian tubules open into the digestive tract at the border region between the midgut and hindgut. The birth organ below the midgut releases the primary larvae after hatching via the birth opening at segment VII. It is likely derived from primary female genital ducts. The presence of six additional birth organs of segments I-VI are de novo formations and a groundplan apomorphy of Stylopidia, the large strepsipteran subgroup with endoparasitic females. The loss of the primary birth organ of segment VII is an apomorphy of Stylopiformia (Stylopidia excl. Corioxenidae).