The morphology of the larval head of Tipulidae (Diptera, Insecta) - The dipteran groundplan and evolutionary trends

External and internal head structures of the larva of Tipula montium are described in detail. The results are compared to conditions found in other representatives of Tipuloidea and other dipteran and antliophoran lineages. Despite of the conceivably basal position of Tipulomorpha within Diptera, the larvae are mainly characterised by derived features. The partially retracted head, the specific hemicephalic condition and several other derived character states support the monophyly of Tipuloidea. A clade comprising Tipuloidea excluding Pediciidae is suggested by the strongly retracted head, by deep dorsolateral incisions of the head capsule, by a distinctly toothed anterior premental margin, by the loss of the second extrinsic maxillary muscle, and possibly by the loss of the pharyngeal filter. Eriopterinae and Hexatominae are characterised by a tendency towards an extreme reduction of the head capsule. Limoniinae, Cylindrotomidae, and Tipulidae form a clade supported by the presence of a premaxillary suture. This implies the non-monophyly of Limoniidae. A feature shared by Cylindrotomidae and Tipulidae is the presence of a movable lacinia mobilis. However, this is arguably a plesiomorphic feature, as it also occurs in Nannochoristidae. Features of the larval head of Trichoceridae, which were included in Tipulomorpha, do not show affinities with those of Tipuloidea. Trichocerid larvae share a specialised subdivided mandible with larvae of psychodomorph groups. Tipuloidea are a highly specialised group. The characters examined did not reveal plesiomorphic features supporting a basal position, and features suggesting closer affinities with Brachycera are vague. The evolution of dipteran larval head structures was apparently strongly affected by the loss of legs and the tendency to live in cryptic habitats. Diptera are the group of Endopterygota with the highest number of apomorphic features of the larval head. The appendages are generally simplified and the muscular apparatus is strongly reduced. Specialised features evolving within dipteran lineages include specifically arranged brushes of hairs on the labrum and epipharynx, movable messores, subdivided mandibles, different mandibular brushes, and a far-reaching reduction of labial parts.     

The larval head morphology of Xyela sp. (Xyelidae, Hymenoptera) and its phylogenetic implications

Larval head structures of Xyela sp. are described in detail. The characters are compared to conditions found in larvae of other groups of Hymenoptera and Endopterygota. Like other symphytan larvae the immature stages of Xyelidae are mainly characterized by presumably plesiomorphic features of the head. The head sutures are well developed and all parts of the tentorium are present. The labrum is free and a complete set of labral muscles is present. The maxillae are in a retracted position. In contrast to other hymenopteran larvae Xyela possesses a clypeofrontal suture, a comparatively long antenna and three well‐developed antennal muscles. Apomorphic features of Xyela are the absence of muscles associated with the salivarium and the complete absence of Musculus craniocardinalis. A clade comprising Orussidae and Apocrita is supported by the unsegmented maxillary and labial palps and the absence of the lacinia. Six potential autapomorphies for the Hymenoptera were revealed: (1) the caudal tentorial apodeme, (2) the bifurcated tendon of Musculus craniomandibularus internus, (3) the lateral lobe of the cardo, (4) the origin of M. tentoriohypopharyngalis from the posterior head capsule, (5) the exceptionally strong prepharyngo‐pharyngeal longitudinal muscle and (6) the longitudinal muscle of the silk press. The maxillolabial complex, the vestigial M. craniocardinalis and a distinctly developed labio‐hypopharyngeal lobe bearing the opening of the salivary duct are potential synapomorphies of Hymenoptera and Mecopterida. The globular, orthognathous head capsule, the modified compound eyes, the occipital furrow and the X‐shaped tentorium are features with unclear polarity shared by Hymenoptera and Mecoptera.     

The larval head of Agathiphaga (Lepidoptera, Agathiphagidae) and the lepidopteran ground plan

Abstract. The larval head of Agathiphaga vitiensis is described. There is a complete hypostomal bridge but no hypostomal ridges. Adfrontal ridges and distinct ecdysial lines are absent. There are two vestigial stemmata (without lenses) on each side. The antenna is one-segmented. All ‘typical lepidopteran’ head setae have been identified. The corporotentorium is very slender; dorsal tentorial arms are present. Intrinsic labral muscles are lacking. The mandible has retained a tentorial muscle. The maxilla is without a discrete cardo and has but a single endite lobe; ‘intrinsic maxillary muscles’ and the ‘cranial flexor of the dististipes’ are lacking. The postlabium is undivided and without setae, the labial palp is one-segmented and the lateral prelabio-hypopharyngeal sclerotization is continued into an oral arm. Some of the ventral pharyngeal dilators arise on the tentorium; mouth-angle retractors and dorsal post-cerebral pharynx dilators are absent. The two brain lobes have almost parallel long axes and are united by a narrow (almost pure neuropile) bridge. The corpora cardiaca and callata are contiguous. The aorta is an open gutter in front of the retrocerebral complex. Available evidence on the ground plan structure of the lepidopteran larval head is reviewed. The ancestral head supposedly was prognathous and was autapomorphic in having the cranio-cardinal articulation far behind the mandible; it had a complete hypostomal bridge but neither hypostomal nor adfrontal ridges, its tentorium was probably stout and with dorsal arms. Paulus & Schmid (1978, Z. zool. Syst. EvolForsch. 16) described a lepidopteran/trichopteran synapomorphy in stemma structure. A tentative table of homologies between cranial setae in Lepidoptera and Trichoptera is presented; it differs considerably from the scheme of Williams & Wiggins (1981, Proc. 3rd Symp. Trichopt.). The mouth parts and their musculature must have been overall very primitive for a panorpid larva, but the number of maxillary palp segments was reduced (three). The ‘dististipes’sensu Hinton is considered to consist of complexly fused parts of the stipes and basal palp segments. The cephalic stomodaeum must have possessed all primitive groups of extrinsic muscles. The incomplete available information on Micropterigidae impedes reconstruction of some details of the lepidopteran ground plan. Larval head structures support the monophyly of an entity comprising the Agathiphagidae + Heterobathmiidae + Glossata. There is one suite of derived characters shared by Heterobathmiidae and Agathiphagidae only and another shared by Heterobathmiidae and the Glossata only; one of these must represent parallelisms.     

The function and phylogenetic implications of the tentorium in adult Neuroptera (Insecta)

Despite several recent analyses on the phylogeny of Neuroptera some questions still remain to be answered. In the present analysis we address these questions by exploring a hitherto unexplored character complex: the tentorium, the internal cuticular support structure of the insect head. We described in detail the tentoria of representatives of all extant neuropteran families and the muscles originating on the tentorium using 3D microCT images and analyzed differences in combination with a large published matrix based on larval characters. We find that the tentorium and associated musculature are a source of phylogenetically informative characters. The addition of the tentorial characters to the larval matrix causes a basad shift of the Sisyridae and clearly supports a clade of all Neuroptera except Sisyridae and Nevrorthidae. A sister group relationship of Coniopterygidae and the dilarid clade is further corroborated. A general trend toward a reduction of the dorsal tentorial arms and the development of laminatentoria is observed. In addition to the phylogenetic analysis, a correlation among the feeding habits, the development of the maxillary muscles, and the laminatentoria is demonstrated.     

First μ‐CT‐based 3D reconstruction of a dipteran larva—the head morphology of protanyderus (tanyderidae) and its phylogenetic implications

The larval head of Protanyderus was examined and documented using innovative techniques, with emphasis on internal structures. A chart listing all head muscles of dipteran larvae and other holometabolan groups is presented in the Supporting Information. The results are compared to conditions found in other nematoceran lineages. The larval head of Protanyderus is characterized mainly by plesiomorphic character states such as the complete and largely exposed head capsule, the long coronal suture, V‐shaped frontal sutures, lateral antennal insertion areas, a transverse labrum, a nearly horizontal plane of mandibular movements, mandibles lacking a movable distal part, a mesal hook and mesal or distal combs, separated maxillary endite lobes, a comparatively complete array of muscles, and a brain only partly located within the head capsule. An anteriorly toothed hypostomal plate and dense labral brushes of microtrichiae are also likely groundplan features of Diptera. The pharyngeal filter is a possible apomorphy of Diptera excl. Deuterophlebiidae (or Deuterophlebiidae + Nymphomyiidae). The messors have also likely evolved early in the dipteran crown group but are absent in the groundplan. The phylogenetic interpretation of externolateral plates with growth lines is ambiguous. Autapomorphies of Tanyderidae are differences between the third and fourth instar larvae, the roof‐like extension above the antennal insertion area, the dorsal endocarina, and the posterodorsal internal ridge. The phylogenetic position of Tanyderidae is controversial, but features of the larval head do not support a proposed sistergroup relationship between Tanyderidae and Psychodidae. Both groups differ in many features of the larval head, and we did not identify a single potential synapomorphy. Larval characters alone are insufficient for a reliable phylogenetic reconstruction, though they vary greatly and apparently contain phylogenetic information. The evaluation of these features in the context of robust molecular phylogenies will be a sound basis for the reconstruction of complex evolutionary scenarios for the megadiverse Diptera. Diptera. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Head morphology of <Emphasis Type="Italic">Osmylus fulvicephalus</Emphasis> (Osmylidae,Neuroptera) and its phylogenetic implications

External and internal head structures of Osmylus fulvicephalus were examined and described in detail. Exo- and endoskeleton, musculature, elements of the central nervous system and tracheae are compared to conditions found in other groups of Neuropterida and other endopterygote lineages. Thirty-six adult cephalic characters were compiled, combined in a datamatrix with 64 characters of the larval head, and analysed cladistically. Mainly because many data on adults remain missing, most branches in the cladogram are mostly or exclusively supported by larval features. The shortening of the mesal mandibular wall and the resulting anterior shift of the adductor tendon possibly constitute an adult groundplan apomorphy of Neuropterida. Raphidioptera and Megaloptera share distinct prognathism and the presence of a sclerotised gula. However, the orthognathous head and the absence of a gula resulted as autapomorphies of Neuroptera in our analyses. Further potential autapomorphies are the asymmetry of the mandibles as well as the respective presence of dorsolateral furrows on the head capsule, of a shovel-like extension on the ventral mandibular cutting edge, and of a row of stiff hairs on the mandible’s ventral surface. The systematic affinities of Osmylidae remain ambiguous. Osmylus is mainly characterised by plesiomorphic features of the adult head such as a complete endoskeleton, long filiform antennae, largely unmodified orthopteroid mouthparts, and particularly the nearly complete set of muscles. The placement with a clade also comprising Hemerobiidae and Chrysopidae is poorly supported. The presence of a dense vestiture of long microtrichia on the distal galeomere resulted as a synapomorphy of the three families. An apparent plesiomorphy preserved in Osmylus but absent in all other groups of Neuroptera is the presence of well developed ocelli. The present study underlines the severe shortage of detailed morphological data on the adults. Intensive study of adult structures is required for a solid reconstruction of the phylogeny of Neuropterida, especially of the hemerobiform lineage of Neuroptera.     

The larval head of Nevrorthidae and the phylogeny of Neuroptera (Insecta)

External and internal head structures of larvae of Nevrorthidae were described in detail. The results were compared to conditions found in other representatives of Neuroptera and the other two neuropterid orders. The cladistic analysis supported the monophyly of Neuroptera, Neuroptera exclusive of Nevrorthidae, Hemerobiiformia, and Myrmeleontiformia. Neuroptera exclusive of Nevrorthidae are supported by the formation of an undivided postmentum and the presence of cryptonephric Malpighian tubules. The highly specialized articulation of the neck (Rollengelenk) and the absence of a salivary duct are autapomorphies of Nevrorthidae. Ithonidae and Polystoechotidae form a clade and are the sister group of the remaining Hemerobiiformia, which are characterized by the complete lack of a gula and a terminal filament of the antenna. Within this lineage, a clade comprising Mantispidae, Dilaridae, Berothidae, and Rhachiberothidae is well supported. Larvae of Myrmeleontiformia are characterized by a complex transformation of head structures, with a hypostomal bridge, a small triangular gula, largely reduced maxillary grooves, and anteriorly shifted posterior tentorial grooves. The slender finger‐like mid‐dorsal apodeme is another autapomorphy of the group. Psychopsidae are placed as the sister group of the remaining Myrmeleontiformia, which are characterized by a conspicuous, protruding ocular region (often less distinct or even absent in Nemopteridae). Ascalaphidae are the sister group of Myrmeleontidae. Larvae of both families share the fusion of the tibia and tarsus in the hind leg. The larval characters analysed were not sufficient for full resolution of the myrmeleontiform and hemerobiiform lineages. The position of several families such as Osmylidae, Sisyridae, and Coniopterygidae remains uncertain. The results are in agreement with an aquatic ancestor of Neuroptera and secondarily acquired terrestrial habits within the lineage (Neuroptera exclusive of Nevrorthidae), and another invasion of the aquatic environment by Sisyridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 533–562.     

Head morphology of Caurinus (Boreidae, Mecoptera) and its phylogenetic implications

External and internal head structures of Caurinus dectes were examined and described in detail. The features are compared to conditions found in other groups of Antliophora. Caurinus is obviously crucial for the reconstruction of the mecopteran and antliophoran groundplan. It displays a remarkable series of plesiomorphic character states such as a complete clypeolabral suture, the presence of M. hypopharyngomandibularis (M. 13) and M. frontohypopharyngalis (M. 41), a subdivided clypeus, a short head without rostrum, a dorsal tentorial arm attached to the head capsule, the absence of a cranial dilator of the antenna, and large mandibles with a well developed apical tooth, two distinct subapical teeth, and a basal molar part. The first three plesiomorphic features render potential autapomorphies of Mecoptera in the traditional sense invalid. Autapomorphies of Caurinus are the distinctly flattened labrum, the absence of the labroepipharyngeal muscle, the very large size of M. 13, the strongly enlarged penultimate palpomeres, the partition of M. 41, the very strongly developed precerebral sucking chamber, strongly curved optic lobes, the presence of a large protocerebral extension in the genal region and deep posterior excavations of the protocerebrum. The maxillolabial plate, the absence of cardines as separate structures, the reduction of ocelli, and the origin of maxillary palp muscles on a median ridge or area of the maxillolabial plate are likely autapomorphies of Boreidae. Another potential autapomorphy of the family is the presence of longitudinal furrows on the mandibles. However, they are absent in Boreus. The thick strongly sclerotised, median ridge of the maxillolabial plate, the missing retractibility of the prementum, the absence of extrinsic labial muscles, and the presence of a median ridge on the prepharyngeal roof suggest a clade Boreus + Hesperoboreus. The origin of extrinsic maxillary muscles from the clypeus has probably evolved independently in Boreus and Hesperoboreus, and in Panorpa, respectively. The absence of M. craniolacinialis and the presence of a row of several subapical mandibular teeth are autapomorphies of Boreus. The presence of a specific intrinsic muscle of the salivary duct and a membranous galea enclosing the labrum and mandibular base are derived features shared by Boreidae and Pistillifera (galea absent in Nannochorista, Siphonaptera and Diptera). The loss of M. frontolabralis (M. 8) is a potential apomorphy of Mecoptera incl. Siphonaptera. A sister group relationship between Boreidae and Siphonaptera is not supported by characters of the adult head. Head structures of Siphonaptera are extremely modified in correlation with ectoparasitic habits.     

The larval head of Exechia (Mycetophilidae) and Bibio (Bibionidae) (Diptera)

Exechia and Bibio have retained several plesiomorphic groundplan features of Diptera and Bibionomorpha, including a fully exposed and sclerotized head capsule, the transverse undivided labrum, the absence of movable premandibles, and undivided mandibles without combs. The fusion of the hypostomal bridge with the head capsule and largely reduced antennae are derived features shared by both taxa. The absence of teeth at the anterior hypostomal margin is a potential autapomorphy of Bibionomorpha. A basal position of Anisopodidae is suggested by a number of plesiomorphies retained in this family. Apomorphies of Bibionomorpha excluding Anisopodidae are the reduction of tentorial elements, the partial fusion of the labrum and clypeus, one-segmented antennae, the absence of a separate submental sclerite, the loss of the labial palpus, and the reduction of the pharyngeal filter apparatus. Head structures of Bibio are largely unmodified. The subprognathous orientation is one of few autapomorphic features. In contrast, the mouthparts of Exechia are highly modified in correlation with the specialized food uptake. The rasping counterrotating movements of maxillae and mandibles with teeth oriented in opposite directions are carried out by strongly developed extensors and flexors of the paired mouthparts. The modified labium mechanically supports the “drill head” formed by the mandibles und maxillae. The necessary stability of the head capsule is provided by the hypostomal bridge which also compensates the far-reaching reduction of the tentorium.     

The larval head structures of Podagricomela shirahatai (Chûjô) (Chrysomelidae,Galerucinae, Alticini) and morphological effects of leaf mining

Head structures of the leaf mining larva of the chrysomelid species Podagricomela shirahatai are described and illustrated. Internal and external structures were reconstructed three dimensionally based on image stacks obtained with microcomputed tomography. The larval head is characterized by prognathism, a dorsoventrally compressed shape, a flattened maxillolabial complex, a completely reduced coronal suture, and the presence of a deep, V‐shaped posterior emargination of the head capsule. Internal structures are not distinctly affected by leaf mining. The cephalic features are compared with conditions found in surface feeding and root feeding alticine larvae and also with characters of chrysomeline larvae of Chrysomela populi Linnaeus. Possible correlations between modifications of the larval head and different feeding behaviors are discussed. Characters are also discussed with respect to possible phylogenetic implications. Some derived features are apparently due to phylogenetic constraints. Apomorphies characterizing alticine larvae with distinctly different life habits are the loss of M. frontoepharyngalis (M. 9), the origin of M. tentoriostipitalis (M. 18) from the head capsule, two insertions of M. tentoriopraementalis inferior (M. 29) and the reduction of stemmata. The study underlines that the anatomical study of chrysomeloid larvae is not only highly desirable in a phylogenetic context, but also crucial for understanding the evolution of different life strategies in this extremely successful group of Coleoptera. J. Morphol. 276:446–457, 2015. © 2014 Wiley Periodicals, Inc.     

The cephalic morphology of the Gondwanan key taxon Hackeriella (Coleorrhyncha,Hemiptera)

External and internal head structures of Coleorrhyncha, a key-taxon within the Hemiptera, are described in detail and documented using modern techniques. The main focus is on Hackeriella veitchi, but two additional representatives of the Gondwanan relict group were also examined, and also head structures of Enicocephalidae, a member of a potentially basal heteropteran lineage. Features were compared to those documented in literature for the Sternorrhyncha, Auchenorrhyncha, and Heteroptera. Coleorrhyncha are characterized by highly modified head structures and correspondingly an entire series of autapomorphies, such as for instance a strongly flattened head capsule with fenestrations. However, they also display features that are likely plesiomorphic compared to members of other hemipteran groups. These include the almost complete tentorium and the lack of the gula. The sistergroup relationship between Coleorrhyncha and Heteroptera is well supported by cephalic features. Potential synapomorphies are the presence of a distinct mandibular sulcus, the reduced number of antennomeres, the absence of clasping organs in the labial groove, coiled accessory salivary ducts, the presence of a small cervical muscle M1a (M. pronotopostoccipitalis medialis), the presence of a second mandibular promotor M14 (M. zygomaticus mandibulae), the presence of M28 (M. verticopharyngalis), and M30 (M. frontobuccalis posterior).     

Development of the nidicolous tadpoles of Eupsophus emiliopugini (Anura: Cycloramphidae) until metamorphosis,with comments on systematic relationships of the species and its endotrophic developmental mode

Vera Candioti, M.F., Nuñez, J.J. and Úbeda, C. 2011. Development of the nidicolous tadpoles of Eupsophus emiliopugini (Anura: Cycloramphidae) until metamorphosis, with comments on systematic relationships of the species and its endotrophic developmental mode. —Acta Zoologica (Stockholm) 92 : 27–45. Species of Eupsophus are unique within Alsodinae in having nidicolous tadpoles. They are characterized by traits typical of generalized exotrophic (e.g., oral disc and spiracular tube) and endotrophic larvae (e.g., scant pigmentation and large hind limbs). The larval morphology and development of E. emiliopugini, including external, buccal, and musculoskeletal features, is described herein. Like the larvae of other alsodines, these larvae have four lingual and four infralabial papillae, quadratoethmoid process, and an m. rectus cervicis with a double insertion. Among the traits exclusive to the genus are: the absence of the pseudopterygoid process and quadrato‐orbital commissure; presence of the m. subarcualis rectus I with two slips; and presence of the m. subarcualis rectus II–IV inserting on Ceratobranchial II. The development and metamorphosis of Eupsophus include some characters that develop later (e.g., degeneration of mouthparts and chondrocranium with minimum calcification), characters that develop earlier (e.g., hind‐limb appearance and jaw and suspensorium ossification), and characters that develop at the same time (e.g., most external features and cranial muscles) than in most exotrophic species. Some distinctive characters (third lower labial ridge absent, general configuration of the hyobranchial skeleton, skeletal development with retention of larval traits) resemble those of other endotrophic species, and the precocious ossification of jaws and suspensorium is shared with several direct‐developing species among recent amphibians.     

Phylogeny of the Neuropterida (Insecta: Holometabola)

The Neuropterida, with about 6500 known species — living fossils in a way — at the base of the Holometabola (as a sister group of the Coleoptera), comprise Raphidioptera (about 210 species, two families), Megaloptera (about 300 species, two families) and Neuroptera (6000 species, 17 families). Megaloptera + Neuroptera is argued vs. the traditional Raphidioptera + Megaloptera. Raphidioptera are undisputedly monophyletic. Monophyly of Megaloptera is the operational hypothesis, although occasionally questioned. Sucking tubes of the larvae are the most spectacular autapomorphy of Neuroptera. The construction of larval head capsules indicates three evolutionary lines: Nevrorthiformia, and Myrmeleontiformia + Hemerobiiformia. Traditional Myrmeleontiformia is Psychopsidae + (Nemopteridae + (Nymphidae + (Myrmeleontidae + Ascalaphidae))), the present approach is (Psychopsidae + Nemopteridae) + all other Myrmeleontiformia. Hemerobiiformia are based on the ‘maxillary head’ concept. The ithonid clade Ithonidae/Rapismatidae + Polystoechothidae and the dilarid clade Dilaridae + (Mantispidae + (Rhachiberothidae + Berothidae)) are based on robust criteria. Other relationships remain unclear: Hemerobiidae + Chrysopidae (on similarity) and the ‘early offshoot’ concept of coniopterygidae (on autapomorphies) should not be perpetuated. Chysopidae + Osmylidae and (Hemerobiidae + (Coniopterygidae + Sisyridae)) + dilarid clade are discussed. Aquatic larvae, regarded as independent apomorphies of megaloptera and neuropteran Nevrorthidae and Sisyridae for a long time, are re‐interpreted as a synapomorphy of Megaloptera + Neuroptera and thus plesiomorphic within these groups. Terrestrial larvae (with cryptonephry to solve osmotic problems) are consequently apomorphic. Aquatic Sisyridae with cryptonephry of a single malpighian tubule, is conflicting, but larvae may have become secondarily aquatic, after a terrestrial intermezzo.     

Larval head anatomy of Heterogynis penella (Zygaenoidea, Heterogynidae), and a general discussion of caterpillar head structure (Insecta, Lepidoptera)

The internal head anatomy (and the peculiar integumental structure of the epicranial notch region) of Heterogynis penella larvae are described; special attention is paid to the skeleto‐muscular and nervous systems and to the cephalic glands. Transverse ligaments connect the apodemes of the mandibular adductor muscles of both sides and the anterior maxillo‐labial articulations of both sides. The two ligaments are linked to each other by a thin, apparently acellular membrane. An accessory, trilobed mandibular gland is present. A putative stretch receptor, connecting the oblique dorsal cibarial dilators of both sides, is described for the first time in a lepidopterous larva and its importance in assessing the homology of these muscles is discussed. The presence of cibarial sensilla, previously predicted in other caterpillars on the basis of behavioural experiments and observations of the nerve pattern, is confirmed. The structural diversity of larval head anatomy in ditrysian Lepidoptera is discussed, with particular emphasis on the innervation of the corpora cardiaca and corpora allata and of the sensilla of the head capsule.     

A longstanding entomological problem finally solved? Head morphology of Nannochorista (Mecoptera,Insecta) and possible phylogenetic implications

External and internal head structures of Nannochorista species were examined and described in detail. The characters are discussed with regard to their functional and phylogenetic implications. The structure of the mouthparts indicates that adults of Nannochorista feed on fluids. The loss of the mandibular muscles and the precerebral pharyngeal dilators are presumptive autapomorphies of the genus. A possible clade comprising Nannomecoptera, Siphonaptera and Diptera is supported by the presence of a labral food channel, the absence of the galea, a sheath for the paired mouthparts formed by the labium, very strongly developed labial palp muscles and cibarial dilators, and the presence of a well‐defined postcerebral pharyngeal pumping chamber. Closer affinities of Nannomecoptera with Diptera are suggested by the presence of a unique sensorial groove on the third maxillary palpomere. Further potential synapomorphies are the presence of a frontal apodeme and a primarily lamelliform mandible without teeth. The presence of a salivary channel on the laciniae and a subdivided labrum are shared derived features of Nannochorista and Siphonaptera. A derived condition present in Mecoptera including Boreidae but excluding Nannochoristidae is the secretion with a strongly developed intrinsic muscle of the salivary duct. The loss of the lateral labral retractor, the cranial muscle of the cardo, and of two of the three premental retractors, and the absence of transverse epipharyngeal muscles are potential autapomorphies of Antliophora. The formation of a maxillolabial complex is a possible synapomorphy of Hymenoptera and Mecopterida.