Cephalic morphology of Hymenopus coronatus (Insecta: Mantodea) and its phylogenetic implications

External and internal head structures of the mantodean Hymenopus coronatus are examined and described in detail. The results are elaborately compared with the literature. Strong crests on the anterior tentorial arms that articulate with the subantennal suture, a parietal suture and glossae and paraglossae with anteriorly bent tips are proposed as new potential apomorphies for Mantodea while a head capsule being wider than long, enlarged compound eyes, the presence of a frontal shield or scutellum, lateral lobes in the anterior tentorial arms, the presence of a transverse and an interantennal suture and the reduction of the mentum are confirmed as apomorphies, As potential apomorphies for Dictyoptera the reduction of Musculus tentoriobuccalis lateralis (M. 49) is newly presented and a “perforate” tentorium, lacinial incisivi that are located in a galeal pouch and the presence of a postmola are confirmed. The present study shows the value of cephalic morphology for phylogenetic analysis but also points out that further studies including evolutionary key taxa are essential for resolving the evolutionary adaptations among dictyopterans.     

An early winged crown group stick insect from the Early Eocene amber of France (Insecta,Phasmatodea)

The new stick insect family Gallophasmatidae, based on Gallophasma longipalpis gen. et sp.n. , from the Earliest Eocene French amber has a pattern of tegmina venation typical of Archaeorthoptera, also present in at least some Mesozoic ‘Phasmatodea’. On the other hand, Gallophasma displays in its body anatomy some apomorphies of the extant Euphasmatodea, e.g. fusion of metatergum and abdominal tergum 1, correlated with the reduction of abdominal sternum 1 to lateral triangular sclerites. A unique autapomorphy of Gallophasma is the presence of annulated and apparently multi‐segmented or pseudo‐segmented cerci; all other Phasmatodea have one‐segmented cerci. The venation of the tegmina of Gallophasma differs from that of extant winged Phasmatodea in the plesiomorphic absence of a knob‐like dorsal eversion. This and other differences in the wing venation between extant and extinct Phasmatodea might have been caused by the loss of wings at some point in the evolutionary history of the order and their secondary gain in a subclade of the extant phasmids.     

Stick insect on unsafe ground: does a fossil from the early Eocene of France really link Mesozoic taxa with the extant crown group of Phasmatodea?

The recently described Gallophasma longipalpis from Earliest Eocene French amber is considered to be a key fossil taxon that phylogenetically links ‘Mesozoic Phasmatodea’ with extant stick and leaf insects. However, our re‐evaluation of the evidence provided for this placement reveals that Gallophasma does not possess any unambiguous synapomorphies with extant forms, e.g. neither with Euphasmatodea nor with the more inclusive Phasmatodea. The fusion of abdominal segment 1 with the metathoracic segment, a derived character state present in both Gallophasma and Euphasmatodea, shows fundamental structural differences, and cannot be homologized between both taxa. We argue that the presence of a well‐developed, externally visible ovipositor and four‐segmented cerci in Gallophasma can be interpreted only as plesiomorphic with regards to all extant Phasmatodea, or even to Phasmatodea plus its putative sister groups Embioptera or Orthoptera. Gallophasma does not belong to the stem lineage of recent Phasmatodea, and is referred to best as ‘lower Neoptera’ or Polyneoptera incertae sedis. Therefore, this fossil may be central to reconstructing the ground pattern of the aforementioned orthopteroid lineages, and to determining wing character polarity within Polyneoptera.     

The spermatogenesis and sperm structure of Timema poppensis (Insecta: Phasmatodea)

The ultrastructure of spermatogenesis and spermatozoa was studied in Timema poppensis Vickery & Sandoval, 1999, a putative basal taxon of Phasmatodea. The apical portion of testis follicles consists of spermatogonial cells with polymorphic nuclei. Primary spermatocytes display very short primary cilia originating from the peripheral centrosomes. Early spermatids develop a conspicuous “nebenkern” consisting of fused mitochondria. They have a single peripheral centriole with microtubular triplets, which expresses a 3.6-μm-long cilium featuring a 9?+?2 axonemal pattern. In a later stage, the centriole and the ciliary shaft displace toward the inner part of the cytoplasm by an infolding of the plasma membrane. Mature spermatids exhibit a derived centriole with microtubule doublets devoid of dynein arms, which is equipped with a dense arc-like outer structure. Ciliary degeneration was not observed during spermiogenesis. Spermatozoa are short flagellate cells about 55–60?μm in length. They are characterized by a three-layered acrosomal complex. The distinctive bell-shaped morphology of the acrosome vesicle is likely an autapomorphic trait of Timema. The flagellum has a 9?+?9?+?2 axoneme, two accessory bodies, two flattened cisterns, and two elongated mitochondrial derivatives. Results support the hypothesis that Phasmatodea, comprising Timema?+?Euphasmatodea, form a monophyletic group. The presence of 17 protofilaments in the wall of accessory microtubules and the flattened configuration of the flagellum are potential apomorphic groundplan features of the order. Within Phasmatodea, a key evolutionary divergence was from the conventional insect spermiogenesis and sperm structure of Timema, to the unusual spermiogenetic process and peculiar sperm structure of Euphasmatodea. As a result, Timema retains more sperm character states found in the polyneopteran ground pattern, while Euphasmatodea have evolved outstanding sperm autapomorphies, like the loss of mitochondria and flattened cisterns, and the presence of strongly expanded accessory bodies.     

Ultra morphology of sensilla on the head and mouth parts of the earwig Anisolabis martima (Bonelli) (Dermaptera: Carcinophoridae)

The earwig Anisolabis maritima is a cosmopolitan insect with a worldwide distribution, omnivorous in feeding, eating the eggs of different insects. Recently recorded in Saudi Arabia as a beneficial predator on eggs and newly hatched larvae of the red palm weevil Rhyncophrous ferrugineus, scanning electron microscopy (SEM) observations revealed the presence of different kinds of receptor sensillae occurring on the head and mouth parts of A. maritime. The sensillae are the same, but differ in numbers on both sexes. The head surface bears three subtypes of trichoid sensillae, one type of ceoloconica. Labrum, labium, labial palps and maxillary palps, possess different kinds of sensillae. Three subtypes of trichoid have been identified on the labrum. The labium contained one type of tricoid and basiconic sensillae. There are great numbers and numerous sensillae cheatica differing in length distributed on the maxillary and the labial palps on all segments between other stutter spines. The last terminal segment of maxillary or labial palps bears the greatest number of the sensillae cheatica. Adult A. maritima may be introduced in the future as a beneficial predator of eggs of R. ferrugineus as a tool for integrated pest management.     

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 earliest Timematids in Burmese amber reveal diverse tarsal pads of stick insects in the mid‐Cretaceous

Many extant insects have developed pad structures, euplantulae or arolia on their tarsi to increase friction or enhance adhesion for better mobility. Many polyneopteran insects with euplantulae, for example, Grylloblattodea, Mantophasmatodea and Orthoptera, have been described from the Mesozoic. However, the origin and evolution of stick insects' euplantulae are poorly understood due to rare fossil records. Here, we report the earliest fossil records of Timematodea hitherto, Tumefactipes prolongates gen. et sp. nov. and Granosicorpes Urates gen. et sp. nov., based on three specimens from mid-Cretaceous Burmese amber. Specimens of Tumefactipes prolongates gen. et sp. nov. have extremely specialized and expanded euplantulae on their tarsomere II. These new findings are the first known and the earliest fossil records about euplantula structure within Phasmatodea, demonstrating the diversity of euplantulae in Polyneoptera during the Mesozoic. Such tarsal pads might have increased friction and helped these mid-Cretaceous stick insects to climb more firmly on various surfaces, such as broad leaves, wetted tree branches or ground. These specimens provide more morphological data for us to understand the relationships of Timematodea, Euphasmatodea, Orthoptera and Embioptera, suggesting that Timematodea might be monophyletic with Euphasmatodea rather than Embioptera and Phasmatodea should have a closer relationship with Orthoptera rather than Embioptera.     

On the evolution of adult head structures and the phylogeny of Hydraenidae (Coleoptera, Staphyliniformia)

External and internal head structures of adults of Orchymontiinae, Prosthetopinae, Hydraeninae and Ochthebiinae were studied and those of Ochthebius semisericeus and Limnebius truncatellus are described in detail. The results are evaluated with respect to their relevance for a reconstruction of hydraenid phylogeny and also compared with structural features found in adults of other staphyliniform families. The monophyly of Hydraenidae is supported by the presence of a plate‐like, trilobed premento‐hypopharyngeal extension, an unusual origin of m. tentoriohypopharyngalis, dorsal tentorial arms firmly fused with the head capsule, modified basal antennomeres, and palpigers connected by a transverse sclerotized bar. Orchymontiinae are monophyletic and the basal sister group of the remaining Hydraenidae. The presence of a ventral transverse genal bulge and of a pubescent antennal club with more than two antennomeres (reversal in some prosthetopines: e.g. Mesoceration abstrictum) are possible apomorphies of Hydraenidae excluding Orchymontiinae. Prosthetopinae are probably monophyletic and the sister group of Ochthebiinae + Hydraeninae. The latter clade is characterized by a distinct cupula formed by antennomere VI, a loose five‐segmented pubescent antennal club, and a modified antennal musculature. The presence of an unusual tentorio‐pharyngeal dilator is a shared derived feature of Ochthebiinae and the genus Davidraena. The monophyly of Ochthebiinae was confirmed and Ochtheosus is the sister group of the remaining ochthebiine genera, which are characterized by a perforated wall‐like structure formed by the posterior tentorial arms. The absence of this tentorial modification and the fimbriate galea are plesiomorphies retained in Ochtheosus. Calobius differs strongly from other subgenera of Ochthebius and a generic status may be appropriate. The monophyly of Hydraeninae is not supported. Hydraena was confirmed as a clade and Laeliaena and Limnebius are sister groups. The latter genus is characterized by several autapomorphies. The basal position of Orchymontiinae and Prosthetopinae suggests a Gondwanan origin of Hydraenidae and a primary preference for life in running water. Important evolutionary changes of head structures are complex transformations of the antennae and related structures. Yet, the use of the antennae as accessory breathing organs is not a groundplan feature of the family. The results of this study strengthen the case of staphylinoid affinities of Hydraenidae.     

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.     

The role of the antennae and maxillary palps in mediating food preference by larvae of the tobacco hornworm, Manduca sexta

The role of antennae and maxillary palps in mediating food preferences by Manduca sexta (Johan.) (Lepidoptera: Sphingidae) was examined. Fifth‐instar larvae, having different chemosensory organs remaining after microsurgery, were tested in two‐choice feeding assays using three solanaceous host species Lycopersicon esculentum (Mill.), Datura innoxia (Mill.), and Solanum pseudocapsicum (L.), an acceptable non‐host Raphanus sativus (L.) (Cruciferae), and an unacceptable non‐host Pelargonium hortorum (L.H. Bailey) (Geraniaceae). Larvae had a choice between leaf discs of two plant species or between one species and moist filter paper discs (water). The antennae are fully competent in mediating normal (unimpaired) food preferences for S. pseudocapsicum vs. water and P. hortorum vs. water. Thus, the antennae alone can mediate both acceptance and rejection behaviour. The latter is the first report of such a function in M. sexta. The antennae are partly competent (reduced preference) for S. pseudocapsicum vs. P. hortorum. No antennal competence could be demonstrated using the other plant species tested. The antennae alone are either partly needed (R. sativus vs. water) or not necessary to elicit normal food preferences for the plant species tested. The maxillary palps are fully competent in mediating normal food preference for S. pseudocapsicum vs. water, and D. innoxia vs. water. The palps are partly competent for S. pseudocapsicum vs. P. hortorum and incompetent for the other plant species tested. Thus, the maxillary palps alone could only mediate acceptance behaviour in this study. The need of the maxillary palps in normal food preferences could not be demonstrated. Interestingly, the maxillary palps alone can mediate food preferences for two normally rejected plants, R. sativus and P. hortorum. Similar results are reported for two other non‐host plant species, Vigna unguiculata (Walp.) (Leguminosae) and Vinca rosea (L.) (Apocynaceae). Perhaps, the maxillary palps inform M. sexta mostly about feeding stimulatory chemicals common to most plant species.     

Functional morphology and neuronal innervation of the prothoracic defence gland in Timema

Timema is the most basal genus of Phasmatodea and the sister group to the remaining stick and leaf-insects (Euphasmatodea). An autapomorphy of all phasmids is the paired prothoracic exocrine defence glands. In this study, the anatomy and innervation of the defence glands in Timema petita and Timema chumash are described and compared with the data on Euphasmatodea. In all phasmids, the glands consist of a cuticular epithelium, a secretory epithelium and muscular fibres that compress the lumen. In Timematodea, the muscular part of the gland is less developed than in Euphasmatodea and the ejection of the defence secretion depends on the dorsal longitudinal neck muscles. On the neuroanatomical level, Timema petita and Timema chumash lack neurons that are involved in the independent contraction of the gland in euphasmids. In both studied species of Timema, neck muscles play an active role in the gland function which is not observed in any other phasmid. Considering the basal position of this genus, this supports the hypothesis that in euphasmids, the muscular part of the gland evolved from the dorsal longitudinal neck muscles. Additionally, the same nerves that innervate the dorsal longitudinal neck muscles in all Polyneoptera also innervate the defence glands in phasmids.     

The cephalic morphology of the troglobiontic cholevine species Troglocharinus ferreri (Coleoptera,Leiodidae)

Leiodidae are the second largest subterranean radiation of beetles at family rank. To explore morphological trends linked with troglobiontic habits and characters with potential phylogenetic significance, the head of the cave-dwelling species Troglocharinus ferreri (Cholevinae, Leptodirini) was examined in detail. Overall, the general pattern is similar to what is found in Catops ventricosus (Cholevini). Shared apomorphic features include a fully exposed anterolateral concavity containing the antennal socket, a distinct bead above this depression, a bilobed lip-like structure anterad the labrum, a flat elevated portion of the ventral mandibular surface, and a ventral process at the proximomesal edge of this mandibular area. The tentorial structures are well-developed as in C. ventricosus, with a large laminatentorium and somewhat shortened dorsal arms. The mouthparts are largely unmodified, with the exception of unusually well-developed extrinsic maxillary muscles. Features of T. ferreri obviously linked with subterranean habits are the complete lack of compound eyes, circumocular ridges, and optic lobes. A series of characters is similar to conditions found in other genera of Leptodirini: the head capsule completely lacks a protruding ocular region, a distinct neck is missing, the transverse occipital crest is indistinct, and the antennae are elongate and lack a distinct club. Two different trends of cephalic transformations occur in troglobiontic Leptodirini, with some genera like Troglocharinus and Leptodirus having elongated head capsules and antennae, and others having broadened, more transverse heads. In contrast, the modifications are more uniform in the closely related Ptomaphagini, with a pattern distinctly differing from Leptodirini: the head is transverse, with a distinctly protruding ocular region, a distinct transverse occipital crest, and a very narrow neck region.     

Evolutionary scenarios for unusual attachment devices of Phasmatodea and Mantophasmatodea (Insecta)

Abstract.  The distal parts of the legs of representatives of Phasmatodea and Mantophasmatodea were examined. The condition found in Mantophasma zephyra and Timema nevadense is described in detail. In both species the arolium is highly modified, i.e. strongly enlarged and pan-shaped and densely covered with acanthae. The presence of acanthae on the euplantulae is another very unusual feature shared by the two taxa. A cladistic analysis based mainly on a data matrix from an earlier study of the authors was carried out, with the inclusion of three new characters derived from attachment devices. The results suggest three possible evolutionary scenarios for the features in question. If Phasmatodea are the sister group of Mantophasmatodea, the apomorphic features of the attachment devices may be synapomorphies of both groups, with different degrees of reversal within the suborder Euphasmatodea. A branching pattern Phasmatodea + (Mantophasmatodea + Grylloblattodea) is consistent with the presence of an enlarged pan-shaped arolium and euplantulae with acanthae in the common ancestor of this lineage, with reversal in Grylloblattodea and within Euphasmatodea. The acanthae on the surface of the arolium may or may not have evolved independently in Timema . A placement of Phasmatodea as sister taxon of Orthoptera, Dictyoptera, or a clade comprising both groups implies that the features in question have evolved independently in phasmids and Mantophasmatodea.     

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.     

Morphological and functional heterogeneity in olfactory perception between antennae and maxillary palps in the pumpkin fruit fly,Bactrocera depressa

The morphology and ultrastructure of the olfactory sensilla on the antennae and maxillary palps were investigated through scanning electron microscopy (SEM) and transmission electron microscopy (TEM), and their responses to five volatile compounds were measured using electroantenogram (EAG) and electropalpogram (EPG) techniques in the pumpkin fruit fly, Bactrocera depressa (Shiraki; Diptera: Tephritidae). Male and female B. depressa displayed distinct morphological types of olfactory sensilla in the antennae and maxillary palps, with predominant populations of trichoid, basiconic, and coeloconic sensilla. Basiconic sensilla, the most abundant type of olfactory sensilla in the antennae, could be further classified into two different types. In contrast, the maxillary palps exhibited predominant populations of a single type of curved basiconic sensilla. High‐resolution SEM observation revealed the presence of multiple nanoscale wall‐pores on the cuticular surface of trichoid and basiconic sensilla, indicating that their primary function is olfactory. In contrast, coeloconic sensilla displayed several longitudinal grooves around the sensillum peg. The TEM observation of individual antennal olfactory sensilla indicates that the basiconic sensilla are thin‐walled, while the trichoid sensilla are thick‐walled. The profile of EAG responses of male B. depressa was different from their EPG response profile, indicating that the olfactory function of maxillary palps is different from that of antennae in this species. The structural and functional variation in the olfactory sensilla between antennae and maxillary palps suggests that each plays an independent role in the perception of olfactory signals in B. depressa.     

The pregenital abdominal musculature in phasmids and its implications for the basal phylogeny of Phasmatodea (Insecta: Polyneoptera)

Recently several conflicting hypotheses concerning the basal phylogenetic relationships within the Phasmatodea (stick and leaf insects) have emerged. In previous studies, musculature of the abdomen proved to be quite informative for identifying basal taxa among Phasmatodea and led to conclusions regarding the basal splitting events within the group. However, this character complex was not studied thoroughly for a representative number of species, and usually muscle innervation was omitted. In the present study the musculature and nerve topography of mid-abdominal segments in both sexes of seven phasmid species are described and compared in detail for the first time including all putative basal taxa, e.g. members of Timema, Agathemera, Phylliinae, Aschiphasmatinae and Heteropteryginae. The ground pattern of the muscle and nerve arrangement of mid-abdominal segments, i.e. of those not modified due to association with the thorax or genitalia, is reconstructed. In Timema, the inner ventral longitudinal muscles are present, whereas they are lost in all remaining Phasmatodea (Euphasmatodea). The ventral longitudinal muscles in the abdomen of Agathemera, which span the whole length of each segment, do not represent the plesiomorphic condition as previously assumed, but might be a result of secondary elongation of the external ventral longitudinal muscles. Sexual dimorphism, common within the Phasmatodea, also applies to the muscle arrangement in the abdomen of some species. Only in the females of Haaniella dehaanii (Heteropteryginae) and Phyllium celebicum (Phylliinae) the ventral external longitudinal muscles are elongated and span the length of the whole segment, possibly as a result of convergent evolution.     

The tentorium and anterior head sulci in Dictyoptera and Mantophasmatodea (Insecta)

The tentorium, the anterior sulci of the head capsule (epistomal, subgenal, subantennal, circumantennal, and circumocular sulci), and the extension of the anterior tentorial pit were studied in 26 species of Blattaria (representing most principal lineages), 4 species of Mantodea (including the basal Mantoida schraderi), and 1 species each of Isoptera (the basal Mastotermes darwiniensis) and Mantophasmatodea (Austrophasma caledonense). The morphology of these head structures is compared with literature data on other insect orders, mainly Phasmatodea, Orthoptera, Dermaptera, Embioptera, and Plecoptera, and partly Odonata and Zygentoma. Characters are defined, presented in a matrix, and evaluated with regard to phylogenetic implications and homoplastic evolution. The structural relationships of the subantennal sulcus to the subgenal, circumocular, and circumantennal sulci, which are highly variable and strongly homoplastic (depending much on the size of the compound eyes) are a focal issue; several types of subantennal sulci are defined. The presence of an anterior transverse bridge in the tentorium (“perforated tentorium”) of all Dictyoptera here studied confirms the monophyly of this group. Mantophasmatodea lacks this element.     

A new genus of moths (Lepidoptera: Gracillarioidea: Douglasiidae) in Myanmar amber

A new genus and species of moths, Tanyglossus orectometopus n. gen. et sp. (Lepidoptera: Gracillarioidea), is described from mid-Cretaceous Myanmar amber. Autapomorphies for the new genus are small size (body length of only 0.9 mm), long proboscis greater than body length, presence of ocelli, 3-segmented porrect labial palps, 5-segmented curved maxillary palps, no eye cap, lanceolate wings with reduced venation and a body and wing covering of several types of scales. Based on the available characters, the fossil is placed in the small, aberrant family Douglasiidae and represents the first fossil member of this family. This study adds to our scant knowledge of the diversity of Mesozoic Glossata.     

The head morphology of Clambidae and its implications for the phylogeny of Scirtoidea (Coleoptera: Polyphaga)

External and internal structures of the head of adults of Clambus are described and illustrated in detail. The results are compared with structural features found in the clambid genus Calyptomerus, in representatives of other scirtoid families, and also in species of other coleopteran suborders, notably Myxophaga. The results tentatively support the monophyly of Scirtoidea and a close relationship between Clambidae and Eucinetidae is suggested by one shared derived feature of the mandible, a long and slender apical tooth with a serrate edge. The monophyly of Clambidae is very strongly supported and Acalyptomerus is probably the sistergroup of a clade Calyptomerus + Clambinae. Potential scirtoid autapomorphies are the loss of the dorsal tentorial arms, a bulging gula, a strongly transverse labrum, and a ridge separating the mediostipes from the lacinia. However, all these features are homoplasious. The monophyly of Clambidae is supported by modifications of the head capsule which is strongly flattened and broadened, by a deep clypeofrontal incision enabling vertical antennal movements, and a series of antennal features. Synapomorphies of Clambinae + Calyptomerus (Clambidae excluding Acalyptomerus) are the conglobate body form with the ventral side of the head capsule in contact with the mesocoxae, and compound eyes integrated in the contour of the head. The completely subdivided eye is an autapomorphy of Clambus. An entire series of features is shared by Clambidae (or Scirtoidea) and Myxophaga. Most of them are apomorphies that apparently evolved independently in both groups. However, the presence of well‐developed maxillary and labial glands is arguably a retained groundplan feature of Coleoptera, with parallel loss in Archostemata, Adephaga and various groups of Polyphaga. J. Morphol. 277:615–633, 2016. © 2016 Wiley Periodicals, Inc.