The thoracic morphology of cave-dwelling and free-living ground beetles from China (Coleoptera,Carabidae, Trechinae)

External and internal thoracic structures of two carabid species (Trechini) were examined and documented with different techniques. The study has a main focus on the eyeless cave-dwelling specialist Sinaphaenops wangorum, but detailed information is also provided for a species occurring in cave entrances. The phylogenetic background of the structural features of the thoracic skeletomuscular system was addressed. The thoracic morphology of the examined species was compared to conditions observed in previously studied carabids and non-related subterranean leiodids (Staphylinoidea) in order to identify cave adaptations. Main thoracic character complexes linked with cavernicolous habits in Trechini are elongation of the pro- and mesothorax and the legs, and a complete and irreversible reduction of the flight apparatus. The lost flight capacity is linked with a far reaching modification of skeletal elements of the metathorax including a strongly shortened and simplified metanotum, a shortened metaventrite, and completely reduced wings and sclerites of the wing base. The elongate prothorax together with the long and slender head and elongated legs distinctly increases the activity range in the subterranean lightless environment, which likely facilitates foraging of the carnivorous beetles. Some of the observed features like wing loss and elongation of the anterior thorax and legs are also found in some cave-dwelling Leiodidae (Leptodirini), whereas some other subterranean members of the staphylinoid family have a compact body and legs of normal length. In contrast to the predaceous Trechini, Leptodirini are scavengers.     

Phylogeny of western Mediterranean Leptodirini, with an emphasis on genital characters (Coleoptera: Leiodidae: Cholevinae)

Abstract The tribe Leptodirini (Leiodidae: Cholevinae) is one of the largest radiations of Coleoptera in the subterranean environment. Although subjected to systematic and evolutionary studies, the phylogeny remains poorly understood. We assessed the phylogeny of the western Mediterranean lineages (Iberian Peninsula, Pyrenees and Sardinia) based on a cladistic analysis of fourteen characters of external morphology and twenty characters of the male and female genitalia, studied in 182 species belonging to thirty‐nine genera. We tested the monophyly of the traditional two main divisions of the group (infraflagellates and supraflagellates), as well as that of some ‘phyletic series’. The final matrix contained fifty‐eight terminal taxa, twenty‐four of which had different character state combinations. The strict consensus of the sixty most parsimonious trees recovered a monophyletic Leptodirini, but not their separation into infraflagellates and supraflagellates. The supraflagellates formed a paraphyletic group with respect to the infraflagellates (corresponding to our sampled ‘Speonomus’ series), with Notidocharis sister to all other included Leptodirini, and Speonomidius sister to Leptodirini excluding Notidocharis. The series ‘Spelaeochlamys’, including the Sardinian genera but excluding Pseudochlamys, was recovered as monophyletic with weak support. The ‘Quaestus’ series formed a polytomy with Pseudochlamys plus the ‘Speonomus’ series (including Bathysciola), which was recovered as monophyletic with strong support. Speonomus, Bathysciola, Quaestus and Troglophyes were para‐ or polyphyletic. Our results suggested the respective monophyletic origin of the Leptodirini from the Pyrenees (Pseudochlamys plus the ‘Speonomus’ series) and the Mediterranean coast plus Sardinia (series ‘Spelaeochlamys’). On the contrary, the Leptodirini of the Atlantic north coast of the Iberian Peninsula (series ‘Quaestus’ and ‘Speonomidius’) were not monophyletic.     

The first fossil streblid bat fly, <Emphasis Type="Italic">Enischnomyia stegosoma</Emphasis> n. g., n. sp. (Diptera: Hippoboscoidea: Streblidae)

The first fossil streblid, Enischnomyia stegosoma n. g., n. sp. (Diptera: Hippoboscoidea: Streblidae), is described from Dominican amber. Placed in the subfamily Nycterophiliinae Wenzel, 1966, which includes two New World extant genera, Nycterophilia Ferris, 1916 and Phalconomus Wenzel, 1984 (=Phalcophila Wenzel, 1976), the male specimen of E. stegosoma is characterised by the following features: a laterally compressed body, well-developed two-segmented antennae with the scape fused with the head, a tubular pedicel with an annulated basal portion and swollen apical portion bearing setae and bristles, a distinct flagellum with a dorsal boss bearing microsetae and a subterminal pectinate arista, a large tubular labium (proboscis) with the tip held upwards, eyes reduced to three facets, an expanded and flattened profemur, an anteriorly curved protarsus, and a well-developed wing with an entire distal margin. The possession of wings separates E. stegosoma from the species of Phalconomus, and the wing outline and venation, as well as the structure of the antennae and palps, distinguish it from species of Nycterophilia.     

Sinocyclocheilus donglanensis, a new cavefish (Teleostei: Cypriniformes) from Guangxi, China

Sinocyclocheilus donglanensis, a new cyprinid species from a subterranean river in Donglan County in the Guangxi Zhuang Autonomous Region of southern China, is described. It is distinguished from all congeners by the following combination of characteristics: a completely scaled body with well-developed eyes; a curved lateral line possessing 57–64 scales; pectoral fin not reaching pelvic fin origin and last unbranched ray of the dorsal fin clearly serrated along its posterior edge; 8–9 predorsal vertebrae; 8–9 gill rakers; joints of dentary-angulars not close to each other at the isthmus; and a slightly inferior mouth with the upper jaw (6.2–7.4% in standard length: SL) protruding slightly beyond the lower one (5.7–6.7% SL). Sinocyclocheilus donglanensis is sympatric with the peculiarly shaped, hunchbacked S. altishoulderus.     

Why be cryptic? Choice of host urchin is not based on camouflage in the caridean shrimp Gnathophylloides mineri

The caridean shrimp Gnathophylloides mineri lives among the spines of various species of sea urchins. The shrimp has cryptic coloration that closely matches the coloration of the Caribbean urchin Tripneustes ventricosus. Despite this camouflage, in Jamaica, G. mineri is three times more abundant on the urchin Lytechinus variegatus than on T. ventricosus. L. variegatus exhibits more pronounced covering behavior than T. ventricosus. In laboratory choice experiments, the shrimp exhibited no preference between live, uncovered T. ventricosus or L. variegatus. Similarly, there was no preference between models (the tests, with spines attached, of hollowed out urchins) of T. ventricosus or L. variegatus without cover, or between models of T. ventricosus or L. variegatus with switched cover (i.e., each model was covered with the debris that had belonged to its partner when collected). There was, however, a significant preference for L. variegatus models with their natural covering over T. ventricosus with their natural covering. The shrimp may prefer L. variegatus with its extensive covering over T. ventricosus with extensive covering because the spines of the former are typically erect, as opposed to the spines of T. ventricosus, which usually lay flat across the test and would not give the shrimp much room to maneuver under the debris layer. Our study indicates that despite effective morphological and behavioral traits that render an organism highly cryptic, other, more effective, means of concealment may undermine the usefulness of this crypsis.     

Taxonomic review of the <Emphasis Type="Italic">Sebastes inermis</Emphasis> species complex (Scorpaeniformes: Scorpaenidae)

A taxonomic review of three color morphotypes of the Sebastes inermis species complex established the existence of three valid species, viz. S. inermis, S. ventricosus, and S. cheni. The complex is defined by having two sharp lachrymal spines, the head weakly armed with nasal, preocular, supraocular and parietal spines, and the caudal fin not distinctly emarginated. Sebastes inermis, known from southern Hokkaido southward to Kyushu, Japan, and the southern part of the Korean Peninsula, is characterized as follows: body dark red or light brown dorsally and laterally when fresh, pectoral fin extending beyond level of anus when depressed, pectoral-fin rays 15, anal-fin rays 7, pored lateral line scales 36–44 and gill rakers 31–37. Sebastes ventricosus, known from Iwate and Ishikawa Prefecture southward to Kyushu, Japan, and the southern part of the Korean Peninsula, is characterized as follows: body somewhat greenish-black dorsally and dark silver ventrally when fresh, pectoral-fin rays 16, anal-fin rays 7–8, pored lateral line scales 43–49, and gill rakers 35–39. Sebastes cheni, known from Iwate and Akita Pref. southward to Kyushu, Japan, and the southern part of the Korean Peninsula, is characterized as follows: body dark golden-brown dorsally and laterally when fresh, pelvic fin extending beyond anus when depressed, pectoral-fin rays 17, anal-fin rays 8, pored lateral line scales 37–46 and gill rakers 32–37. In the genetic analysis, the presence or absence of two AFLP fragments was completely fixed among the three species. Sebastes tokionis and S. guentheri are regarded as junior synonyms of S. inermis and S. ventricosus, respectively. A lectotype is designated for S. cheni, and a key to the three species of the S. inermis species complex provided.     

A new Polygordius (Annelida: Polychaeta) from Terra Nova Bay, Ross Sea, Antarctica

Polygordius antarcticus sp. nov. is described from benthic material collected at depths ranging between 31 and 61 m in Terra Nova Bay, Ross Sea, during the 1987–1988 and 1988–1989 expeditions of the Italian National Research Program in Antarctica. This is the first named species of the genus from Antarctic waters. Although no fully mature specimens were available, the new species appears unique in combining the following features: a regular pygidium, inflated and with a single ring of round adhesive pads, a conical prostomium with short antennae (only half as long as the prostomium) and shallow head fold, and a well-developed circulatory apparatus, with circumoesophageal commissures entering the prostomium and long intersegmental commissures insinuating into the ventrolateral compartments of the trunk. These and other significant features were investigated by light and scanning electron microscopy. In the construction of the head and the scheme of the blood system, the new species resembles Polygordius triestinus Woltereck, described by Hempelmann in 1906, an aberrant species inhabiting muddy, anoxic sediments in the Adriatic Sea, but it differs significantly from it in the shape of the mouth (in P. triestinus the upper lip is hypertrophied and strongly protruding) and the pygidium (in P. triestinus this region is stump-like and non-adhesive). The pygidium of P. antarcticus sp. nov. is encircled by 28–30 small adhesive pads, each with 15–20 glandular openings. Preterminal cirri are lacking, as are perianal appendages. The anus is surrounded by six to seven lobes of which the midventral is largest and longest. A world distribution map of the genus is provided. Accepted: 27 August 1998     

Gross and fine structure of the antennal circulatory organ in cockroaches (Blattodea,Insecta)

The antennal circulatory organ of Periplaneta americana and Blaberus craniifer was investigated by light and electron microscopy. This organ consists of two pulsatile ampullae located near the antennal base which are interconnected by a large transverse muscle and associated blood vessels which run into the antennae. Diastole is caused simultaneously in both ampullae by the transverse muscle. Systole is produced passively by the elasticity of the wall of the ampullae and minute accessory tendons. Both elastic structures contain fine unbanded extracellular filaments. The antennal vessels possess two distinct regions: a proximal convoluted region lying within the hemocoel of the head and a narrower distal region running through the antenna and opening near the antennal apex. The length of the proximal portion increases markedly during ontogeny in correlation with the growing antenna. Its wall consists of a high-prismatic epithelium ensheathed by a thick layer of collagen fibrils. The structure of the wall cells is comparable to that found in some salt transporting epithelia: it shows a polar organization with basal infoldings, a large number of mitochondria, and typical arrangement of the junctions or mitochondrial-scalariform junctional complexes. The possible physiological function of this epithelium in ionic or osmoregulation of the hemolymph entering the antenna is discussed. The wall of the distal vessel region consists of a flat single-layered epithelium and seems to be specialized only for delivery of hemolymph to antennae. The structure and function of the antennal heart in cockroaches is compared to that found in other insects.     

Ultrastructure and fate of the nephridial anlagen in the antennal segment of Epiperipatus biolleyi (Onychophora, Peripatidae)—evidence for the onychophoran antennae being modified legs

In this study, new ultrastructural data on the nephridiogenesis in Epiperipatus biolleyi (Onychophora) are provided, and the general distribution of nephridial organs, their vestigia, and derivatives within the onychophoran body is revised. Transient anlagen of nephridial organs proved to be present in the anteriormost segment bearing the antennae. These nephridial anlagen were never found to open to the exterior in any developmental stage studied, but are nevertheless equipped with well-developed cilia. The ciliated nephridial canals are situated at the antennal bases, hence in a more dorsal position than in the remaining body segments. In postantennal segments, the nephridial anlagen constantly arise at the bases of the presumptive legs or their derivatives. These results provide the first evidence that onychophoran antennae are modified legs that retained the original arrangement of the nephridial anlagen at their bases, despite the evolutionary change in position and function of these legs. Current assumptions are accordingly confirmed that the onychophoran antennae and the first antennae in Mandibulata (Euarthropoda) are non-homologous, since they have to be attributed to different head segments. Although the fate of the homologue of the onychophoran antennae in the Euarthropoda remains to be clarified, insights obtained in the present study question previous claims that an ocular segment is absent in extant Mandibulata.     

Ancient origin of a Western Mediterranean radiation of subterranean beetles

Background  

Cave organisms have been used as models for evolution and biogeography, as their reduced above-ground dispersal produces phylogenetic patterns of area distribution that largely match the geological history of mountain ranges and cave habitats. Most current hypotheses assume that subterranean lineages arose recently from surface dwelling, dispersive close relatives, but for terrestrial organisms there is scant phylogenetic evidence to support this view. We study here with molecular methods the evolutionary history of a highly diverse assemblage of subterranean beetles in the tribe Leptodirini (Coleoptera, Leiodidae, Cholevinae) in the mountain systems of the Western Mediterranean.     

The cephalic morphology of free-living and cave-dwelling species of trechine ground beetles from China (Coleoptera,Carabidae)

Morphological adaptations of cave-dwelling organisms including different groups of Coleoptera have fascinated researchers since a long time. Nevertheless, very few detailed documentations of the anatomy of cave beetles using modern techniques are available. In this study, we describe and illustrate external and internal cephalic features of free-living and cave-dwelling trechine carabid beetles using digital microscopy, SEM, micro-CT, and computer-based 3D reconstruction. Morphological characteristics found in three selected species with different habitat preferences are compared. The following derived features distinguish a troglobite species (Sinaphaenops wangorum Ueno et Ran 1998) from a species living in the entrance of caves (Trechiotes perroti Jeannel 1954) and from a fully epigean species (Bembidion sp.): (1) the optic lobes are completely missing; (2) the head capsule, mouth parts, and antennae are distinctly elongated; (3) some muscle attachment areas are shifted; and (4) an additional dilator of the pharynx is present. Despite of conspicuous differences likely related with subterranean habits, the link of some cephalic features to this specific habitat preference remains uncertain.     

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.     

Addition to the revision of the Pristaulacus comptipennis species-group: Description of two new species from Laos and Thailand (Hymenoptera: Aulacidae)

Two species belonging to the recently revised Pristaulacus comptipennis species-group, P. manuelae Turrisi and Madl, sp. n. from Laos and P. iosephi Turrisi and Madl, sp. n. from Thailand, are described, illustrated and compared with most related species. The former species most resembles P. watanabei Turrisi and Smith, 2011, from Thailand, but is readily recognizable, mainly by the shape of the head, which is elongated, with a well-developed temple. The latter species resembles two other species, P. nobilei Turrisi and Smith, 2011, from south China, and P. emarginaticeps Turner, 1922, from Vietnam, but it is easily recognizable mainly by the shape of the medial occipital groove, which is shallow and V-shaped, as well as the shape of the mesosoma, with anterior lobe of mesoscutum, anterior to notauli, strongly elevated dorsally and anteriorly and distinctly overhanging pronotum.     

Mandibular glands of male European beewolves, Philanthus triangulum (Hymenoptera, Crabronidae)

Males of a solitary digger wasp, the European beewolf, Philanthus triangulum, possess large mandibular glands that have been reported to produce a scent marking pheromone. We analysed the morphology and ultrastructure of these glands using light microscopy as well as scanning and transmission electron microscopy. The paired glands are located laterally in the head and each side consists of a larger and a smaller part. Both parts possess a collecting duct each with distinct openings at the mandible base. However, the collecting duct of the larger part is additionally connected to the pharynx through a lateral extension. The collecting ducts are bordered by a monolayered epithelium lined with cuticle that exhibits conspicuous ramified protuberances. About 1400 acini consisting of class 3 gland cells surround the ducts and are connected to them through conducting canals. The main components in the cytoplasm of these gland cells are mitochondria, well-developed smooth endoplasmatic reticulum, and electron lucent vesicles suggesting a high secretory activity. The connection between the large gland parts and the pharynx suggests that the secretion of the mandibular glands might not only be delivered directly onto the mandibles but might also be transported to and stored in the postpharyngeal gland.     

A new species of Platydracus Thomson, 1858 (Coleoptera,Staphylinidae, Staphylininae) from the upper Eocene Florissant beds,Colorado, USA

Platydracus breviantennatus n. sp., is described and figured based on an impression fossil from the upper Eocene Florissant beds of Colorado, the United States. Based on the large and densely setose body, relatively small eyes, and robust mandibles (right mandible seemingly with one simple preapical tooth), tibiae and antennae, the new species is placed in the modern genus Platydracus Thomson, 1858. It differs from other species of Platydracus by its large body, small head, and distinctly short antennae.     

Habitat-dependent growth in a Caribbean sea urchin <Emphasis Type="Italic">Tripneustes ventricosus</Emphasis>: the importance of food type

The sea urchin Tripneustes ventricosus is a common, yet relatively poorly known, grazer of seagrass beds and coral reefs throughout the Caribbean. We compared the size and abundance of urchins between adjacent seagrass and coral reef habitats (where macroalgae are the dominant primary producers). We also conducted a laboratory experiment comparing the growth rate of juvenile urchins fed a diet of either macroalgae or seagrass. Reef urchins had significantly larger test diameter than those in the seagrass on some sampling dates. This size difference may be at least partially explained by diet, because laboratory-reared urchins fed macroalgae grew significantly faster than those fed seagrass. The seagrass population, however, was stable over time, whereas the reef population exhibited strong fluctuations in abundance. Overall, our study indicates that both the seagrass and coral reef habitats are capable of supporting healthy, reproductive populations of T. ventricosus. Each, however, appears to offer a distinct advantage: faster growth on the reef and greater population stability in the seagrass.     

Requirement for Twist1 in frontonasal and skull vault development in the mouse embryo

Using a Cre-mediated conditional deletion approach, we have dissected the function of Twist1 in the morphogenesis of the craniofacial skeleton. Loss of Twist1 in neural crest cells and their derivatives impairs skeletogenic differentiation and leads to the loss of bones of the snout, upper face and skull vault. While no anatomically recognizable maxilla is formed, a malformed mandible is present. Since Twist1 is expressed in the tissues of the maxillary eminence and the mandibular arch, this finding suggests that the requirement for Twist1 is not the same in all neural crest derivatives. The effect of the loss of Twist1 function is not restricted to neural crest-derived bones, since the predominantly mesoderm-derived parietal and interparietal bones are also affected, presumably as a consequence of lost interactions with neural crest-derived tissues. In contrast, the formation of other mesodermal skeletal derivatives such as the occipital bones and most of the chondrocranium are not affected by the loss of Twist1 in the neural crest cells.     

Comparative brain architecture of the European shore crab <Emphasis Type="Italic">Carcinus maenas</Emphasis> (Brachyura) and the common hermit crab <Emphasis Type="Italic">Pagurus bernhardus</Emphasis> (Anomura) with notes on other marine hermit crabs

The European shore crab Carcinus maenas and the common hermit crab Pagurus bernhardus are members of the sister taxa Brachyura and Anomura (together forming the taxon Meiura) respectively. Both species share similar coastal marine habitats and thus are confronted with similar environmental conditions. This study sets out to explore variations of general brain architecture of species that live in seemingly similar habitats but belong to different major malacostracan taxa and to understand possible differences of sensory systems and related brain compartments. We examined the brains of Carcinus maenas, Pagurus bernhardus, and three other hermit crab species with immunohistochemistry against tyrosinated tubulin, f-actin, synaptic proteins, RF-amides and allatostatin. Our comparison showed that their optic neuropils within the eyestalks display strong resemblance in gross morphology as well as in detailed organization, suggesting a rather similar potential of processing visual input. Besides the well-developed visual system, the olfactory neuropils are distinct components in the brain of both C. maenas and P. bernhardus as well as the other hermit crabs, suggesting that close integration of olfactory and visual information may be useful in turbid marine environments with low visibility, as is typical for many habitats such as, e.g., the Baltic and the North Sea. Comparing the shape of the olfactory glomeruli in the anomurans showed some variations, ranging from a wedge shape to an elongate morphology. Furthermore, the tritocerebrum and the organization of the second antennae associated with the tritocerebrum seem to differ markedly in C. maenas and P. bernhardus, indicating better mechanosensory abilities in the latter close to those of other Decapoda with long second antennae, such as Astacida, Homarida, or Achelata. This aspect may also represent an adaptation to the “hermit lifestyle” in which competition for shells is a major aspect of their life history. The shore crab C. maenas, on the other hand seems to rely much less on mechanosensory information mediated by the second antennae but in water, the visual and the olfactory senses seem to be the most important modalities.     

Head patterning and Hox gene expression in an onychophoran and its implications for the arthropod head problem

The arthropod head problem has puzzled zoologists for more than a century. The head of adult arthropods is a complex structure resulting from the modification, fusion and migration of an uncertain number of segments. In contrast, onychophorans, which are the probable sister group to the arthropods, have a rather simple head comprising three segments that are well defined during development, and give rise to the adult head with three pairs of appendages specialised for sensory and food capture/manipulative purposes. Based on the expression pattern of the anterior Hox genes labial, proboscipedia, Hox3 and Deformed, we show that the third of these onychophoran segments, bearing the slime papillae, can be correlated to the tritocerebrum, the most anterior Hox-expressing arthropod segment. This implies that both the onychophoran antennae and jaws are derived from a more anterior, Hox-free region corresponding to the proto and deutocerebrum of arthropods. Our data provide molecular support for the proposal that the onychophoran head possesses a well-developed appendage that corresponds to the anterior, apparently appendage-less region of the arthropod head.     

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.