The larval abdomen of the enigmatic Nannochoristidae (Mecoptera,Insecta)

External and internal structures of the larval abdomen of Nannochorista are described in detail, with emphasis on the posterior segments. The results are compared with conditions found in other groups of Antliophora, especially the mecopteran subgroups Boreidae and Pistillifera. Like the entire postcephalic body, the larval abdomen of Nannochorista is extremely slender and nearly cylindrical. The anterior segments are largely unmodified. The surface is smooth and lacks any protuberances or prolegs. The term “cloaca” for the posterior membranous pouch of Nannochorista sp. is morphologically unjustified. A list of muscles of segments IX and X is presented. The abdominal musculature was partly homologized following Snodgrass. The muscles of segment X are highly modified. They move the membranous pouch, the anal papillae, and the terminal lobes. The presence of these structures is likely an adaptation to the specific aquatic life style of nannochoristid larvae. The anal papillae are possibly homologous to the 4-lobed terminal attachment apparatus of larvae of Caurinus (Boreidae) and Pistillifera (Panorpidae, Bittacidae, Choristidae) but this is uncertain. The specific condition in both groups, i.e. two retractile papillae with tracheae and Malpighian tubules in Nannochoristidae, and a 4-lobed exposed attachment device in Pistillifera + Boreidae (groundplan) are very likely autapomorphic for both groups, respectively. A slender abdomen with smooth surface is very likely plesiomorphic within Antliophora and Mecopterida. This condition is found in Trichoptera (partim), Nannochoristidae, Siphonaptera, and many basal groups of Diptera. An eruciform or scarabaeiform body shape with a soft, largely unsclerotised cuticle is probably a synapomorphy of Boreidae and Pistillifera. The presence of ventral protuberances resembling prolegs on the anterior segments is an autapomorphy of the latter group. The homology of paired or unpaired terminal appendages of segment X is uncertain. However, the specific condition of paired and 3-segmented appendages with hooks in Nannochoristidae is almost certainly autapomorphic for this family. The protracted opening of the hind gut on the membranous pouch is another potential autapomorphy of Nannochoristidae. Aquatic habits of larvae, also very likely an apomorphic condition, have likely evolved several times independently in Antliophora.     

The abdomen of a free-living female of Strepsiptera and the evolution of the birth organs

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

Morphology of the terminalia of the stick insect Dajaca napolovi from Vietnam (Insecta: Phasmatodea)

Aschiphasmatinae is a small group of stick insects from the Oriental region whose genital morphology has been rarely described in detail. The subfamily is of particular interest, as phylogenetic studies have shown Aschiphasmatinae to be the sister group to the remaining Euphasmatodea. In this paper, the male and female terminalia are described for the first time in Dajaca napolovi Brock, a little known aschiphasmatine species from Vietnam. In the male, the transversally undivided abdominal sternum IX and gently incurved cerci with a conspicuous apical tooth represent apomorphies of Aschiphasmatinae. Thorn pads on the hind margin of abdominal tergum X consist of only a single row of 6–7 ventrally oriented teeth. The simple thorn pad structure of Aschiphasmatinae can represent an ancestral condition for Euphasmatodea. The vomer on venter X is well‐developed and features two unusually large basal apodemes and a strongly developed apical spine showing a specialized streaked surface micropattern. Female terminalia are characterized by an unkeeled abdominal sternum VIII covering the reduced primary ovipositor. Gonapophysis VIII does not extend beyond the tip of gonapophysis IX. An asymmetry concerning the size of the paired gonapophyses is reported for the first time in Phasmatodea.     

Observations on the biology of Hemimems talpoides (Insecta: Dermaptera)

The little known ectoparasitic insects of the genus Hemimerus lived and bred on a captive giant rat Cricetomys , allowing prolonged observation of their habits. They never normally left the host for more than a few seconds and never fed elsewhere than on the host. Food was taken from the skin, ears, eyes and mouth but the rat never showed serious irritation. Breeding was slow and possibly closely controlled by temperature. The position in copulation was quite different from that of typical Dermaptera.     

External female genitalia of six species of the genus Meccus (Hemiptera: Reduviidae: Triatominae)

Triatomine classification is based on morphological characteristics. Studies have been conducted to improve their identification by observing many characteristics. However, there are problems of differentiating among highly interrelated species and new criteria are required. The purpose of this study was to determine the morphological differences in the external female genitalia of M. pallidipennis, M. longipennis, M. picturatus, M. bassolsae, M. mazzottii, and M. phyllosomus in order to distinguish among species using scanning electron microscopy. Observations were made of the dorsal, posterior, lateral, and ventral views of the female external genitalia for each species. In the six species we studied, relevant differences were observed in the dorsal view of the X segment, as well as the IX, VIII, and VII tergites. In the posterior and lateral view, the most visible differences were registered in the gonocoxite size of the segments VIII, IX, and X. Finally, in the ventral view of the VII sternite, differences among species were observed in the size of the inflection in the top and upper corner. Our results show that it was possible to differentiate among the triatominae species for each of the four views by using a scanning electron microscope to analyze morphological characteristics of the VII, VIII, IX, and X abdomen segments.     

Phylogeny of earwigs (Insecta: Dermaptera) based on molecular and morphological evidence: reconsidering the classification of Dermaptera

Abstract. Dermaptera (earwigs) is a cosmopolitan order of insects, the phylogenetic relationships of which are poorly understood. The phylogeny of Dermaptera was inferred from large subunit ribosomal (28S), small subunit ribosomal (18S), histone-3 (H3) nuclear DNA sequences, and forty-three morphological characters. Sequence data were collected for thirty-two earwig exemplar taxa representing eight families in two suborders: Hemimeridae (suborder Hemimerina); Pygidicranidae, Anisolabididae, Labiduridae, Apachyidae, Spongiphoridae, Chelisochidae and Forficulidae (suborder Forficulina). Eighteen taxa from ten additional orders were also included, representing Ephemeroptera, Odonata, Orthoptera, Phasmida, Embiidina, Mantodea, Isoptera, Blattaria, Grylloblattodea and Zoraptera. These data were analysed via direct optimization in poy under a range of gap and substitution values to test the sensitivity of the data to variations in parameter values. These results indicate that the epizoic Hemimerus is not sister to the remaining Dermaptera, but rather nested as sister to Forficulidae + Chelisochidae. These analyses support the paraphyly of Pygidicranidae and Spongiphoridae and the monophyly of Chelisochidae, Forficulidae, Anisolabididae and Labiduridae.     

春尺蠖蛹雌雄外部形态特征快速鉴别分析

为依据春尺蠖Apocheima cinerarius Erschoff蛹的形态特征快速、无损、准确鉴别雌雄个体,对其头部、胸部、腹部和体色外部鉴别特征进行了分析,并通过解剖生殖系统验证准确性.结果表明:以春尺蠖蛹的胸部和腹部特征识别雌雄准确率明显高于头部和体色特征,识别率可达100％.首先,雌蛹第8腹节腹板前缘中部具有"Y"型沟,与第7腹节腹板后缘形成倒三角状,而雄蛹无此特征.其次,雌蛹生殖孔与产卵孔连接形成裂缝,两侧平坦无突起,而雄蛹第9腹节腹板中央有一纵裂缝的生殖孔,两侧各有半圆状瘤状突起.最后,雌蛹的胸部背板各节间相对长度均小于雄蛹,而雄蛹中胸背板最宽,其后缘明显向外凸起.因此,春尺蠖蛹胸部或腹部特征可用于快速、准确地鉴别雌雄性别.     

When the Body Hides the Ancestry: Phylogeny of Morphologically Modified Epizoic Earwigs Based on Molecular Evidence

Here, we present a study regarding the phylogenetic positions of two enigmatic earwig lineages whose unique phenotypic traits evolved in connection with ectoparasitic relationships with mammals. Extant earwigs (Dermaptera) have traditionally been divided into three suborders: the Hemimerina, Arixeniina, and Forficulina. While the Forficulina are typical, well-known, free-living earwigs, the Hemimerina and Arixeniina are unusual epizoic groups living on molossid bats (Arixeniina) or murid rodents (Hemimerina). The monophyly of both epizoic lineages is well established, but their relationship to the remainder of the Dermaptera is controversial because of their extremely modified morphology with paedomorphic features. We present phylogenetic analyses that include molecular data (18S and 28S ribosomal DNA and histone-3) for both Arixeniina and Hemimerina for the first time. This data set enabled us to apply a rigorous cladistics approach and to test competing hypotheses that were previously scattered in the literature. Our results demonstrate that Arixeniidae and Hemimeridae belong in the dermapteran suborder Neodermaptera, infraorder Epidermaptera, and superfamily Forficuloidea. The results support the sister group relationships of Arixeniidae+Chelisochidae and Hemimeridae+Forficulidae. This study demonstrates the potential for rapid and substantial macroevolutionary changes at the morphological level as related to adaptive evolution, in this case linked to the utilization of a novel trophic niche based on an epizoic life strategy. Our results also indicate that the evolutionary consequences of the transition to an ectoparazitic mode of living, which is extremely rare in earwigs, have biased previous morphology-based hypotheses regarding the phylogeny of this insect group.     

An unusual new species of the genus Brachypalpus Macquart (Diptera: Syrphidae) from Eastern Asia

Brachypalpus (Brachypalpus) longifacies sp. nov. is described and illustrated from Eastern Asia (Russian Far East and Japan). It differs from other species of the nominative subgenus by several unusual characters, i.e. face in profile weakly convex dorsally and concave ventrally, antennae very elongated, and abdomen with reddish transversal maculae on 2nd tergum (male) or 2nd and 3rd terga (female). A key is given for known species of the subgenus Brachypalpus.LSID: urn:lsid:zoobank.org:pub:634CB6E2-B2C1-40EC-8811-BF3DD4BDE69A     

Revision of the Neotropical genus Chimarrhodella Lestage (Trichoptera: Philopotamidae)

Abstract. A revision and phylogenetic analysis of the Neotropical genus Chimarrhodella Lestage is presented. Four described species are recognized, C.galeata (Martynov), C.peruviana (Ross), C.ulmeri (Ross) and C.nigra Flint. In addition, five new species are described, C.costaricensis sp.n. (Costa Rica), C.tapanti sp.n. (Costa Rica), C.flinti sp.n. (Venezuela), C.pilcopata , sp.n. (Peru) and C.tobagoensis sp.n. (Tobago). These species are organized into two monophyletic species groups, designated here the Galeata Group and the Peruviana Group. The Galeata Group is characterized by having male genitalia with tergum X possessing well developed lateral lobes bearing sclerotized, spine-like processes and includes C.galeata, C.costaricensis, C.flinti, C.pilcopata and C.tapanti. The Peruviana Group is characterized by having male genitalia with tergum X only slightly divided apically, tergum IX bearing hook-like processes posteriorly, and female genitalia with segment VIII much elongated in comparison to the Galeata Group and includes C.peruviana, C.nigra, C.tobagoensis and C.ulmeri. Records of the genus from Costa Rica and Tobago represent significant extensions of the known range for the genus, previously unknown north of Panama. A key to males and known females and a species-level phylogenetic analysis are included.     

Characterization of muscles associated with the articular membrane in the dorsal surface of the crayfish abdomen

The anatomy, physiology, and biochemistry of the dorsal membrane muscle (DMA) and the superficial extensor muscle accessory head (SEAcc) in the abdomen of the crayfish, Procambarus clarkii and lobster, Homarus americanus, are reported. These muscles have not been previously characterized physiologically or biochemically. The anatomy was originally described by Pilgrim and Wiersma (1963. J Morph 113:453-587). The arrangement of these muscles varies depending on the abdominal segment. The function of the dorsal membrane muscle is to retract the thin articulating membrane joining the cuticular segments so that the dorsal membrane does not evert during extension of the abdomen. Consequently, the articular membrane does not protrude, and thus potential damage to the membrane is minimized. Examination of nerve terminal morphology revealed strings of varicosities, usually only associated with tonic terminals. The electrophysiological data indicate that there are at least four tonic excitatory and one inhibitory motor neuron innervating these muscles. Facilitation indices and fatigue-resistance indicate physiologically the tonic nature of innervation. Anti-GABA antibodies demonstrate the anatomical presence of an inhibitor motor neuron. The SDS electrophoretic analysis of myofibrillar proteins and Western blots of key protein isoforms for these muscles in crayfish and lobsters also indicate that the DMA and SEAcc muscles are tonic phenotype. J. Exp. Zool. 287:353-377, 2000.     

Evolution of locomotory attachment pads in the Dermaptera (Insecta)

This contribution is the first comparative SEM study of tarsal and pretarsal structures of 18 dermapteran species, including epizoic Hemimeridae, rare Apachyidae, as well as basal Pygidicranidae. Our data reject the apparent uniformity of this taxon and show that representatives of Dermaptera have independently evolved both types of attachment mechanisms: hairy and smooth. Dermaptera possess a wide spectrum of attachment devices: arolia, euplantulae, tarsal surfaces covered with specialised tenent setae and other types of cuticular outgrowths. The groundpattern of the pretarsal and tarsal attachment structures was reconstructed by mapping their characters onto a cladogram, generated without tarsal characters. In the groundpattern of recent Dermaptera, the tarsus consists of three tarsomeres. Presumably, the last common ancestor of the Dermaptera possessed an arolium, since this structure occurs in the most basal taxa: Diplatyidae, Karschiellidae (partim, adults), Pygidicranidae partim, and Apachyidae. The absence of arolium in two of the pygidicranid taxa is probably due to a secondary loss. The arolium seems to be reduced in the 'higher Dermaptera' and amongst them, only the Geracinae, which belong to the Spongiphoridae and, hence, to the well supported Eudermaptera [European Journal of Entomology, 98 (2001), 445], evolved this structure convergently. The character state distribution for euplantulae suggests their evolution being similar to that of the arolium. All species of Tagalina possess a specialised tarsus with a strongly dilated second tarsomere. The same applies to the Forficulidae. However, their relatively remote phylogenetic position to Tagalina burri is a convincing reason to assume convergent evolution of this character. The Chelisochidae, with a slender, elongated second tarsomere, possess a unique structure, which supports their monophyly. The special, heart shaped structure of the second tarsal segments in the Forficulidae suggests their monophyly. The attachment structures of Hemimerus vosseleri are highly derived and probably autapomorphic for this taxon.     

Two new species of caddisflies from East Asia （Trichoptera： Philopotamidae, Psychomyiidae）

A new species of Kisaura Ross （Philopotamidae） from Japan and a new species of Psychomyia Latreille （Psychomyiidae） from Russia are described. Kisaura imparis n. sp. is closest to Kisaura aurascens （Martynov, 1934）, but the lateral processes of tergum X are asymmetrical and the apex of tergum X is different. Psychomyia poltavkaensis n. sp. resembles Psychomyia n. sp., but the shape of tergum IX＋X is more rounded, the apex of the basoventral process of each preanal appendage is unforked, and the inferior appendages are branched.     

The gonangulum: A reassessment of its morphology,homology, and phylogenetic significance

The gonangulum is a sclerite in the female genitalic region of insects. Its presence or full development has long been considered an apomorphy supporting Zygentoma + Pterygota. Recent studies of female genitalia in several insect orders (K.-D. Klass and co-workers) revealed many new data on the gonangulum and homologous sclerotisations (laterocoxa LC9). Herein the gonangulum area is described (including articulations, muscle attachments, sulci) and compared among Archaeognatha, Zygentoma, Odonata, Dermaptera, Dictyoptera, and Notoptera. A wider perspective is provided to the topic by addressing some novel issues: identification of LC9 sclerotisations in non-insect taxa and in insects that secondarily lack an ovipositor; occurrence of homonomous sclerotisations in other abdominal segments of both sexes; morphological interpretation of LC9; and the role of paedomorphosis in LC9 evolution. As a result, there is currently no support for any insect lineage from this character system. For gonangulum-related characters both a significant intra-ordinal variation and frequent homoplasy are demonstrated using various Odonata, Dermaptera, and Dictyoptera as examples. Divergent fates of LC9 in simplified genitalia are shown using a dermapteran and an odonatan. We view all this as a showcase of how a renewed and more detailed examination of a character system can dramatically change the phylogenetic evidence drawn from it.     

An enigmatic new stick insect from the Philippine Islands (Insecta: Phasmatodea)

A new genus and species of stick insect is described and figured from Mount Halcon, on the Philippine island of Mindoro. Conlephasma enigma gen. et sp. n. is a stout, flightless, and apparently ground-dwelling species with vivid integumental colors. When disturbed, specimens spray a defensive secretion from the prothoracic exocrine glands. The systematic position of Conlephasma within Euphasmatodea is unclear. The elongated galealobulus and the trichome area located laterally in the galea, represent unusual apomorphic characters of the maxilla that could indicate affinities with Necrosciinae or Pseudophasmatinae. All tibiae exhibit the anareolate condition. Euplantulae are of two types: those of tarsomeres I–IV feature a nubby microstructure, whilst the one on the ventral side of the pretarsus is smooth. Males are characterized by the presence of a well-developed vomer on the tenth abdominal segment. A distinctive and apomorphic trait of female terminalia is represented by the elongated tenth abdominal tergum. Conlephasma can represent an interesting taxon for studies on the evolution of the stick and leaf insects.     

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.     

Ecological aspects of the external morphology and functionality of the psychomyiid female ovipositor (Insecta,Trichoptera)

Females of most psychomyiid species bear an elongated ovipositor enabling them to oviposit their eggs into grooves and cavities in different kinds of substrates in freshwaters. Although the preference of psychomyiid species for oviposition into distinct substrates and the subsequent larval life performance in aquatic habitats is already known, the mode of oviposition and the functional morphology of the elongated ovipositor have not been described until now. In this study we present SEM photographs of some psychomyiid species with female ovipositors of different lengths and shapes, explaining their preference for oviposition into distinct substrates. Additionally, we discuss some ecological aspects of the ovipositor shapes and the mode of egg laying for psychomyiids. The female ovipositor consists of two body segments, with an elongated IXth and a distinctly shorter Xth segment, which is flexible and can be pushed up dorsally when releasing the eggs. On the basis of SEM photographs, we suppose that the opening of the channel through which the eggs were released is located ventrally near the ovipositor tip. The ovipositor itself is characterised by a ventral cleft reaching from the gonoporus of segment IX to the tip of the last segment X. We stored adult females abdomina overnight in different aqueous dilutions of ethanol leading to different stages of swelling of the ovipositors (grade of swelling in distilled water >30% ethanol >70% ethanol). Some internal membrane-like structures, normally infolded into the ovipositor, became visible by the swelling of the ovipositor. We discuss the possible rearrangement of the organisation of segment IX resulting in the development of an ovipositor on the basis of SEM photographs of the differently swollen ovipositors.     

A new species of flea in the genus Ctenophthalmus (Siphonaptera: Hystrichopsyllidae)]

A new species, Ctenophthalmus (Euctenophthalmus) parthicus sp. n., is described from social vole (Microtus socialis Pallas) from four localities of the West Kopetdag Mountains of Turkmenia. The new species is close to C.(E.) secundus Wagner and C.(E.) congener nadimi Farhang-Azad. It differs from males of the other known species by the following characters. The movable process has an infra-acetabular process whose length is the same as the height of its supra-acetabular portion. The movable process has the straight front margin, right anterior apical angle. The anterior part of the dorsal margin of the movable process is elevated, the hind part is slightly concave. The posterior apical angle of the movable process is cut. The hind margin of the movable process is convex. The apex of the distal arm of sternum IX is straight, not oblique. The female differs by the following peculiarities. Sternum VII has a well developed dorsal lobe below which there is a smaller rounded lobe and a distinct small ventral process. Unciform sclerotization of tergum VIII (or the place of connection between tergum VIII and its internal lobe) resembles an arc. The front margin of the internal lobe of tergum VIII is not sclerotized. The bursa copulatrix is as long as the dorsal spines of pronotum.     

Morphology of the dorsal vessel in the abdomen of the blood-feeding insect Rhodnius prolixus

The dorsal vessel (DV) in the abdomen of the blood-feeding insect Rhodnius prolixus was divided functionally into two regions, the heart, into which haemolymph entered the DV through four pairs of ostia located in abdominal segment VII, and the aorta, along which the haemolymph was propelled from abdominal segment VI to the thorax. Osmium-fixed whole mounts revealed the DV to consist of spirally arranged striated muscle fibers and to possess two rows of ventrally attached longitudinal fibers extending the length of the abdomen. Seven pairs of alary muscles were found attached to the DV in the posterior abdominal segments. Contractions of the alary muscles attached to the ventral surface of abdominal segments VII and VIII served to expand the heart. Electron microscopy revealed the DV to consist of a thin layer of contractile elements surrounded by an inner (intima) and outer (adventitia) connective tissue layer. Embedded in the intima along each lateral side of the DV were two large groups of endocardial cells extending the length of the DV. A small group of pericardial cells was embedded in the adventitia along the mid-ventral side of the DV, and clusters of pericardial cells were found attached to the alary muscles. Nerve terminals were found only on the heart: they contained agranular synaptic vesicles approximately 30 nm in diameter and densely stained granules approximately 100-120 nm in diameter. These structural components are discussed in relation to the role of the DV in circulation.