The thorax of Mantophasmatodea,the morphology of flightlessness,and the evolution of the neopteran insects

Mantophasmatodea was described as a new insect order in 2002. Since then, this small group of wingless insects has developed into one of the best investigated insect taxa. Nevertheless, many aspects of mantophasmatodean morphology as well as their evolutionary relationships remain ambiguous. To determine the phylogenetic relationships of Mantophasmatodea based on an extended character set and to elucidate possible morphological adaptions towards flightlessness, we investigated the thoracic morphology of two species, Austrophasma caledonensis and Mantophasma sp. The morphological similarity between these two species is striking and no differences in musculature were found. The mantophasmatodean thorax strongly resembles that of ice crawlers (Grylloblattodea), especially with respect to the presence of pleural processes in the meso‐ and metathorax, branched furcae in all segments, and similar muscle equipment. In a cladistic analysis containing all major lineages of Neoptera, the monophyly of Polyneoptera is supported by the presence of an anal fan and several modifications of the wing joint. Within Polyneoptera, a sister‐group relationship between stoneflies and the remaining Polyneoptera is supported. A clade comprising Mantophasmatodea and the Grylloblattodea gains strong support from thoracic morphology and can be considered assured. Potential thoracic apomorphies include prothoracic paracoxal invaginations, pterothoracic pleural arms that originate from the epimeron, and a unique metathoracic sterno‐coxal musculature. The monophyly of Orthoptera and Dictyoptera is further supported while the deeper polyneopteran nodes remain unresolved. Among the wingless taxa investigated we found few general morphological adaptations whereas, in other aspects, especially in the musculature, strong differences could be observed. However, much more research on the strongly neglected topic of flightlessness is required to make reliable statements.     

Embryonic development of Carabus insulicola (Insecta,Coleoptera, Carabidae) with special reference to external morphology and tangible evidence for the subcoxal theory

The egg morphology and successive changes in the developing embryos of the carabid ground beetle Carabus insulicola (Carabidae) are described based on light and scanning electron microscopy observations. Newly laid eggs of this species are ellipsoid and measure approximately 6.1 × 2.9 mm, before increasing to 6.6 × 3.4 mm at hatching. The egg period is about 11 days at 23°C. The egg shell is characterized by a thin fragile chorion covering a hard serosal cuticle. The embryo forms on the ventral egg surface, where it develops for the duration of the egg period. During the process of thoracic leg formation, two subcoxal rings, subcoxae‐1 and 2, are clearly discernible at the basalmost region of the leg rudiments, and these subcoxae participate in the formation of the larval pleura and sterna. The result thus provides tangible evidence for the subcoxal theory, that is, that thoracic pleura and sterna are derived from subcoxal regions. Despite the complete absence of abdominal appendages in the larvae of this species, two pairs of appendage‐like swellings, the medial and lateral ones, temporarily arise in the first eight abdominal segments during the middle of embryonic development. The medial swellings are assumed to be serially homologous with the coxal part of the thoracic leg, and they later flatten out and participate in the formation of the larval pleura (hypopleurites). In the light of the serially homologous relationships among gnathal appendages, thoracic legs, and abdominal appendage‐like swellings, we identified the subcoxal regions in both the gnathal and abdominal segments. Although, the lateral swellings soon degenerate and disappear, it is considered that the swellings originate in the abdominal subcoxae‐2 and may be homologous to the tracheal gills of larvae of Gyrinidae. Based on the embryological results, new interpretations for the constituent of gnathal appendages are proposed. J. Morphol. 274:1323–1352, 2013. © 2013 Wiley Periodicals, Inc.     

Different requirements for l(1) giant in two embryonic domains of Drosophila melanogaster

l(1) giant is a zygotic lethal mutation which affects the embryonic development of both the labial/thoracic segments and a subset of posterior abdominal segments. Using antibodies specific for proteins encoded by several Drosophila genes to identify the compartmental origin of the defects, we show that the requirement of giant activity is different in these two embryonic domains. Anteriorly, the posterior compartment of the labial segment is missing at the blastoderm stage. Posteriorly, cells are specifically deleted by cell death within the anterior compartments of abdominal segments 5-7 during germ band elongation. In mature embryos, posterior compartment structures of the peripheral nervous system of A5-7 are fused. In addition to a different pattern of defect in the two parts of the embryo, the kind of action appears different. Anteriorly, giant resembles a gap mutation in that a particular region is missing from the blastoderm fate map, whereas in the abdominal domain, giant affects the development of anterior compartment-specific structures.     

Embryonic development of a whirligig beetle,Dineutus mellyi,with special reference to external morphology (insecta: Coleoptera,Gyrinidae)

The egg morphology and successive changes of developing embryos of the whirligig beetle, Dineutus mellyi (Adephaga: Gyrinidae) are described from observations based on light and scanning electron microscopy. The egg surface is characterized by minute conical projections covering the entire egg surface, a stalk‐like micropylar projection at the anterior pole of the egg, and a longitudinal split line along which the chorion is cleaved during the middle embryonic stages. The germ band or embryo is formed on the ventral egg surface, and develops on the surface throughout the egg period; thus, the egg is a superficial type, as is the case in most coleopteran species. A pair of lateral tracheal gills (LTGs) of the first abdominal segment originates from appendage‐like projections arising at the lateral side of pleuropodia, and the LTGs of the second to ninth abdominal segments are arranged in a row with that of the first segment. Therefore, LTGs are structures with serial homology. The paired dorsal tracheal gills (DTGs) of the ninth abdominal segment are formed on the regions just latero‐dorsal to the LTGs of this segment. Regarding the pleuropodia as the structures being homologous with thoracic legs, neither the LTGs nor DTGs are homologous with thoracic legs, but originate in the more lateral region corresponding to the future pleura of the thoracic segments. The last (10th) abdominal segment in the larva is formed by the fusion of the embryonic 10th and 11th abdominal segments. Four terminal hooks at the end of the last abdominal segment originate from two pairs of swellings on the posterior end of the embryonic 11th abdominal segment. It is proposed that the terminal hooks possibly correspond to the claws of medially fused cerci of the embryonic 11th abdominal segment. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Different requirements for l(1)giant in two embryonic domains of Drosophila melanogaster

L(1)giant is a zygotic lethal mutation which affects the embryonic development of both the labial/thoracic segments and a subset of posterior abdominal segments. Using antibodies specific for proteins encoded by several Drosophila genes to identify the compartmental origin of the defects, we show that the requirement of giant activity is different in these two embryonic domains. Anteriorly, the posterior compartment of the labial segment is missing at the blastoderm stage. Posteriorly, cells are specifically deleted by cell death within the anterior compartments of abdominal segments 5–7 during germ band elongation. In mature embryos, posterior compartment structures of the peripheral nervous system of A5–7 are fused. In addition to a different pattern of defect in the two parts of the embryo, the kind of action appears different. Anteriorly, giant resembles a gap mutation in that a particular region is missing from the blastoderm fate map, whereas in the abdominal domain, giant affects the development of anterior compartment-specific structures.     

Revaluation of the prothoracic pleuron of the membracidae (Homoptera): The presence of an epimeron and a subdivided episternum in Stictocephala bisonia Kopp and Yonke,Oxyrhachis taranda (Fabr.), and Centrotus cornutus (L.)

The propleuron of adults of Stictocephala bisonia Kopp and Yonke, Oxyrhachis taranda (Fabr.) and Centrotus cornutus (L.) (Homoptera: Membracidae) and 5th-instar nymphs of S. bisonia is investigated using scanning electron microscopy of the external and internal skeletal surface. Criteria for homologizing propleural features in the Pterygota are detailed in an attempt to resolve contradictions of earlier interpretations. In the species examined, the following features occur in adults and in the nymphs of S. bisonia: the pleural sulcus, pleural apophysis, episternum, epimeron, precoxale, and trochantin. In adults, the episternum is separated into an anterior and a lateral part. The prothoracic tooth of O. taranda is an epimeral feature.     

Head structures of Karoophasma sp. (Hexapoda, Mantophasmatodea) with phylogenetic implications

External and internal head structures of adults of Karoophasma sp. were examined and described. The results are compared with conditions found in other representatives of Mantophasmatodea and members of other lower neopteran groups. The X-shaped apodeme of the frons, the unpigmented oval area enclosed by apical branches of the anterior tentorial arms, the oval sclerotisation at the base of the labrum, the sclerotized rounded apical part of the galea, and the loss of M. labroepipharyngalis are probably autapomorphic for Mantophasmatodea. Plesiomorphic features (groundplan of Neoptera) are the orthognathous condition, the absence of parietal ridges, the absence of a gula, the absence of a 'perforation of the corpotentorium', the multisegmented antennae inserted between the compound eyes, the general arrangement of the mouthparts, the shape and composition of the maxillae and labium, and the nearly complete set of muscles. The presence of a transverse muscle connecting the antennal ampullae is a potential synapomorphy of Orthoptera, Phasmatodea and Dictyoptera. Character states suggesting affinities with Grylloblattodea are the absence of ocelli, the elongation of the corpotentorium, and the very similar mandibles with widely separated bases and completely reduced molae. Whether predacious habits are a synapomorphic feature of Mantophasmatodea and Grylloblattodea is uncertain. The retained orthognathous condition in Mantophasmatodea and Mantodea is likely related with different specialized preying techniques in both groups, i.e. rapid forward pushes of the head–prothorax complex, and the use of raptorial legs, respectively.     

东亚飞蝗胚胎体节的形成过程

体节形成是昆虫胚胎发育过程中的关键问题.东亚飞蝗Locusta migratoria manilensis(Meyen)是一种重要的农业害虫,其体节形成的时序过程尚无详细报道.本研究采用免疫组化和品红染色方法研究了室内人工饲养东亚飞蝗的体节形成过程.结果表明:完成受精后,细胞核开始分裂并向卵表面迁移.细胞核到达卵表面的...     

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.     

The sperm structure of <Emphasis Type="Italic">Galloisiana yuasai</Emphasis> (Insecta,Grylloblattodea) and implications for the phylogenetic position of Grylloblattodea

The sperm ultrastructure of the Grylloblattodea Galloisiana yuasai was described and the sperm characters were comparatively examined in several orthopteroid insect orders for inferring the phylogenetic placement of the Grylloblattodea. The spermatozoa of G. yuasai are joined in bundles (spermatodesms) containing 200 units. Major features of these spermatozoa include a monolayered acrosome, a 9+9+2 axoneme with 16-pfs accessory microtubules and expanded intertubular material, and an evident “centriole adjunct”. The diffused material observed between the axoneme and the mitochondrial derivatives is considered to be an extension of the three connecting bands observed in other orthopteroid taxa, similar to what happens in some orthopteran lineages. The presence of the connecting bands, even though modified in G. yuasai, suggests that the Grylloblattodea are to be placed in a clade with Mantophasmatodea, Mantodea and Orthoptera.     

What is the epipleurite? A contribution to the subcoxal theory as applied to the insect abdomen

The epipleurites were originally described by Hopkins in 1909 on the imago and larva of a beetle. Then this term was widely used in insect morphology, mainly for larvae, to designate certain sclerites of the pleural region. They have recently been interpreted as tergopleural (i.e. pleural but not strictly appendicular) by Deuve in 2001, but a study of embryonic development by Kobayashi et al. in 2013 has shown that they are instead eupleural (i.e. appendicular) and correspond to a dorsal part of the subcoxa. Their presence in the abdominal segments of insects illustrates the fundamental importance of the subcoxa in segmental structure, with a function of anchoring and supporting the appendage when the latter is present. However, the epipleurites are normally separated and functionally dissociated from the coxosternum, which integrates the ventral component of the subcoxa. In females, the epipleurite of segment IX of the abdomen corresponds to the gonangulum, as already pointed out by Deuve in 1994 and 2001, and it is involved in gonopod articulation. At segments VIII and IX of both males and females of holometabolans, the formation process of the genital ducts leads to an internalisation of the whole subcoxosternum (i.e. the coxosternum with the exception of the coxal and telopodal territories), and it is the two flanking epipleurites that ventrally close the abdomen in relation to the rearward displacement of the gonopore. This model may be generalised, in its broad lines, to a large part of the hemimetabolans. The body plan of the insect abdomen underlines the morphological and functional importance of the subcoxa in its fundamental structure, but the study of the Hexapoda in general also indicates the presence of a more proximal segment, the precoxa, which would belong to the groundplan but is more cryptic because it is often closely associated with the subcoxa and/or the paranotal lobe. Its location, which is sometimes on the ventral flank of the paranotal lobe, is in line with the hypothesis of a dual origin of the pterygote wing.     

A possible evolutionary pathway to insect flight starting from lepismatid organization

Starting from the hypothesis that flight in Pterygota evolved from lepismatid organization of their ancestors, the functional anatomy of the thorax was studied in Lepisma saccharina Linnaeus, 1758, and a Ctenolepisma sp. in regard to both the adaptations to the adaptive zone of Lepismatidae and to pre‐adaptations for the evolution of Pterygota. Well‐preserved parts of three subcoxal leg segments were found in the pleural zone participating in leg movement. The lepismatid strategy of escaping predators by running fast and hiding in narrow flat retreats led to a dorso‐ventrally flattened body which enabled gliding effects when dropped, followed by flight on the ground. The presumed exploitation of soft tissue at the tips of low growing Devonian vascular plants opened a canalized pathway to the evolution of the flying ability. Locomotion to another plant was facilitated by dropping. It is possible that threat by spider‐like predators favoured falling and gliding as escape reactions by selection. Falling experiments with `lepismatid' models revealed a narrow `window' for gliding, with optimum dimensions of 8 mm body length and 8 mg weight. An equation was derived which describes the glide distance as function of weight, area of the horizontal outline, the specific glide efficiency of the body, and a non‐linear function of the falling height. Improved gliding was made possible by enlarging thoracic paratergites into broad wing‐like extensions of light‐weight organization. The disadvantage of the lateral lobes for locomotion on the ground could be minimized by tilting them vertically when running and horizontally when gliding. This movability could be attained by the intercalation of a membranous strip between tergite and paratergite and the utilization of the pre‐existing muscular system and the articulation between the two most basal subcoxal sclerites as a pivot. The dorsal part of the most basal subcoxa was thus integrated into the wing. Initiation of active flight was possible by flapping movements during gliding. Morphological, ontogenetic and ecological aspects of the origin of Pterygota are discussed.     

The lateral thoracico-abdominal region in adults and fifth instar nymphs of an aquatic bug,Notonecta undulata say (Insecta,Heteroptera)

Skeletal differences in the lateral thoracico-abdominal regions of fifth instar and adult Notonecta appear to reflect respiratory differences in the two stages. Changes in the epidermis of this region during the last instar are described, and the possible relationships between the epidermis, nymphal cuticle, and imaginal cuticle are discussed.Explanation of Figures A Abdominal segment - AC Antecosta (anterior boundary of abdominal segment) - AP Abdominal projection of second abdominal segment - AT Anterior tracheolar branch - B Posteromedial boundary of posterior projection - C Metacoxa - CM Metacoxal membrane - CP Metacoxal process - CS Coxal-subalar muscle - DL Dorsal longitudinal muscle of abdomen - EL External lateral muscle of abdomen - EM Metathoracic epimeron - ES Metathoracic episternum - EV Posterior evagination of metanotum - F Fold of metacoxal membrane - FM Functional thoracico-abdominal membrane - FW Metathoracic wing or wingpad - H Horizontal ridge or sulcus on metathoracic postnotum - HW Metathoracic wing or wingpad - HWM Costal margin of hindwing - ILL Internal lateral muscle of abdomen, lateral portion - ILM Internal lateral muscle of abdomen, medial portion - IN Insertions of wingbase muscles on subalar epidermis - LE Lobe of metathoracic episternum - MM Posterior moulting muscle - MP Process for attachment of Muscle EL 1 - MSP Pleuron of mesothorax - MST Tergum of mesothorax - N Metathoracic notum - P Metathoracic pleural ridge or sulcus - PA Postalar portion of metathorax - PAB Metathoracic postalar bridge - PH Third phragma - PN Metathoracic postnotum - PP Posterior projection of metathoracic epimeron - PT Posterior tracheolar branch - PW Posterior end of metathoracic wingbase - S Abdominal spiracle - SA Subalar portion of metathorax - SAM Subalar membrane - SAS Subalar sclerite - SAV Widened ventrolateral portion of subalar air space - SL Scutellum of metathoracic notum - SM Spiracular membrane - ST Spiracular tracheole - SU Scutum of metathoracic notum - T Tendon of metacoxal remotor muscle - TA Thoracico-abdominal sclerite - TAA Anterior portion of thoracicoabdominal sclerite - TAC Posterior portion of thoraeicoabdominal sclerite - TAM Membranous portion of lateral thoracico-abdominal region - TS Metathoracic spiracle - VA Anterior end of ventral abdominal air channel - WB Junction between subalar wall and metathoracic wing or wingpad - WH Hairs on margin of mesothoracic wingpad     

Physiological roles of a yeast-like symbiote in reproduction and embryonic development of the brown planthopper,Nilaparvata lugens Stål

The physiological roles of a yeast-like symbiote in oviposition and embryonic development of its host, the brown planthopper, Nilaparvata lugens Stål, were studied using heat treatment, lysozyme injection and ligation of eggs. The eggs laid by the heat-treated females harboured only a few of the symbiotes, and their symbiote ball through embryonic development was free of symbiotes. The embryos of subsymbiotic eggs could not undergo blastokinesis and dorsal closure, and failed to hatch due to lack of differentiation of the abdominal segments. Electrophoretic profile of the eggs laid by the heat-treated females indicated the absence of several minor proteins which are usually found in the fat body of normal females. A protein (Y) of 131 kD was barely detectable in the heat-treated insects, and could not be found in the ligated eggs in which the symbiote ball was completely separated from the developing germ band. It is suggested that the symbiote supplies its host with proteins for normal embryonic and postembryonic development. The number of yeast-like symbiotes in female adults was reduced after injection with lysozyme solution, and some of the eggs were unable to hatch due to failure in blastokinesis; this was similar to the heat-treated insects. The embryos of ligated eggs could complete segmentation and differentiation normally before 110 h, but the abdominal segments failed to differentiate after dorsal closure, and regressed leaving only the head. Partially ligated eggs harboured some symbiotes and could produce normal larvae. It is concluded that the yeast-like symbiote is significant in abdominal segmentation and differentiation of the planthopper embryo.     

Embryonic development of the hemipteran insect Rhodnius prolixus

The embryonic development of the hemipteran insect Rhodnius prolixus was studied by use of contemporary light and electron microscopy. Embryos were staged according to days postoviposition. Eggs laid on day one complete blastoderm formation and anatrepsis, the first phase of blastokinesis, by day 5. The embryo develops in a cephalocaudal orientation which is 180° to the anteroposterior axis of the egg. Subsequent development, prior to the second phase of blastokinesis (katatrepsis), leads to segmentation of the germ band, evagination of appendages, and histogenesis of germ layers. Concomitantly with these events, the amnion undergoes dramatic change. By day 7 the embryo begins a 180° revolution while migrating to the ventral surface of the yolk. This restores its polarity with respect to that of the egg and facilitates hatching. The serosa contracts, pulling the amnion and embryo anteriorly. Eventually the serosa is internalized at a point dorsal to the head and the lateral walls of the embryo grow up and surround the yolk. Development continues until day 15 when the embryo hatches as a first instar larva.     

Embryonic development of Eucorydia yasumatsui Asahina,with special reference to external morphology (Insecta: Blattodea,Corydiidae)

As the first step in the comparative embryological study of Blattodea, with the aim of reconstructing the groundplan and phylogeny of Dictyoptera and Polyneoptera, the embryonic development of a corydiid was examined and described in detail using Eucorydia yasumatsui . Ten to fifteen micropyles are localized on the ventral side of the egg, and aggregated symbiont bacterial “mycetomes” are found in the egg. The embryo is formed by the fusion of paired blastodermal regions, with higher cellular density on the ventral side of the egg. This type of embryo formation, regarded as one of the embryological autapomorphies of Polyneoptera, was first demonstrated for “Blattaria” in the present study. The embryo undergoes embryogenesis of the short germ band type, and elongates to its full length on the ventral side of the egg. The embryo undergoes katatrepsis and dorsal closure, and then finally, it acquires its definitive form, keeping its original position on the ventral side of the egg, with its anteroposterior axis never reversed throughout development. The information obtained was compared with that of previous studies on other insects. “Micropyles grouped on the ventral side of the egg” is thought to be a part of the groundplan of Dictyoptera, and “possession of bacteria in the form of mycetomes” to be an apomorphic groundplan of Blattodea. Corydiid embryos were revealed to perform blastokinesis of the “non‐reversion type (N)”, as reported in blaberoid cockroaches other than Corydiidae (“Ectobiidae,” Blaberidae, etc.) and in Mantodea; the embryos of blattoid cockroaches (Blattidae and Cryptocercidae) and Isoptera undergo blastokinesis of the “reversion type (R),” in which the anteroposterior axis of the embryo is reversed during blastokinesis. Dictyopteran blastokinesis types can be summarized as “Mantodea (N) + Blattodea [= Blaberoidea (N) + Blattoidea (R) + Isoptera (R)]”.     

Embryogenesis of the dipluran Lepidocampa weberi Oudemans (Hexapoda,Diplura, Campodeidae): External morphology

The external features of the embryo of the dipluran, Lepidocampa weberi Oudemans are described. The long germ band is formed, and blastokinesis is a simple flexion of the germ band. The primary dorsal organ is formed between the cephalic and abdominal ends by concentration of serosal cells. The mouth fold is formed by ventral extension of the intercalary, mandibular, and maxillary terga, through which entognathy is completed. The posteroventral region of the mouth fold develops into the admentum. Rotation of the labial anlagen is involved in labial formation, and the glossa, paraglossa, and labial palp acquire a tandem arrangement. The postmentum is formed by fusion of the labial subcoxae and is appendicular in origin. The styli and exertile vesicles are derived from the distal parts of bifurcated appendicular anlagen of the second to seventh abdominal segments. The columnar appendage of the first abdominal segment is serially homologous with the exertile vesicles of the following segments. The abdomen is composed of ten segments, and the cercus is the appendage of the tenth, last abdominal segment. Embryogenesis of Lepidocampa weberi resembles that of the rhabduran Campodea staphylinus (Uzel, 1898) as well as that of the dicelluratan Japyx major (Silvestri, '33). It may be emphasized that the rhabduran and dicelluratan diplurans share important features such as entognathy formation and abdominal organization, and the resemblance between them seems to be close enough to postulate their close affinity. Some embryogenetic features, which Diplura and Collembola share, are recognized as plesiomorphic and the manner of entognathy formation may significantly differ. J. Morphol. 237:101–115, 1998. © 1998 Wiley-Liss, Inc.     

Anatomical changes in the neuromuscular complex of the proleg of Galleria mellonella (L.) (Lepidoptera: Pyralidiadae) during metamorphosis

The lateral and ventral external surfaces of the third and fourth abdominal segments were described and muscle attachments were correlated with surface indentations of the larva. The proleg of this species has a symmetrical planta with a complete circle of crochets. Furthermore, it differs externally from the grasping type of proleg in having a largely membranous coxal region confluent with the body wall, and a relatively large subcoxal lobe. The body wall musculature and innervation of the third and fourth abdominal segments are similar in many respects to those described for other lepidopteran larvae to which they are here compared, but differ from most because of the simpler structure of the prolegs which lack highly developed adductor muscles. Like most muscles innervated by the ventral nerve, the principal plantar retractors of these two segments cease to function in the first day of the pupal stage and have completely degenerated by the forty-fifth hour of pupal life. The ventral nerve retains its four primary branches in the adult, in which many smaller rami can be traced to the cuticle and to the neoblastic body wall muscles.     

Mantophasmatodea and phylogeny of the lower neopterous insects

Polyneoptera is a name sometimes applied to an assemblage of 11 insect orders comprising the lower neopterous or “orthopteroid” insects. These orders include familiar insects such as Orthoptera (grasshoppers), Blattodea (roaches), Isoptera (termites) (Mantodea) praying mantises, Dermaptera (earwigs), Phasmatodea (stick insects), Plecoptera (stoneflies), as well as the more obscure, Embiidina (web‐spinners), Zoraptera (angel insects) and Grylloblattodea (ice‐crawlers). Many of these insect orders exhibit a high degree of morphological specialization, a condition that has led to multiple phylogenetic hypotheses and little consensus among investigators. We present a phylogenetic analysis of the polyneopteran orders representing a broad range of their phylogenetic diversity and including the recently described Mantophasmatodea. These analyses are based on complete 18S rDNA, 28S rDNA, Histone 3 DNA sequences, and a previously published morphology matrix coded at the ordinal level. Extensive analyses utilizing different alignment methodologies and parameter values across a majority of possible ranges were employed to test for sensitivity of the results to ribosomal alignment and to explore patterns across the theoretical alignment landscape. Multiple methodologies support the paraphyly of Polyneoptera, the monophyly of Dictyoptera, Orthopteroidea (sensu Kukalova‐Peck; i.e. Orthoptera + Phasmatodea + Embiidina), and a group composed of Plecoptera + Dermaptera + Zoraptera. Sister taxon relationships between Embiidina + Phasmatodea in a group called “Eukinolabia”, and Dermaptera + Zoraptera (“Haplocercata”) are also supported by multiple analyses. This analysis also supports a sister taxon relationship between the newly described Mantophasmatodea, which are endemic to arid portions of southern Africa, and Grylloblattodea, a small order of cryophilic insects confined to the north‐western Americas and north‐eastern Asia, in a group termed “Xenonomia”. This placement, coupled with the morphological disparity of the two groups, validates the ordinal status of Mantophasmatodea. © The Willi Hennig Society 2005.     

Scanning electron microscopy studies on the first-instar larva of Dermatobia hominis

Abstract. First-instar larvae of Dermatobia hominis collected 1, 4 and 7 days after having penetrated experimentally infected rats, were studied by scanning electron microscope (SEM) observation. On the pseudocephalon there are basiconic and trichoid sensilla (antennal sensory complex), and basiconic, coeloconic and campaniform sensilla (maxillary sensory complex). The thoracic segments bear several rows of small, backwardly pointed, spines, and trichoid, campaniform, coeloconic and pit sensilla. The anterior spiracle is a minute opening. Both small and large spines directed posteriorly are on the first to fourth abdominal segments, which also bear coeloconic and companiform sensilla. These sensilla are present on the unarmed (fifth and sixth) and armed (seventh) abdominal segments. The seventh and the last (eight) abdominal segments have forwardly directed spines. Each spiracular plate has two spiracular openings and four spatulate-like structures called sun rays. The anus and the coeloconic sensilla are proeminent on the last segment. The results are compared with other parasitic dipteran larvae, and emphasize that the multiple types of sensilla on D. hominis larva may have importance in establishing the parasitic phase of the life cycle of this insect.