A new genus and species of Braconinae from Australia (Insecta, Hymenoptera, Braconidae)

Euurobraconoides gen.n. (type-species: Euurobraconoides longicaudis sp.n.) from Australia is described and illustrated. The relationships of Euurobraconoides gen.n. to the Euurobraconini Ashmead and to the Braconini are discussed.     

Cockroach mating behaviors,sex pheromones,and abdominal glands (Dictyoptera: Blaberidae)

Two chemical signals are essential in all cockroach sexual behavioral sequences: the sex pheromone released by one partner, generally the female (for long distance attraction), and an aphrodisiac sex pheromone produced exclusively by male tergal glands (for female mounting and tergal contact or feeding behavior). Unlike the other cockroach groups, the males of the Oxyhaloinae species produce both chemical signals: the pheromone and the aphrodisiac. The occurrence of three patterns of mating behavior (A, B, and C), the production of male sex pheromones, and the existence in the male of developed sternal and tergal glands in seven related Oxyhaloinae species, make these cockroaches a useful model for studying the evolution of mating behavior patterns. The various types of mating behavior were not classified in the previous studies by Roth and Barth. In this report, they have been named type A (female in upper position), B (male in upper position), and C (male and female end to end). In type A mating, the male tergal glands, which are licked by the females, are well developed, whereas in types B and C, there is no licking of the male's tergal secretion by the females and the tergal glands are much less developed; the aphrodisiacs secreted by the tergal glands may no longer act in this case through contact chemoreception, but through an olfactory process involving volatile components. One common sex pheromone component seems to be acetoin. I suggest that the mating behavior tends from A toward B and C during the evolutionary process with a concomitant regression of the tergal glands and changes in the aphrodisiac emission levels. The mating behavioral sequences of cockroaches (Dictyoptera) and crickets (Orthoptera) show a striking degree of similarity and are probably examples of convergent evolution.     

Ultrastructure of the glands producing sex pheromones of the male Nauphoeta cinerea (Insecta,Dictyoptera)

Summary The abdominal glands described here play a decisive role in the typical sexual behavior of Nauphoeta cinerea. Unlike other cockroaches, the males of this species produce two successive chemical signals: the sex pheromone itself, produced by the sternal glands, attracts the female from a distance, and the aphrodisiacs, secreted by the tergal glands, are licked by the female who is thus in a favorable position for mating. The well developed glandular apparatus is composed of 5 sternal glands (St3 to St7) and 7 tergal glands (T2 to T8). These glands appear as a thickening of the epithelium without significant modification of the external cuticle. The glandular epithelium is made up of several kinds of cells: ordinary epidermal cells (which only exist in the sternal glands), cells with microtubules, type 2 cells (oenocytes), and especially type 3 glandular units (composed of a secretory cell and a canal cell). The products secreted by the sternal glands are chiefly volatile products and fatty acids and those secreted by the tergal glands are primarily fatty acids and proteins. In this work, the relationship between the cytology of the glandular cells and the nature of the secreted products is discussed.     

Fissuroderes, a new genus of Kinorhyncha (Cyclorhagida) from the deep sea and continental shelf of New Zealand and from the continental shelf of Costa Rica

Specimens of Kinorhyncha collected by RV Sonne from the continental shelf off the coast of Costa Rica and from the deep sea East of New Zealand as well as by RV Kaharoa in the Firth of Thames are identified as five new species of a new genus Fissuroderes gen. nov. which is distinguished from all other kinorhynch genera by a ring-like cuticle in the first trunk segment and a cuticle with midventral and lateral articulations resulting in two sternal plates and one tergal plate in segments 2-10. Fissuroderes higginsi sp. nov. can be distinguished from all other species of Fissuroderes by a laterodorsal spine in segment 2 and the lack of a lateroventral spine in segment 8. Fissuroderes rangi sp. nov. is recognized from all other species by the lack of a ventrolateral spine in segment 2, the lack of a lateroventral spine in segment 5, an extremely elongate spinose tergal extension of segment 11 and a prominent midventral spinose process of the sternal plates in segment 11. Fissuroderes papai sp. nov. is identified by the short spinose process of the sternal plate of segment 11. Only Fissuroderes thermoi sp. nov. possesses bilobed sternal plates in segment 11 and a short conical tergal extension. Fissuroderes novaezealandia sp. nov. differs from F. higginsi sp. nov. in the lack of a laterodorsal spine in segment 2, the lack of an accessory spine in a lateral position in segment 8, the lack of a sublateral spine in segment 10, the existence of a lateroventral spine in segment 8 and in the lack of a subdorsal type 2-gland cell outlet in segment 2. The former species differs from F. papai sp. nov. in the lack of a lateroventral spine in segment 8, the lack of longitudinal indentations in the posterior part of the trunk cuticle of each segment and in the existence of an elongate spinose tergal extension of segment 11 and of many fine hairs on the surface of the cuticle. Fissuroderes is included into the Echinoderidae which is suggested to enclose also the genera Echinoderes and Cephalorhyncha. Echinoderes nybakkeni Higgins, 1988 is now combined as Cephalorhyncha nybakkeni (Higgins, 1988) comb. nov.     

Morphology of the sternal and tergal glands producing the sexual pheromones and the aphrodisiacs among the cockroaches of the subfamily oxyhaloinae

Among cockroaches in the subfamily of Oxyhaloinae, the adult males produce two essential and successively active chemical signals: the sex pheromone attracting females from a distance secreted by the sternal glands, and the aphrodisiac required for mating which is secreted by the tergal glands. The adult males of the seven species studied, Nauphoeta cinerea, Henschoutedenia flexivitta, Leucophaea maderae, Jagrehnia madecassa, Gromphadorhina portentosa, G. laevigata, and G. chopardi, possess well-developed sternal and tergal glands whose number varies according to the species and methods of mating (three to six sternal glands and four to seven tergal glands). These glands are basically composed of type three glandular units (glandular cell + duct cell) and type 2 cells (modified oenocytes) which exhibit no significant external cuticular modification except for tergite 2 of L. maderae. The extreme variance in development of these glands can be linked to sexual behavior. The hypothesis put forward here is that of a regressive evolution of the tergal glands, related to a modification of the role played by the aphrodisiacs which they secrete.     

The nasus gland: A new gland in soldiers of Angularitermes (Termitidae,Nasutitermitinae)

Termites have developed many exocrine glands, generally dedicated to defence or communication. Although a few of these glands occur in all termite species, or represent synapomorphies of larger clades, others are morphological innovations of a single species, or a few related species. Here, we describe the nasus gland, a new gland occurring at the base of the nasus of Angularitermes soldiers. The nasus gland is composed of class 1, 2, and 3 secretory cells, a rare combination that is only shared by the sternal and tergal glands of some termites and cockroaches. The ultrastructural observations suggest that the secretion is produced by class 2 and 3 secretory cells, and released mostly by class 3 cells. The base of the nasus has a rough appearance due to numerous pits bearing openings of canals conducting the secretion from class 3 secretory cells to the exterior. We tentatively assign a defensive function to the nasus gland, although further research is needed to confirm this function. Although the gland is described only from species of Angularitermes, other genera of Nasutitermitinae also present a rough nasus base, suggesting the presence of a similar, possibly homologous, gland.     

Detailed analysis of the prothoracic tissues transforming into wings in the Cephalothorax mutants of the Tribolium beetle

Despite the immense importance of the wing in the evolution and successful radiation of the insect lineages, the origin of this critical structure remains a hotly-debated mystery. Two possible tissues have been identified as an evolutionary origin of wings; the lateral expansion of the dorsal body wall (tergal edge) and structures related to an ancestral proximal leg segment (pleural tissues). Through studying wing-related tissues in the red flour beetle, Tribolium castaneum, we have previously presented evidence in support of a dual origin of insect wings, a third hypothesis proposing that wings evolved from a combination of both tergal and pleural tissues. One key finding came from the investigation of a Cephalothorax (Cx) mutant, in which the ectopic wing characteristic to this mutant was found to be formed from both tergal and pleural contributions. However, the degree of contribution of the two tissues to the wing remains elusive. Here, we took advantage of multiple Cx alleles available in Tribolium, and produced a variety of degrees and types of ectopic wing tissues in their prothoracic segments. Through detailed phenotypic scoring of the Cx phenotypes based on nine categories of mutant traits, along with comprehensive morphological analysis of the ectopic wing tissues, we found that (i) ectopic wing tissues can be formed at various locations in the prothorax, even internally, (ii) the lateral external ectopic wing tissues have tergal origin, while the internal and posterior external ectopic wing tissues appear to be of pleural origin, and (iii) the ectopic wing tissues of both tergal and pleural origin are capable of transforming into wing surface tissues. Collectively, these outcomes suggest that the evolutionary contribution of each tissue to a complete wing may be more complex than the simple binary view that is typically invoked by a dual origin model (i.e. the wing blade from the tergal contribution + musculature and articulation from the pleural contribution).     

Sexual communication in the termite Prorhinotermes simplex (Isoptera, Rhinotermitidae) mediated by a pheromone from female tergal glands

We studied the post-flight behavior and sex attraction in imagoes of the termite Prorhinotermes simplex (Rhinotermitidae, Prorhinotermitinae). Pairing is mediated by the secretion from tergal glands, exposed by females in a calling posture and highly attractive to males. Analysis of extracts of these glands by means of gas chromatography with electroantennographic detection indicated a chromatographic area corresponding to an intense physiological response of males. The retention characteristics of this area proved to be identical with those of (3Z,6Z,8E)-dodeca-3,6,8-trien-1-ol. Electroantennographic and behavioral assays revealed that units of picograms of the compound represent a stimulus qualitatively and quantitatively equivalent to one female tergal gland. Thus, we hypothesize that (3Z,6Z,8E)-dodeca-3,6,8-trien-1-ol is a major component of the female sex pheromone in P. simplex.     

Structure and evolution of stridulatory mechanisms in New Zealand wetas (Orthoptera : Stenopelmatidae)

Four types of stridulatory mechanisms were found in 35 species (7 genera) of New Zealand wetas (Orthoptera : Stenopelmatidae). These include: (1) tergo-femoral; (2) tergo-tergal; (3) mandbbuao-mandibular; and (4) pleuro-coxal. These fell into 12 groups, based upon shape, density and pattern of tergal and femoral pegs, number of terga bearing pegs and presence of mandibular tusks bearing stridulatory tubercles. The plesiomorphic condition, consisting of spinose peg patches on 3 or more abdominal tergites rubbed by bands of pegs on the femur, is found in Hemiandrus spp. Apomorphically-derived tergal files have arisen separately in 3 other genera. In Hemideina, (2 species groups) well-formed files are very similar among species. In Deinacrida, (4 species groups), the trend is toward file reduction into one or 2 massive tergal ridges, and embellished femoral pegs. In the tusked wetas (2 gen., 2 spp.), one species has a crude file of many broken ridges. Stridulatory structures on mandibular tusks of both New Zealand species are unique, although the tusks appear to have their origin in African stenopelmatids. The plesiomorphic condition appears to have been deployed originally for defense stridulation (inter-specific communication). Additional intea-specific stridulatory communication developed in Hemideina and Deinacrida, using the apomorphic file and peg mechanisms. Here, the stridulation is associated with defense, calling, mating and disturbance behaviors.     

Morphology of the male and female tergal glands of the woodroach Cryptocercus punctulatus (Insecta,Dictyoptera)

Summary Males and females of Cryptocercus punctulatus possess tergal glands which differ in position, size, morphology, and chemical composition of their secretions. Ultrastructural studies reveal the presence of class 1 and class 3 glandular units interspersed throughout the glands; class 3 glandular units are 30 times as numerous as in the female, but no cytological difference was found between the sexes. The morphology of the tergal glands is characterized by the occurrence of a subcuticular space reservoir, a dense tracheal system, and a thick collagenous layer instead of the typical basement membrane. Comparison with the termite abdominal gland system indicates a great similarity in the fine structure. The biological function of the tergal glands is discussed.Abbreviations A Class 1 gland cell - B Class 3 gland cell - cf Collagen fibrils - d Duct - ea End-apparatus - EC Endocuticle - ef Epicuticular filaments - ep Epicuticle - j Septate junctions - m Mitochondria - mt Microtubules - mv Microvilli - pc Pore-canal - R Intracuticular reservoir - rc Receptor canal - S Saccule cell - t 6–10 Terga 6 to 10 - TG Tergal gland - tr Trachea - * Subcuticular space - Secretory material - Secretory vesicles - Pinocytosis vacuoles     

Phylogeny of the Braconinae (Hymenoptera: Braconidae): A new tribal order!

Generic relationships within the parasitoid wasp subfamily Braconinae are assessed based on a molecular phylogenetic analysis of four gene fragments: mitochondrial cytochrome c oxidase subunit I, 16S rDNA, nuclear 28S D2-D3 rDNA and elongation factor 1-alpha. Our results support the recognition of Aphrastobraconini, Braconini and Coeloidini plus three new tribes: Compsobraconini tribus nov ., Tropobraconini tribus nov . and Virgulibraconini tribus nov . The first of these new tribes is restricted to the New World; the second includes the Old World genera Tropobracon Cameron, Trispinaria Quicke and Grangerbracon Samartsev and Belokobylskij and possibly others, whereas the third comprises the Australian genus Virgulibracon Quicke, plus several other described and undescribed Australian genera. Consistent placement of Amyosoma Viereck with members of the Virgulibraconini tribus nov . is discussed, whereas Amyosoma is left currently unplaced. A preliminary key to tribes is presented, and the characters used to differentiate between Aphrastobraconini and Braconini are revised. Megacoeloides Quicke was never recovered with Coeloides, the type genus of Coeloidini, so it is treated as Braconinae incertae sedis. By combining molecular and morphological traits, nearly all valid genera are assigned to tribes, and the possible relationships of the remainder are discussed. Alienoclypeus Shenefelt, 1978 is synonymised with Atanycolus Förster, 1862 (Atanycolus insolitus (Shenefelt) comb. nov .). Several new genera have been revealed and will be described elsewhere.     

A new kinorhynch genus,Mixtophyes (Kinorhyncha: Homalorhagida), from the Guinea Basin deep-sea,with new data on the family Neocentrophyidae

A new homalorhagid genus and species, Mixtophyes abyssalis gen. et sp. nov., is described from the deep-sea of the Guinea Basin in the Atlantic Ocean. The new genus and species was collected during the R/V Meteor Diva2 M63/2 Cruise, and it is characterized by a trunk with segment 1 consisting of a tergal and a single sternal plate, without any subdivisions or differentiations, segment 2–10 of one tergal and two sternal plates and segment 11 of a tergal and a sternal plate. Lateral terminal spines are present in both sexes. As part of the study of the new taxon, the type material of Neocentrophyes intermedius and N. satyai was re-examined, and new information is provided for both species. Based on the information from Mixtophyes abyssalis gen. et sp. nov. and re-examination of the two Neocentrophyes species, emended diagnoses for Neocentrophyes and Neocentrophyidae are proposed, and an updated identification key to homalorhagid genera and species of Neocentrophyidae is provided. Mixtophyes abyssalis gen. et sp. nov. is tentatively assigned to Neocentrophyidae, but evaluation of our current hypotheses about homalorhagid evolution and phylogeny also indicated that the family most probably is paraphyletic and that revision of homalorhagid classification is required when new results from phylogenetic analyses become available.     

Comparative morphology of the venom gland and reservoir in opiine and alysiine braconid wasps (Insecta, Hymenoptera, Braconidae)

Detailed venom reservoir and venom gland intima morphology was investigated in representatives of 178 species and 76 genera of Braconidae belonging to the subfamilies Opiinae (23 genera and 69 species) and Alysiinae (36 genera and 67 species of Alysiini, and 17 genera and 42 species of Dacnusini). The presence of an unsculptured sack–like expansion of the anterior end of the reservoir suggests a relationship between some members of the alysiine tribe Dacnusini, viz. Exotela and Chorebus , and the genera Oenonogaster and Glyphogaster , both of which are currently placed in the Alysiini. The same feature also suggests that Exotela, Chorebus, Oenonogaster and Glyphogaster may be derived from the Opiinae separately from other Alysiinae. Other dacnusines examined are indicated as forming a monophyletic taxon on the basis of their reduced and narrow reservoirs. An anterior insertion of the venom glands on to the reservoir generally supports the Phaenocarpa group of alysiine genera, with some modification, whereas the presence of a discrete, narrow posterior extension of the reservoir represents a synapomorphy for members of the Aspilota group of alysiine genera. The phylogenetic inferences from this venom apparatus study are discussed in the light of the biology of the taxa concerned. Twenty-one new combinations are made in the Opiinae and seven in the Alysiinae.     

Effects of the juvenile hormone analog pyriproxyfen on German cockroach, Blattella germanica (L.), tergal gland development and production of tergal gland secretion proteins

Male German cockroaches possess secretory glands that secrete fluid into a pair of transverse depressions on the seventh and eighth abdominal tergites. We investigated the effects of altered juvenoid titer during the first part of the last instar on tergal gland secretory tissue development and the production of tergal gland secretion proteins. Male fifth (last) instar nymphs (1-3 days post-emergent) were topically treated with the JH analog pyriproxyfen. Light and transmission electron microscopy demonstrated that treatment with pyriproxyfen produced a visible decrease in the amount of tergal gland tissue present, a deformation of the overall shape of the gland located on tergite seven, and a less orderly arrangement of the secretory cells in the tissue. The protein fraction of tergal gland secretion was examined in pyriproxyfen-treated and control insects 1, 5, and 15 days after the insects molted to the adult stage. Amounts of all tergal secretion proteins were reduced in treated insects.     

A remarkable new genus of Braconini (Hymenoptera: Braconidae: Braconinae) from Papua New Guinea,reared from fallen fruit

During an investigation of the insect food web in Papua New Guinea several specimens of a new genus and species of large, black and orange braconine braconid wasp were reared from fruits of Rubiaceae. These are described here as Karposibracon papuensis Quicke & Butcher gen. et sp. nov. Karposibracon gen. nov. appears to belong to the tribe Braconini and is the largest known species in this group, with body length up to 12?mm. The new genus is illustrated photographically, distinguished from other genera, and comments are on its likely biology provided.     

Chemical signals of queens in kin recognition of honeybees,Apis mellifera L.

Summary The volatile signals of live queens as well as head and tergite extracts were analyzed by capillary gas chromatography. Multivariate analysis revealed a significant decrease in the variability of the volatile signals of queens the more closely related they were. Though extracts of the tergal glands showed the same phenomenon, head extracts of closely related queens had no significantly reduced variabilities compared to extracts of unrelated queens. Workers, tested in a metabolic bioassay, could not discriminate among head extracts of related and unrelated queens. Therefore the classical queen substance and the semiochemicals of the mandibular glands are unlikely to be used as kin recognition labels by the workers. Workers could, however, discriminate among the odours of tergite extracts and the volatile signals of queens, indicating that the signal of the tergal gland secretions is a possible source for kin recognition labels which the mandibular gland secretions are not. Furthermore the results show that learning is strongly involved in the kin recognition system of honeybees.     

Sex-pairing pheromones and reproductive isolation in three sympatric Cornitermes species (Isoptera, Termitidae, Syntermitinae)

The species-specificity of pairing has been studied in three sympatric Neotropical termites: Cornitermes bequaerti, Cornitermes cumulans and Cornitermes silvestrii (Termitidae, Syntermitinae). Bioassays showed that sex attraction was highly species-specific between C. bequaerti and C. cumulans but not between C. cumulans and C. silvestrii. The sex-pairing pheromone of the three species is secreted by the tergal glands of female alates. It consists of a common compound (3Z,6Z,8E)-dodeca-3,6,8-trien-1-ol. In C. bequaerti, this polyunsaturated alcohol is the only compound of the sex-pairing pheromone, whereas it is associated with the oxygenated sesquiterpene (E)-nerolidol in C. cumulans, and with (E)-nerolidol and (Z)-dodec-3-en-1-ol in C. silvestrii. (3Z,6Z,8E)-Dodeca-3,6,8-trien-1-ol is responsible for sexual attraction, whereas (E)-nerolidol, which is inactive in eliciting attraction of male alates, is responsible for the species-specificity of the attraction. This is the first time that a multicomponent sex-pairing pheromone has been identified in termites. The role of (Z)-dodec-3-en-1-ol present on the surface of the tergal glands of the female alates of C. silvestrii could not be definitively determined, but it is suggested that this compound could be involved in the species-specificity of sex attraction with other sympatric species of Cornitermes. Our study shows that the reproductive isolation in termites is due to a succession of factors, as the chronology of dispersal flights, the species-specificity of sex-pairing pheromones and the species-specific recognition.     

Ultrastructure of posterior sternal glands of Macrotermes annandalei (Silvestri): new members of the sexual glandular set found in termites (Insecta)

In female alates of Macrotermes annandalei, two types of abdominal glands are involved in the secretion of sex pheromone. Tergal glands are found at the anterior margin of tergites 6-10 and posterior sternal glands (PSGs) are located at the anterior margin of sternites 6-7. The cytological features of both types of glands are quite similar. The fine structural organization of PSGs is studied more precisely and described for the first time. The glandular cuticle is pitted with narrow apertures corresponding to the openings of numerous subcuticular pouches. Several Class 3 glandular units open in each pouch. One canal cell and one secretory cell make an individual glandular unit. The canal cell is enlarged apically and is connected with the other canal cells to form a common pouch. Based on the structural features found in these glands, we propose a common secretory process for PSGs and tergal glands. During the physiological maturation of alates inside the nest, secretory vesicles amass in the cytoplasm of secretory cells, while large intercellular spaces collapse the cuticular pouches. At the time of dispersal flight, pouches are filled with the content of secretory vesicles while intercellular spaces are sharply reduced. After calling behavior, no secretion remains in the glands and pouches collapse again, while secretory cells are drastically reduced in size. The structure and the secretory processes of PSGs and tergal glands are compared to those of abdominal sexual glands known in termites.