Venom gland and reservoir morphology in the Doryctinae and related braconid wasps (Insecta, Hymenoptera, Braconidae)

Detailed venom reservoir and venom gland intima morphology has been investigated in reprsentatives of 84 genera and 10 subfamilies of Braconidae with particular reference to the Doryctinac, basal cyclostomes and related groups including the Opiinae, Alysiinae and Rogadinae sens. lat . Several new phylogenetically significant characters are described and illustrated. Extcnsivc secretory ductules on the primary venom duct is suggested as a synapomorphy for the subfamilies Braconinac, Doryctinae, Opiinae, Alysiinae and Gnamptodontinae. The presence of two separate venom gland insertions and their position beyond the spiral sculpture of the reservoir are suggested as synapomorphies for the subfamilies Doryctinae and Ypsistocerinac. An anterior, weakly sclerotized, probably glandular. expansion of the reservoir characterizes the Histeromerinae and Rhyssalinae and also the enigmatic genera Thoracoplites Fischer and Doryctomorpha Ashmead which are hcre transferred to the Rhyssalinae. Within the Doryctinac, venom reservoir morphology suggests three generic groups, one comprising the genera of the Odontohraconini and possibly also the genera Acanthodoryctes , Binareu and Monarea , all of which have an essentially undivided reservoir with extremely fine and uniform spiral sculpture. the second comprising the genera Doryctophasmus , Euscelinus, Gildoria, Halycea and Schlettereriella , which share horn-like processes at the base of the venom duct insertions, and a third including Acrophasmus. Dendro- soter, Heterospilus, Megaloproctus, Rutheia, Paraspathius, Schlettereriella. Spathius and Syngaster based on the presence of two markedly different densities of annulation of the reservoir. The degree to which external and other characters support these groups is discusscd.     

Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa

Zaldivar‐River ó n, A., Areekul, B., Shaw, M. R. & Quicke, D. L. J. (2004). Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa. —Zoologica Scripta, 33, 223–237. The morphology of the venom apparatus intima in representatives of 38 genera of the problematic braconid wasp subfamily Rogadinae and other cyclostome braconids was investigated and a preliminary phylogenetic analysis for the group was performed with the information obtained. Despite the limited number of characters, the data suggest several relationships at various taxonomic levels. The venom apparatus in the Clinocentrini and the Stiropiini is relatively unmodified and similar to that found in other genera previously placed within a broader concept of the Rogadinae (e.g. genera of Lysitermini, Pentatermini, Tetratermini, Hormiini) and also to that of the Betylobraconinae. The presence of a cone of filaments located inside the secondary venom duct near to its insertion on the venom reservoir/primary venom duct is proposed as a synapomorphy for the tribe Rogadini to the exclusion of Stiropiini, Clinocentrini and Yeliconini. Other features of the secondary venom duct and its insertion on the venom reservoir/primary venom duct support a number of relationships between the genera of the Rogadini and also within the large genus Aleiodes. A clade containing 15 Rogadini genera (Bathoteca, Bathotecoides, Bulborogas, Canalirogas, Colastomion, Conspinaria, Cystomastacoides, Macrostomion, Megarhogas, Myocron, Pholichora, Rectivena, Rogas, Spinaria and Triraphis) is supported by the presence of a thickened and short secondary venom duct, whereas the different members of Aleiodes (excluding members of the subgenus Heterogamus) and Cordylorhogas are distinguished by having a recessed secondary venom duct with well‐defined and numerous internal filaments. New World Rogas species exhibit a unique venom apparatus and may not be closely related to the Old World ones. Features of the venom apparatus of the enigmatic genus Telengaia and the exothecine genera Shawiana and Colastes suggest that the Telengainae and Exothecinae are both closely related to the Braconinae, Gnamptodontinae, and possibly to the Opiinae and Alysiinae. An unsculptured venom reservoir was found in one specimen of the type species of Avga, A. choaspes, which is consistent with it occupying either a very basal position within the cyclostome braconids or belonging to a recently recognized ‘Gondwanan’ clade that also includes the Aphidiinae.     

Morphology and ultrastructure of a secretory region enclosed by the venom reservoir in social wasps (Insecta,Hymenoptera)

The morphology and ultrastructure of the convoluted gland inside the venom reservoir of four species of social Vespidae are described. The cells of the venom gland (including the convoluted gland) can be divided into six groups: (1) epithelial cells, (2) glandular cells with the end apparatus secreting into the tubule inside the convoluted gland (internal or embedded tubule), (3) a continuous arrangement of glandular cells with the end apparatus secreting directly into the venom reservoir, (4) glandular cells that are loosely dispersed along the tubule lumen between the free tubules and the embedded tubule of the convoluted gland, (5) secretory cells of the free tubules and (6) duct cells. One kind of secretory cell, hitherto unknown and described in this paper (group 3), is characterized by the presence of a well-developed end apparatus, usually with enlarged extracellular spaces, but lacking the normally associated duct cells. The secretory cells contain several stacks of granular endoplasmic reticulum, but these are mainly concentrated in the middle of the cell. The basal half of the cells contains many lipid droplets. Although the function of the convoluted gland is not yet understood, an hypothesis is related to what is known of the function of reservoir secretory cells in solitary wasps. All wasp species studied showed the same organization of the convoluted gland, which clearly distinguishes their venom gland from that of Sphecidae.     

Ovipositor internal microsculpture and other features in doryctine wasps (Insecta, Hymenoptera, Braconidae)

Detailed ovipositor morphology has been investigated in representatives of 81 of the more than 120 genera of the parasitic wasp subfamily Doryctinae. Potentially phylogenetically informative variation was found in the microsculpture of the egg canal wall and the presence and degree of development of the valvillus. In all the genera, a socketed seta arises from below each member of the line of ctenidia (comb-like scales) distal to the valvillus (or valvillar region). These subctenidial setae are modified in two groups of genera. In the Holcobraconini (= Odontobraconini) together with the genera Acanthodoryctes, Binarea, Monarea , and Liobrucon , the setae are extremely flattened producing overlapping leaflet-like structures. In some of these, groups of leaflets are further modified to form fans that protrude into the egg canal. In Schlettereriella and Leptospathius the basal seta is highly branched and rather flattened and may serve a similar function to the valvillus which in these two genera is extremely reduced. In Doryctes and Neodoryctes , there is a distinctive type of bar-like sculpture anterior to the valvillus and a single large crescentic bar posterior to the valvillus. These modifications are suggested as possible synapomorphies (in lieu of an analysis) for their respective groups of genera. Several other characters are also described and illustrated. CO 1998 The Norwegian Academy of Science and Letters     

Ovipositor internal microsculpture in the Braconidae (Insecta, Hymenoptera)

Variation in the microsculpture of the ovipositor egg canal, the morphology of the tip of the upper valve, the structure of the valvillus and the apex of the lower valve seals, as revealed by scanning electron microscopy is described for members of 52 genera representing 30 subfamilies of Braconidae. The possible phylogenetic implications of this variation are discussed. Characters supporting monophyly of the cyclostome clade, and of the non-cyclostome subfamilies Agathidinae, Pselaphaninae and Sigalphinae are reported. Functional and phylogenetic implications are discussed and possible convergent evolution of some features associated with endoparasitism are considered in the light of recent molecular systematics findings. © 1998 The Norwegian Academy of Science and Letters     

Mendesellinae, a new subfamily of braconid wasps (Hymenoptera, Braconidae) with a review of relationships within the microgastroid assemblage

Abstract. A new subfamily of braconid wasps, the Mendesellinae, is proposed and described. It is represented by two genera and nine species, all newly described: Mendesella gen.n., type species M.amazonica sp.n., M.braziliensls , sp.n., M.magna sp.n., Epsilogaster gen.n., type species E.bicolor sp.n., E.brazMensis sp.n., E.dureno sp.n., E.palenque sp.n., E.tico sp.n. and E.panama sp.n. The phylogenetic relationships among the subfamilies of the 'microgastroid assemblage' of Braconidae are reviewed and analysed, to clarify the status and relationships of the new subfamily.     

New braconid wasps from French Cretaceous amber (Hymenoptera, Braconidae): synonymization with Eoichneumonidae and implications for the phylogeny of Ichneumonoidea

Two new fossils of Braconidae are described from Albian-Cenomanian amber of south-western France, Protorhyssalodes arnaudi gen. n., sp. n., and Aenigmabracon capdoliensis gen. n., sp. n. The former appears superficially similar to the type genus and species of the extinct sub-family Protorhyssalinae, from Turonian New Jersey amber specimens, and the latter both to Protorhyssalus and to members of the extinct family Eoichneumonidae. However, both new taxa display unique combinations of wing venation characters making confident assignment to sub-family impossible. Indeed, they are the first braconids ever known to possess both vein 2-CU and a distinct trace of vein 2-1A on hindwing. The new fossil taxa are incorporated into a morphological analysis of extinct and extant ichneumonoids. As a result of the analyses we synonymize the Eoichneumonidae with the Braconidae.     

Venom gland and reservoir morphology in cynipoid wasps

The venom apparatus morphology was examined in 25 species of Cynipoidea, representing 11 parasitoid taxa; 12 gall inducers and two inquilines. Typically the venom apparatus consists of an oval or bilobed reservoir connected to the ovipositor apparatus by a very short venom duct at the anterior end and to a single elongate unbranched venom gland at or near its posterior end. The Dufour's gland was not found in any of the examined species. The elongate unbranched venom gland and the absence of the Dufour's gland are putative cynipoid synapomorphies. The shape and size of especially the venom reservoir were found to vary considerably within the Cynipoidea. It is typically less prominent in the parasitoid taxa than in the gall inducers. Exceptions include the poppy gallers Barbotinia and Aylax, in which the venom reservoirs were remarkably small and in the rose galler Diplolepis, where only a rudimentary venom apparatus was found. Possible functional and phylogenetic implications of cynipoid venom apparatus features are discussed.     

A new opiine (Hymenoptera,Braconidae) from Australia with discussion of Diachasma Foerster

A new species of Opiinae, Diachasma dentatum Shirley, Restuccia & Ly, is described from Australia. This species is similar to several other Australian opiines previously described or included in the genus Diachasma, but the mandibles are unusually broad, nearly exodont. Notable differences between Australian and Palaearctic Diachasma are discussed. Diachasma tasmaniae Fischer, 1995, originally described from Tasmania and New South Wales, is newly recorded from Victoria. Diachasma rufipes Szépligeti, 1905 is transferred to Notiopambolus, new combination.     

A new Indo-Australian genus of Braconinae (Insecta, Hymenoptera, Braconidae)

Bicarinibracon gen.n. (type species Atanyco1us tricolor . Szépligeti from Papua New Guinea and Irian Jaya). Bicarinibracon luteus gen. et. 3p.n. from India and Malaya and B. lepicius gen. et. sp.n. from Thailand and the Philippines are described and illustrated. A key to the specics of Bicarinibracon is provided and its systematic placement is discussed.     

Ichneumon wasps of the Khasurty locality in Transbaikalia (Insecta, Hymenoptera, Ichneumonidae)

One new genus and four new species of ichneumon wasps are described from the Lower Cretaceous of Buryatia (Khasurty locality): Amplicella mininae, sp. nov., A. shcherbakovi, sp. nov., A. bashkuyevi, sp. nov., Khasurtella buriatica, gen. et sp. nov. The species Tanychora sinensis Zhang, 1991 is transferred to the new genus Khasurtella. Similarity of the ichneumonid faunas of Khasurty and the Chinese localities Yixian and Laiyang indicates the age similarity between these localities. Identification key to genera and species of Tanychorinae is given.     

Inter-generic variation in the male genitalia of the Braconinae (Insecta, Hymenoptera, Braconidae)

Male genitalia belonging to representatives of a total of 72 genera of Old and New World Braconinae have been examined by light and scanning electron microscopy. On the basis of features of the volsellae (setosity), the parameres (setosity), the digitus (number of teeth, shape) and basal ring (posterior extension) it was possible to identify certain generic groupings which support the general suprageneric divisions of the subfamily based on other taxonomic characters. The male genitalia are thus shown to provide a potentially useful new set of characters for the higher level classification of the braconine genera.     

Terga1 and inter–tergal metasomal glands of male braconine wasps (Insecta, Hymenoptera, Braconidae)

The distribution of cxocrine tergal and inter–tergal glands among males of more than sixty genera of Braconinac has been surveyed. The morphology of the glands and their reservoirs is described. Two gland types were found. Large, sac–like inter–tergal glands, usually most prominent between the 7th and 8th abdominal tergites, were found in all members of the Braconini, Adeshini, Aphrastobraconini, Glyptomorphini and the Atanycolus and Merinotus groups of genera. Inter–tergal glands were generally considerably smaller in members of the Bathyaulacini, Coeloidini. Euurobraconini and Iphiaulacini. Small pore–plate, vase–like or tubular tergal glands opening on the medio–lateral or antero–lateral selerotized parts of the 8th metasomal tergite were found in many genera, but were largely absent from members of the Braconini, Euurobraconini and Iphiaulacini ss. The phylogenetic implications of these findings are discussed.     

Three new genera of Braconini from Australasia and Malaysia (Insecta, Hymenoptera, Braconidae)

Ancilibracon gen.n. (type species: Ancilinracon townesi sp.n.). Ancilibracon bakeri sp.n., Gelasinibracon gen.n. (type species: Gelasinibracon sedlaceki sp.n.), Gelasinibracon simplicicaudatus sp.n., Esengoides gen.n. (type species: Esengoides crentdatus sp.n.) and Esengoides fulvus sp.n. are each described and illustrated. Ancilibracon towensi is from Malaysia. A. bakeri is from Borneo, both Gelasinibracon species are from Guinéa, E. crenulatus is from the solomon Islands and E. fulvus is from Australia. These three new genera belong to the Plesiobrscon Cameron group of the Braconina. Their relationship are discussed.     

A new species of Paxillibracon Quicke (Insecta, Hymenoptera, Braconidae) from Tanzania

Paxillibracon inornatus sp.n. from Tanzania is described, illustrated and distinguished from Paxillibracon tessmanni (Szépligeti), the only other known species of the genus. The scapus, antennal sockets, features of the venation and metasomal sculpture of P. inornatus sp.n. reinforce the earlier interpretation that Paxillibracon is derived from close to the base of the Atanycolus Foerster group of genera.     

Two new genera and species of Braconinae (Insecta, Hymenoptera, Braconidae)from Brunei

Fuvihracon gen.n. (typc-species: Fuvibracon gauldi sp.n.) and Spurhulibrucon (type-species: Sparhulibracon dayi sp.n.), both lrom Brunei, are described and illustrated. The relationships of the two new genera and of the Carnpyloneurus Szépligeti and Chelonogaslra Ashmead generie groups are discussed.     

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.     

New maimetshid wasps in Cretaceous amber from Myanmar (Insecta: Hymenoptera)

The extinct, exclusively Cretaceous wasp family Maimetshidae is newly recorded from Earliest Cenomanian Burmese amber. Two new genera and species are described. Burmaimetsha concava gen. et sp. nov., based on a male and a female, is most similar to Guyotemaimetsha Perrichot, Nel & Néraudeau, from Albian-Cenomanian French amber, but differs in its larger mandibles, distinctly concave face, elongate antennomeres, and forewing with cell [1Rs] smaller and fourth abscissa of Rs shorter. Maimetshasia kachinensis gen. et sp. nov., is based on a male, and is characterized by asymmetric mandibles with two and three teeth, by its forewing venation without cross-vein 2rs-m, with cell [1M] large and trapezoidal, and vein 2Rs + M very short, and by the hind wing without a free apex of Rs. The family was evidently widespread in the Cretaceous, and the new records extend the paleobiogeographical range to the South-East of Eurasia. A discussion about the possible biology of Maimetshidae is provided.     

A new, putatively primitive Cretaceous fossil braconid subfamily from New Jersey amber (Hymenoptera, Braconidae)

Protorhyssalus goldmani gen. n., sp. n., in a new subfamily of braconid wasps, the Protorhyssalinae, is described from Late Cretaceous amber fossils from New Jersey, USA. The Protorhyssalinae appears to be cyclostome and shows a similar set of plesiomorphic characters to the extant Rhyssalinae. However, it possesses hindwing vein 2-CU, a feature only found among the cyclostome braconids in the rare and putatively primitive Chilean subfamily Apozyginae.