The ovipositor musculature of Carpomya schineri (Loew) (Diptera, Tephritidae)

The structure of the ovipositor sclerites and musculature was studied in the tephritid fly Carpomya schineri (Loew) whose larvae develop in the fruits of Rosa. The structural characters of the ovipositor of this species facilitating laying eggs into such fruits are discussed. The ovipositor musculature and functioning in Carpomya are very similar to those in Ceratitis and Bactrocera; species of all these genera lay eggs in fruits. In the details of the ovipositor musculature, Carpomya is more similar to Ceratitis than to Bactrocera. The genera Carpomya, Ceratitis and Bactrocera are phylogenetically close but Ceratitis and Bactrocera belong to sister tribes within one subfamily, Dacinae. Thus, morpho-functional similarity of the ovipositor structure may not only reflect close relationship but also result from common adaptive transformations.     

The ovipositor musculature of Bactrocera depressa Shiraki (Diptera, Tephritidae)

Structure of the ovipositor sclerites and musculature was studied in the tephritid fly Bactrocera depressa Shiraki, whose larvae develop in the fruits of Cucurbita moschata and other Cucurbitaceae. The functioning and adaptations of the ovipositor of this species to laying eggs into soft fruits are discussed with respect to their differences from the ovipositor characteristics in species with different host specializations, e.g., Ceratitis capitata (Wiedemann) (Hanna, 1938), Lenitovena trigona Matsumura (Ovtshinnikova, 2008), Urophora affinis Frauenfeld and U. quadrifasciata (Meigen) (Berube and Zacharuk, 1983) and Campiglossa plantaginis (Haliday, 1833) (Ovtshinnikova, 2010). The ovipositor musculature of Bactrocera depressa is very similar to that of Ceratitis capitata, a species which lays eggs in soft tissues of various fruits. The sclerites and musculature of the ovipositor, especially the basal part of syntergosternite VII, in the fruit infesting Bactrocera (also in Ceratitis) is intermediate between that in Lenitovena laying eggs in the (soft) decaying wood, and Urophora and Campiglossa that deposit eggs in the hard flowerheads of asteraceans. Species of the latter genera have still more complicated structure of the basal part of syntergosternite VII enabling a more intense protruding of the ovipositor and its more closely controlled operating during oviposition in the harder plant tissues. The discussed morpho-functional rearrangements proceed within species of a single lineage of the modern classification (Korneyev, 1999) from the basal Lenitovena to the terminal Urophora and Campiglossa, Bactrocera occupying an intermediate position in this row. In this way, the technology changes (from a muscul lever to a pump) resulting in an increase of the ovipositor mobility during piercing of the substrate.     

Musculature of the Ovipositor of <Emphasis Type="Italic">Timia erythrocephala</Emphasis> Wiedemann, 1824 (Diptera,Ulidiidae)

The structure of the ovipositor sclerites and musculature was investigated in Timia erythrocephala Wiedemann, 1824 (Ulidiidae), whose larvae are saprophagous. The ovipositor musculature of this species is compared with that of the previously investigated Tephritidae. Based on morphological analysis, characters common to Ulidiidae and primitive Tephritidae and supposedly plesiomorphic for Tephritidae are revealed. Abdominal segment VII in Timia is compact and consists of tergite and stemite VII fused into a syntergostemite with a visible suture. The preserved suture between the stemite and tergite of segment VII should be regarded as an intermediate state in the process of syntergostemite formation in tephritoid flies, which was completed already in Lenitovena.     

Anatomy and mechanics of the telescopic ovipositor system of Scelio latreille (Hymenoptera: Scelionidae) and related genera

The telescopic ovipositor system possessed by the genus Scelio Latreille (Hymentoptera : Scelionidae) and related genera is described in detail for the first time, and found to be anatomically and mechanically unique amongst the parasitic Hymenoptera. Its basic plan is similar to the Ceratobaeus-type ovipositor system described previously for other scelionids, in that the ovipositor is invaginated entirely into the body cavity when at rest and attached to the terminal metasomal segment only by a lightly sclerotized collapsible membranous tube. There are, however, significant anatomical and mechanical modifications that permit the Scelio-type system greater extendability. Telescopic extension of multiple sections of greatly elongated intersegmental membrane, operated by changes in hydrostatic pressure, allows the entire ovipositor system, including the terminal metasomal tergite (T7 + T8), to be exserted from the body cavity during oviposition, thus extending the range of the ovipositor by as much as 3.5 times its length. These changes are accompanied by the incoporation of the lateral apodemes into the wall of the most distal segment of telescopic tube, and the loss of their associated musculature. Similar to other scelionids, orientation of the ovipositor in Scelio is controlled by contraction of muscles connecting the proximal head of the ovipositor with the fused terminal metasomal tergite (T7 + T8). A model for the mechanics of extension and retraction of the Scelio-type ovipositor system is proposed, and is supported by anatomical evidence, behavioural observations and direct manipulation of the system. The evolution of this system in relation to the exploitation of a particular host group, the eggs of Orthoptera, and its significance in resolving phylogenetic problems within the Scelionidae, are also discussed.     

Pigmy moths (Lepidoptera, Nepticulidae) from the Middle Volga Basin

Fifty-seven species of the family Nepticulidae were found in the Middle Volga Basin. The family includes four genera: Bohemannia Stainton, 1859 (1 species), Ectoedemia Busck, 1907 (15), Stigmella Schrank, 1802 (38), and Trifurcula Zeller, 1848 (3 species). The mines made by pigmy moth larvae are very specific in the study area. Larvae of the largest genus, Stigmella, make typical serpentine leaf mines. Larvae of the genera Bohemannia and Ectoedemia make blotchy mines; some larvae develop in fruits and tree bark. Most species of the genera Bohemannia, Ectoedemia, and Stigmella were recorded by their mines. At the larval stage, representatives of Trifurcula feed on stems of legumes. The pupae develop in the soil. Most species in the family Nepticulidae are leaf miners, mainly associated with Rosaceae (34.7%), Salicaceae (15.8%), Fagaceae (12.3%), and Betulaceae (12.3%). All the species investigated do not act as pests of forage plants in the area studied.     

The ovipositor morphology in the members of the family Scathophagidae (Diptera) with reference to their biology

Relation of the ovipositor structure and the egg-laying habit with the larval biology in the family Scathophagidae was investigated. The ovipositor was examined and illustrated for 29 species including Phrosia albilabris (F.), Cordilura umbrosa (Lw.), Spathephilus breviventris (Lw.), Scathophaga stercoraria (L.), Chaetosa punctipes (Mg.), Pogonota barbata (Ztt.), Cochliarium cuneiventris (Ztt.), and Delina nigrita (Fll.). Species with different bionomics and different larval feeding tissues, i.e., different substrates for oviposition, were distinguished. Comparative morphological analysis of the author’s and literature data on 29 species belonging to 15 genera has shown the constancy of the ovipositor structure within the genera examined. Two principal types of the ovipositor structure were distinguished: (a) the elongate ovipositor with the distal position of the terminal part of the abdomen (the proctiger), this type is characteristic of phytophagous species (the delinine type); and (b) the shortened ovipositor with the proctiger shifted dorsally (the scathophagine type), this type is typical of predatory and saprophagous species. The delinine type of the ovipositor is adapted to inserting eggs into hard plant tissues. Changes in the feeding habits of the larvae were found to be associated with changes in the ovipositor structure. The dependence of the ovipositor structure on the egg-laying habit and on the larval biology enables hypothesizing biological features of those species for which the relevant data are not available yet. The morphogenesis of the ovipositor is speculated based on the data on its structure.     

Musculature of the ovipositor of <Emphasis Type="Italic">Oxyna parietina</Emphasis> (Linnaeus, 1758) (Diptera,Tephritidae) and its relation to the larval habits

The structure of the ovipositor sclerites and muscles was studied in the tephritid fly Oxyna parietina (Linnaeus, 1758) inducing galls on the stems of Artemisia vulgaris. Adaptations of the ovipositor structure associated with changes of the oviposition substrate due to new larval habits are analyzed. The ovipositor muscles of Oxyna parietina are compared to those of Campiglossa. The genera Oxyna Robineau-Desvoidy, 1830 and Campiglossa Rondani, 1870 belong to the same genus-group of Tephritini, but differ in their host associations and oviposition habits: the larvae of Campiglossa plantaginis (Haliday, 1833) develop in the capitula of Aster tripolium and other asters (Asteraceae). The ovipositor sclerites and muscles are similar in the two genera. Progression of hemolymph into the membranous sheath of the ovipositor and protraction of the aculeus are necessary conditions for laying eggs into the capitula of Asteraceae (Campiglossa plantaginis), as well as for insertion of eggs into the axils of Artemisia leaves (Oxyna parietina). The most significant characters of Oxyna and Campiglossa include the presence of the hyaline apodeme and associated muscles MVM 6 and MVM 7 (instead of the median apodeme in Urophora). Thus, in other tephritid flies similarity in the structure of the ovipositor muscles may reflect similar morphofunctional adaptations to laying eggs into similar substrates, but similarity of Campiglossa and Oxyna in the structure of the ovipositor muscles is due to their close relations.     

Functional anatomy of the ovipositor clip in the parasitoid leptopilina heterotoma (Thompson) (Hymenoptera : Eucoilidae), a structure to grip escaping host larvae

Observations on the host attack behaviour of the parasitoid Leptopilina heterotoma (Hymenoptera : Eucoilidae) led to the supposition that this wasp should possess a structure on its ovipositor by which it can hold a host larvae in a fixed position until the larva is paralyzed. The ovipositor was studied by light, scanning and transmission electron microscopy and appeared to have a clip with teeth on the unpaired valve of the ovipositor, about 50 μm from the tip. Based on the structure of this ovipositor clip and the oviposition behaviour, the functioning of the ovipositor clip is discussed.     

Spider silk felting—functional morphology of the ovipositor tip of Clistopyga sp. (Ichneumonidae) reveals a novel use of the hymenopteran ovipositor

Apical serrations of the hymenopteran ovipositor have been widely postulated to originally constitute adaptations for cutting through hard substrates. Simplifications of the ovipositor tip have occurred in several ichneumonid wasp genera associated with spiders. Despite such reduction in Clistopyga (Hymenoptera, Ichneumonidae), the ovipositor still possesses some apical serrations. Through the first detailed study, we believe, on the behaviour of an ovipositing Clistopyga species, we show that it can alter its ovipositor for different purposes and that the primary function of the apical serrations is clinging to its spider host as the spider attempts to escape. Intriguingly, we also discover a hitherto undocumented adaptation for the hymenopteran ovipositor. The female wasp seals openings in the silken spider nest by using its ovipositor on the silk in a highly sophisticated way that is comparable to how humans entangle wool by needle felting. By studying the ovipositor morphology through a scanning electron microscope, we elucidate how this works, and we hypothesize that by closing the nest the female wasp protects its developing kin.     

Geometrid moths of the genus Eupithecia Curtis, 1825 (Lepidoptera,Geometridae): Prerequisites and characteristic features of high species diversity

The prerequisites and characteristics of high species diversity of the genus Eupithecia (Geometridae), the largest in the order Lepidoptera, are discussed. The surge of speciation in this genus was facilitated by the evolutionarily advanced morpho-functional characters of the adults and larvae, as well as by the plasticity of a number of morphological structures, especially those of the genitalia in both sexes, in the ancestral Eupithecia. The wide range of flowering plants has served as a basis for close associations of the larvae predominantly with generative plant organs and for broad adaptive radiation of Eupithecia into a variety of ecological niches in the areas lacking their competitors from other genera of the tribe Eupitheciini. Transition of the larvae of some Eupithecia to facultative or obligate ambush predation was facilitated by adaptation of the antho- and carpophagous species to high-calorie food rich in proteins, lipids, and carbohydrates. Based on the distribution of the genus Eupithecia and its most morphologically primitive species-groups, its East Asian origin is hypothesized. It is noted that many of the largest genera of animals and plants include relatively small-sized species.     

Sense organs on the ovipositor of Macrocentrus cingulum Brischke (Hymenoptera: Braconidae): their probable role in stinging,oviposition and host selection process

Parasitoid wasps from the insect order Hymenoptera can be deployed successfully as biological control agents for a number of pests, and have previously been introduced for the control of corn pest insect species from the Lepidopteran genus Ostrinia. Organs on the ovipositor of parasitoid wasps have mechanical and tactile senses that coordinate the complex movements of egg laying, and the ovipositor of Hymenopteran insects have evolved associated venom glands as part of their stinging defense. The ovipositor of parasitic wasps has evolved an additional function as a piercing organ that is required for the deposition of eggs within suitable host larvae. The morphology and ultrastructure of sense organs on the ovipositor and sheath of Macrocentrus cingulum Brischke (Hymenoptera: Braconidae) are described using scanning and transmission electron microscopy. Three types of sensilla trichodea were shown to be abundant on the outer sheath of the ovipositor, with types II and III being most distal, and the inner surface of the ovipositor covered with microtrichia, more densely near the apex. Sensilla coeloconica are distributed on both ventral and dorsal valves, while campaniform sensilla and secretory pores are only located on the dorsal valve. The olistheter-like interlocking mechanism, as well as the morphology of the ventral and dorsal valve tips and the ventral valve seal may be important in stinging, oviposition and in the host selection process.     

Peculiarities of muscles and skeleton structure in <Emphasis Type="Italic">Campiglossa plantaginis</Emphasis> (Haliday, 1833) ovipositor (Diptera,Terthritidae) in association with larval mode of life

Sclerites and muscles of the ovipositor in the tephritid fly Campiglossa plantaginis (Haliday, 1833), with phytophagous larvae developing in inflorescences of Aster tripolium L. and other Asteraceae, were examined. Structural adaptations of the ovipositor, providing oviposition into different substrates, are discussed in relation to different trophic specialization of larvae.     

Revision of the tribe Colpopterini Gnezdilov, 2003 (Homoptera, Fulgoroidea: Nogodinidae)

The tribe Colpopterini Gnezdilov, 2003 is revised, upgraded to the subfamily Colpopterinae, stat. n., and transferred from the family Issidae Spinola to Nogodinidae Melichar. The tribe Colpopterini is recorded from the Afrotropical Region for the first time—Bumerangum deckerti gen. et sp. n. is described from Southern Africa. The tribe Colpopterini s. str. comprises 6 genera: Bumerangum gen. n., Caudibeccus Gnezdilov et O’Brien, 2008, Colpoptera Burmeister, 1835, Jamaha Gnezdilov et O’Brien, 2008, Neocolpoptera Dozier, 1931, and Ugoa Fennah, 1945. The genera Cheiloceps Uhler, 1895, Tempsa Stål, 1866, Eupilis Walker, 1857, and Gabaloeca Walker, 1870 are transferred to the tribe Issini Spinola of the family Issidae. Issus longulus Lethierry, 1890 is transferred to the genus Colpoptera Burmeister. A key to the genera and a list of the species of the tribe Colpopterini are given. Morphological data confirming independent evolution of similar ovipositor types in the families Issidae and Nogodinidae are provided. The term “styletization” is suggested for describing the transformation of the ovipositor from a rounded to an elongate type.     

Phylogeny of embiopterans (Insecta)

A cladistic analysis of embiopterans, based on 157 species (representing 70% of the known genera) and 186 morphological characters, is presented, as well as a molecular analysis for 22 taxa using genes encoding 16S, 18S and 28S rDNA and COI. Species of all known families are included, except Andesembiidae Ross (specimens of which are in a private collection). The evidence presented supports the monophyly of four of the families (Australembiidae, Oligotomidae, Teratembiidae, and Anisembiidae). Notoligotomidae is paraphyletic and included within the Afro‐neotropical family Archembiidae (which is also paraphyletic). The genera Embia, Cleomia, Macrembia, and Dihybocercus (Embiidae) form, together with Australembiidae, a group strongly supported by morphology; the position of the remaining genera of Embiidae has two quite different resolutions. Almost 80% of the genera of Anisembiidae recently described appear as either paraphyletic or polyphyletic. Contrary to the opinion of other specialists, the major groups as well as the monophyly of some families are supported by features which have been ignored in classical approaches to the systematics of Embioptera, such as the ovipositor and cephalic and leg structures, characters with an almost perfect fit.     

Phylogeny of the platygastroid wasps (Hymenoptera) based on sequences from the 18S rRNA, 28S rRNA and cytochrome oxidase I genes: implications for the evolution of the ovipositor system and host relationships

The Platygastroidea are a diverse group of mostly small to tiny wasps, the common biology for which is endoparasitism of insect and spider eggs. No analytically‐based phylogeny exists for the superfamily, and the current suprageneric classification is flawed in part because of its reliance on homoplasious and pleisiomorphic morphological characters. To determine platygastroid relationships as a basis for investigating host and ovipositor evolution, phylogenies of > 70 in‐group species (representing 55 genera) were reconstructed by parsimony and Bayesian methods using three molecular markers; the mitochondrial cytochrome oxidase I and the nuclear genes 28S and 18S rRNA. The results strongly support the monophyly of the superfamily and one of the two families, Platygastridae, but the Scelionidae are most likely polyphyletic. However, within the Scelionidae, there is a well supported ‘main scelionid clade’ that contains the majority of genera assigned to the family. At the subfamilial level, both putative subfamilies of Platygastridae, the Platygastrinae, and Sceliotrachelinae, are likely to be polyphyletic. Within the Scelionidae, both the Teleasinae and Telenominae are monophyletic, but the Scelioninae is clearly not so. The current tribal classification for the Scelionidae is in need of major reassessment because no tribes, with the exception of the Scelionini s.s., were found to be monophyletic. Further illustrating the problems associated with the current classification is the nonmonophyly of a number of genera, namely Opisthacantha Caloteleia, Telenomus, Trimorus, Teleas and Idris. Analysis of ovipositor evolution in the superfamily revealed that the Ceratobaeus‐type ovipositor system is ancestral; however, this trait was lost prior to the evolution of the main scelionid clade, for which the Scelio‐type ovipositor system is ancestral and defines a mostly paraphyletic assemblage. Ancestral state analysis indicates that the Ceratobaeus‐type ovipositor was subsequently re‐evolved in the main scelionid clade, representing a possible contradiction of Dollo’s law. Previously, the tribal placement has been used to predict the host associations of genera for which host data were unavailable. However, the fact that most tribes are not monophyletic throws into doubt any such speculation based on the current classification. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 653–669.     

Evaluation of the DNA Barcodes in Dendrobium (Orchidaceae) from Mainland Asia

DNA barcoding has been proposed to be one of the most promising tools for accurate and rapid identification of taxa. However, few publications have evaluated the efficiency of DNA barcoding for the large genera of flowering plants. Dendrobium, one of the largest genera of flowering plants, contains many species that are important in horticulture, medicine and biodiversity conservation. Besides, Dendrobium is a notoriously difficult group to identify. DNA barcoding was expected to be a supplementary means for species identification, conservation and future studies in Dendrobium. We assessed the power of 11 candidate barcodes on the basis of 1,698 accessions of 184 Dendrobium species obtained primarily from mainland Asia. Our results indicated that five single barcodes, i.e., ITS, ITS2, matK, rbcL and trnH-psbA, can be easily amplified and sequenced with the currently established primers. Four barcodes, ITS, ITS2, ITS+matK, and ITS2+matK, have distinct barcoding gaps. ITS+matK was the optimal barcode based on all evaluation methods. Furthermore, the efficiency of ITS+matK was verified in four other large genera including Ficus, Lysimachia, Paphiopedilum, and Pedicularis in this study. Therefore, we tentatively recommend the combination of ITS+matK as a core DNA barcode for large flowering plant genera.     

Species specific responses of common grassland plants to a generalist root herbivore (Agriotes spp. larvae)

Insect root herbivores can alter plant community structure by affecting the competitive ability of single plants. Our study aims at increasing knowledge on the impact of click beetle larvae (Elateridae, genus Agriotes) on grassland plant communities, by determining biomass responses as well as responses of the arbuscular mycorrhizal fungi (AMF) symbiosis to this widespread generalist root herbivore for eight common grassland plant species (Festuca rubra, Holcus lanatus, Poa pratensis, Achillea millefolium, Plantago lanceolata, Veronica arvensis, Medicago lupulina, Trifolium repens), belonging to three functional groups (grasses, herbs, legumes).The presence of larvae in the rhizosphere of individual plants had an overall negative effect on root biomass, which varied in degree between plant species, with more root biomass being removed from larger root systems. The effect of larvae on shoot biomass, total plant biomass and shoot/root ratio also differed in strength between plant species. Relative changes in root and total plant biomass ranged from a 71% and 55% loss, respectively, in V. arvensis to an 11% and 1% increase in T. repens, but were not related to root- or plant size. Root colonization by AMF and the length of extraradical AMF hyphae were not affected by larvae. The plant's functional group did not determine the response of the plant to feeding by larvae. Growth of larvae was positively correlated with root biomass, but did not depend on plant species or group identity.The results confirm the generalist nature of Agriotes spp. larvae, which depend on sufficient root quantity and are likely to feed most on those plant species whose roots are most abundant in their habitat. Their effect on the plant community may be generated through the ability of the respective plant species to cope with the herbivore damage, with tolerances being plant species rather than plant group specific.     

Evaluation of <Emphasis Type="Italic">Nastus fausti</Emphasis> Reitter (Coleoptera: Curculionidae: Entiminae: Nastini) for biological control of invasive giant hogweeds (<Emphasis Type="Italic">Heracleum</Emphasis> spp.)

The weevil Nastus fausti Reitter (Coleoptera, Curculionidae) was evaluated for its potential in the biological control of invasive giant hogweeds (Heracleum spp.). Quantitative sampling suggested that at a high population density (more that 3–4 mature larvae per plant) damage by N. fausti larvae could have some negative impact on the above-ground part of the plant. However, no-choice laboratory tests showed that N. fausti females were able to feed on a number of Apiaceae genera, including such important cultivated crops as carrot, parsnip, and celeriac. Feeding on these plants did not cause any significant decrease in female survival or fecundity. Moreover, at least part of N. fausti larvae may feed and develop on roots of these plants, and the rate of their growth and development does not differ significantly from that in larvae fed on roots of H. mantegazzianum. N. fausti adult and larval feeding on Angelica purpurascens, representative of related genus of the same tribe, was recorded under natural conditions, too. In combination, these data suggest that N. fausti is an oligophagous species connected with plants from at least several genera of Apiaceae and thus it cannot be considered a potential agent for biological control of invasive Heracleum species.     

Muscle development in the abdominal region of Larval Hylidae (Amphibia: Anura)

Larval muscle development in the abdominal region of five species of hylid frogs (Scinax nasicum, S. fuscovarium, Hyla andina, Phyllomedusa boliviana, Gastrotheca gracilis) was studied using differential staining techniques. These five species represent three major hylid subfamilies. The development of the main abdominal muscles, the rectus abdominis, the two lateral muscles (obliquus externus and transversus), and the lateral pectoralis abdominalis is described. The number of myotomes of the rectus abdominis varies between five and six, and the abdominal muscles associated with the rectus abdominis (obliquus externus, pectoralis abdominalis, and rectus cervicis) vary interspecifically in time of appearance and configuration. The presence of gaps in the configuration of the rectus abdominis has been related to the lotic habits of the larvae. However, our observations indicate the presence of such gaps in larvae that inhabit lentic environments as well. These results suggest that the presence of these gaps is unrelated to larval habitat. There are relatively small differences in muscle morphology among these closely related species, which apparently cannot be explained by morphological adaptations related to their ecology. In the species studied, the number of elements that form the abdominal musculature in larvae is equal to that observed in adults. Likewise, the general morphology of the muscles is ontogenetically conserved. This suggests that both the axial skeleton and musculature are more ontogenetically conserved in relation to the substantial changes that are observed in the skull and head muscles of developing anurans. J. Morphol. 241:275–282, 1999. © 1999 Wiley‐Liss, Inc.