Connecting biology, optics and nanomechanical properties in micro-wasps.

Coloration in insects provides a fruitful opportunity for interdisciplinary research involving both physics and biology, and for a better understanding of the design principles of biological structures. In this research we used nanometric and micrometric analyses to investigate the morphological and mechanical properties of the black-orange-black (BOB) color pattern in scelionid wasps, which has never been studied. The primary objective of the present investigation was to explore the structural and mechanical differences in the mesoscutum of four species: Baryconus with an orange mesosoma (i.e. BOB pattern), all black Baryconus, Scelio with an orange mesosoma (i.e. BOB pattern), and all black Scelio. The most outstanding findings include the absence of multilayer structures that generate structural color, a pigment concentrated in the upper surface of the epicuticle, and surprising differences between the four species. Three of the four species showed an accordion-like structure in the furrow (notaulus), whereas the adjacent mesoscutum was different in each species. Moreover, the normalized color component spectra for blue, green and red colors of the black mesoscutum of each genus showed the same spectral dependence while the orange color manifested small changes in the dominant wavelength, resulting in slightly different orange tones.

Sperm morphology of Elasmus polistis Burks, 1971 (Hymenoptera: Chalcidoidea: Eulophidae).

The sperm morphology of the parasitoid Elasmus polistis (Eulophidae) has been investigated with light and transmission electron microscopy. The sperm were filiform and spiraled, with 165.6 (± 4.6) µm in length, and showed a distinctive head, formed by a one-layered small acrosome and a nucleus, and a flagellar region. An extracellular sheath from which many long filaments radiated out covered the acrosome and part of the nucleus. The spiral nucleus, with 24.1 (± 1.3) µm in length, was filled with homogeneously compact chromatin. In the nucleus-flagellum transition, the centriole adjunct extended posteriorly from the nuclear base in a spiral around the basal body, which has two central microtubules, and axoneme for approximately 1.1 µm. The two mitochondrial derivatives began roughly at the same level and at the base of the centriole adjunct. In cross-section, they were symmetrical, with a slightly oval shape and a smaller diameter in comparison to the axoneme. The latter, also spiraled, consisted of 9 + 9 + 2 microtubules that was formed from the basal body situated just below and aligned with the nucleus. The E. polistis sperm showed the same basic structures and morphological characteristics as observed in other Chalcidoidea. However, it was possible to distinguish the sperm of this species from those of other Eulophidae by (i) the long length of the centriole adjunct on the flagellum, and (ii) the presence of two central microtubules within the basal body. The sperm characteristics suggest that Eulophidae is closely related to Trichogrammatidae and both families are more similar to Eurytomidae, Pteromalidae, and Torymidae than Agaonidae.

Sperm morphology of Muscidifurax uniraptor (Hymenoptera: Chalcidoidea: Pteromalidae).

Sperm morphology of the parasitoid Muscidifurax uniraptor was investigated under light and transmission electron microscopy. M. uniraptor sperm are filiform, spiraled, approximately 150 µm in length, with a distinctive head, hooded by an extracellular sheath and a flagellum. This extracellular layer, from which many filaments radiate, measures approximately 90 nm in thickness and covers a small acrosome and the anterior nuclear region. The acrosome is composed of an acrosomal vesicle and a perforatorium with its base inserted in the nuclear tip. The nucleus is filled with homogeneously compacted chromatin. The centriolar adjunct extends towards the anterior portion in a spiral around the nucleus for 3.5 µm in length. The two mitochondrial derivatives begin exactly at the centriole adjunct base and, in cross-section, have a circular shape with equal areas that are smaller than the axoneme diameter. It is coiled, with 9 + 9 + 2 microtubules and begins from the centriole, just below the nuclear base. The axoneme is connected to the mitochondrial derivatives by two small irregularly shaped masses. Between the derivatives and the axoneme, the 'center-flagellar material' is observed. Overall, these characteristics are recognized in other Chalcidoidea, especially in the eurytomids, but together they form a set of species-specific data.

Redescription of Campoletis sonorensis (Cameron, 1886) (Hymenoptera, Ichneumonidae, Campopleginae), parasitoid of Spodoptera frugiperda (J. E. Smith, 1797) (Lepidoptera, Noctuidae) in Brazil.

The fall armyworm Spodoptera frugiperda (Lepidoptera; Noctuidae) is a voracious pest of numerous crops of economic importance throughout the New World. In Brazil, its larvae are attacked by several species of parasitoid wasps, making them potential candidate as biological control agents against this pest. A survey of the parasitoid fauna on S. frugiperda in maize crops throughout Brazil reveals two species of Campoletis, which are morphologicaly very similar species. In this paper we combine these data with pictures from the type material of C. sonorensis and C. flavicincta, as well as their descriptions to provide a redescription to Campoletis sonorensis (Cameron, 1886) using for this both morphological characters and DNA Barcoding (Hebert et al., 2003) information, in an attempt to help with the correct identification of the taxa to improve biological control studies.

Redescription of Campoletis sonorensis (Cameron, 1886) (Hymenoptera, Ichneumonidae, Campopleginae), parasitoid of Spodoptera frugiperda (J. E. Smith, 1797) (Lepidoptera, Noctuidae) in Brazil / Redescrição de Campoletis sonorensis (Cameron, 1886) (Hymenoptera, Ichneumonidae, Campopleginae), parasitoide de Spodoptera frugiperda (J. E. Smith, 1797) (Lepidoptera, Noctuidae) no Brasil

The fall armyworm Spodoptera frugiperda (Lepidoptera; Noctuidae) is a voracious pest of numerous crops of economic importance throughout the New World. In Brazil, its larvae are attacked by several species of parasitoid wasps, making them potential candidate as biological control agents against this pest. A survey of the parasitoid fauna on S. frugiperda in maize crops throughout Brazil reveals two species of Campoletis, which are morphologicaly very similar species. In this paper we combine these data with pictures from the type material of C. sonorensis and C. flavicincta, as well as their descriptions to provide a redescription to Campoletis sonorensis (Cameron, 1886) using for this both morphological characters and DNA Barcoding (Hebert et al., 2003) information, in an attempt to help with the correct identification of the taxa to improve biological control studies.(AU)

Spodoptera frugiperda é uma praga voraz de diversas culturas de importância econômica no Novo Mundo. No Brasil, suas larvas podem ser atacadas por diversas espécies de vespas parasitóides que são candidatos a agentes de controle biológico contra essa praga. Pesquisando os parasitóides da fauna de Spodoptera frugiperda em cultivos de milho no Brasil foram encontradas duas espécies do gênero Campoletis, muito similares morfologicamente. Este trabalho apresenta uma redescrição para Campoletis sonorensis usando caracteres morfológicos e DNA Barcoding (Hebert et al., 2003) com o objetivo de evitar erros de identificação desse grupo, aprimorando estudos de controle biológico.(AU)

Redescription of Campoletis sonorensis (Cameron, 1886) (Hymenoptera, Ichneumonidae, Campopleginae), parasitoid of Spodoptera frugiperda (J. E. Smith, 1797) (Lepidoptera, Noctuidae) in Brazil / Redescrição de Campoletis sonorensis (Cameron, 1886) (Hymenoptera, Ichneumonidae, Campopleginae), parasitoide de Spodoptera frugiperda (J. E. Smith, 1797) (Lepidoptera, Noctuidae) no Brasil

Abstract The fall armyworm Spodoptera frugiperda (Lepidoptera; Noctuidae) is a voracious pest of numerous crops of economic importance throughout the New World. In Brazil, its larvae are attacked by several species of parasitoid wasps, making them potential candidate as biological control agents against this pest. A survey of the parasitoid fauna on S. frugiperda in maize crops throughout Brazil reveals two species of Campoletis, which are morphologicaly very similar species. In this paper we combine these data with pictures from the type material of C. sonorensis and C. flavicincta, as well as their descriptions to provide a redescription to Campoletis sonorensis (Cameron, 1886) using for this both morphological characters and DNA Barcoding (Hebert et al., 2003) information, in an attempt to help with the correct identification of the taxa to improve biological control studies.

Resumo Spodoptera frugiperda é uma praga voraz de diversas culturas de importância econômica no Novo Mundo. No Brasil, suas larvas podem ser atacadas por diversas espécies de vespas parasitóides que são candidatos a agentes de controle biológico contra essa praga. Pesquisando os parasitóides da fauna de Spodoptera frugiperda em cultivos de milho no Brasil foram encontradas duas espécies do gênero Campoletis, muito similares morfologicamente. Este trabalho apresenta uma redescrição para Campoletis sonorensis usando caracteres morfológicos e DNA Barcoding (Hebert et al., 2003) com o objetivo de evitar erros de identificação desse grupo, aprimorando estudos de controle biológico.

Uncovering head gland diversity in neotropical Polistinae wasps (Hymenoptera, Vespidae): comparative analysis and description of new glands.

Exocrine glands are involved in several wasp colony activities; however, the number of known glands in the Vespidae is rather low when compared to other social insect groups. The aim of this study is to survey the head of Neotropical social wasps and to provide a detailed comparative study of the glands found in the Polistinae. A total of 33 species distributed over 13 genera were studied with serial histological sections of the head, excluding the labiomaxillary complex. Additionally, the exoskeleton was explored using scanning electron microscopy looking for associated modifications. A total of eleven exocrine glands were observed, five are structures recorded for the first time for the Hymenoptera, three are new records for the Polistinae and three are previously known organs. The glands studied are: ocellar gland I, ocellar gland II, periocular gland, subantennal gland, hypopharyngeal gland, clypeal gland, posterobasal genal gland, ectal mandibular gland, mesal mandibular gland, intramandibular gland I, and intramandibular gland II. The widespread distribution of most of these glands suggests an origin prior to the evolution of the Polistinae. Our results highlight the importance of detailed morphological studies to unveil the significance of chemical communication in one of the most characteristic groups of social animals.

A mandible arresting system in neotropical social wasps (Vespidae; Polistinae): structural diversity within homogeneous functionality.

Microtrichia are epidermal protuberances that may serve as temporary adhesive devices. Several insects possess these structures; however, they have not previously been reported in social wasps. With scanning electron microscopy, we characterize the shape and abundance of microtrichia in ten species of social wasps (Vespidae: Polistinae) and three species of related taxa (Vespidae: Eumeninae, Pompilidae, and Scoliidae). Semi-thin sections of the head of Leipomeles spilogastra and Apoica albimacula were also studied. We found microtrichia on a thin, flexible membrane connected to the mandible in all the Vespidae specimens. The flexible membrane can be divided into three regions: the basal region that covers the mandibular mesial emargination, the medial region located around the height of the mandibular condyles, and the distal region that appears anterior to the apodeme folding. Basal and distal regions of the membrane are extensively covered by microtrichia while the medial region has either less microtrichia or is entirely devoid of them. The shape and density of the microtrichia differed between species, and these traits are unrelated with nest material construction or phylogenetic closeness. We propose that the microtrichial membrane described is a passive mechanism to keep the wasps' mandibles retracted through a mechanical interlocking system. It is possible that this energy-saving mechanism is present in other mandibulate insects.

Revision of Apocharips Fergusson (Hymenoptera: Figitidae: Charipinae) with description of three new species from Colombia.

The genus Apocharips is here revised. Three previously described species are considered as valid: Apocharips angelicae Pujade-Villar & Evenhuis, 2002, A. hansoni Menke, 1993, and A. trapezoidea (Hartig, 1841). Apocharips eleaphila (Silvestri, 1915) and A. peraperta (Silvestri, 1915) are synonymyzed with A. trapezoidea. Three new species are here described: Apocharips colombiana Ferrer-Suay & Pujade-Villar n. sp., Apocharips tamanii Paretas-Martinez & Pujade-Villar n. sp. and Apocharips tropicale Ferrer-Suay & Paretas-Martinez n. sp. A key to the six species included in Apocharips is given.

Sperm morphology of the Prorops nasuta (Waterston, 1923) (Hymenoptera: Bethylidae).

In the present study, spermatozoa of the Prorops nasuta (Hymenoptera: Bethylidae) parasitoid were described morphologically. This is the first publication to describe a species belonging to the superfamily Chrysidoidea. Light and transmission electron microscopy were used. The spermatozoa of P. nasuta are linear, with a mean length of 665microm. The acrosome is composed of an acrosomal vesicle and a perforatorium. The nucleus measures approximately 17microm in length and is circular at its cross-section; however, its anterior extremity is oval. The chromatin is electron-dense and compact, although there are clear areas in the posterior peripheral regions. In the nucleus-flagellum transition region, the cross-section of the centriole adjunct is oval, with a pleated border and an E-PTA-positive peripheral region. The axoneme shows a 9+9+2 microtubule arrangement. The microtubules are E-PTA positive and, at the posterior extremity, the accessories are the last to terminate. The diameters and shapes of the two mitochondrial derivatives are almost identical. One begins beside the nuclear base and the other after the centriole adjunct. Posteriorly, they terminate together, immediately before the axoneme. Both have mitochondrial cristae and a region of paracrystalline material; however, the format and arrangement of this material differs from those of all other species previously studied. The paracrystalline material is more strongly E-PTA positive than the cristae region. Accessories bodies are electron-dense and located between the mitochondrial derivatives and the axoneme. In general, P. nasuta spermatozoa are similar to those of the majority of Hymenoptera; however, they have various exclusive characteristics that may be useful for studying the phylogeny and taxonomy of the superfamily Chrysidoidea and of Hymenoptera in general.

Structure and ultrastructure of the spermatozoa of Trypoxylon (Trypargilum) albitarse Fabricius 1804 (Hymenoptera: Apoidea: Crabronidae).

The ultrastructure of the spermatozoa of Trypoxylon (Trypargilum) albitarse is described here for the first time within this genus. Testes and seminal vesicles were dissected and processed for transmission electron microscopy. In the testicular follicles, the spermatids are arranged in a maximum number of 32 for each cyst. The spermatozoa are slender and they measure approximately 150 microm in length. The head is about 17 microm long and is formed by the acrosome and the nucleus. The flagellum consists in an axoneme, two mitochondrial derivatives, two accessory bodies and, at the nucleus-flagellum transition, a symmetric centriolar adjunct. The axoneme presents the typical 9+9+2 microtubule pattern. In the terminal region, the central microtubules and nine doublets finish first, followed by the accessory microtubules. Both mitochondrial derivatives begin together and are inserted in the base of the centriolar adjunct. Along the middle region, the larger derivative has almost twice the area of the smaller one and includes a discrete paracrystalline region. At the tip, the smaller derivative ends before the larger one and both before the axoneme. The characters derived from the ultrastructure of the spermatozoa of T. albitarse show synapomorphies shared with the Apoidea and present characters that are probably apomorphic for the Crabroninae subfamily.

Morphology of male reproductive system in three species of Trypoxylon (Trypargilum) Richards (Hymenoptera: Crabronidae).

Variations in the adult male reproductive system among different groups of Hymenoptera offer characteristics that help studies on behavior and phylogenetics. The objective of this study was to describe the adult male reproductive system of three Trypoxylon (Trypargilum) species. For that, tissues were disseced, fixed in 2.5% glutaraldehyde in 0.1 M sodium cacodylate buffer, pH 7.2 and postfixed in 1% osmium tetroxide. The material was dehydratated and embedded for light and electron transmission microscopes. The species have similar reproductive systems, which are formed by a pair of testes, each one with three fusiforme follicles, from which emerges an efferent duct that later joins forming a deferent duct. The deferent duct opens into an ejaculatory duct. The first half of the deferent duct is enlarged and differentiated in a region specialized in sperm storage, the seminal vesicle. The accessory gland flows in the post-vesicular region of the deferent duct. The testes and vesicles are both covered with a conjunctive capsule. Sexually mature individuals have all spermatogenesis stages in their follicles. Sperms are released from testes in bundles which are disorganized inside seminal vesicles.

Morphology of male reproductive system in three species of Trypoxylon (Trypargilum) Richards (Hymenoptera: Crabronidae) / Morfologia do sistema reprodutor masculino em três espécies de Trypoxylon (Trypargilum) Richards (Hymenoptera: Crabronidae)

Variations in the adult male reproductive system among different groups of Hymenoptera offer characteristics that help studies on behavior and phylogenetics. The objective of this study was to describe the adult male reproductive system of three Trypoxylon (Trypargilum) species. For that, tissues were disseced, fixed in 2.5 percent glutaraldehyde in 0.1 M sodium cacodylate buffer, pH 7.2 and postfixed in 1 percent osmium tetroxide. The material was dehydratated and embedded for light and electron transmission microscopes. The species have similar reproductive systems, which are formed by a pair of testes, each one with three fusiforme follicles, from which emerges an efferent duct that later joins forming a deferent duct. The deferent duct opens into an ejaculatory duct. The first half of the deferent duct is enlarged and differentiated in a region specialized in sperm storage, the seminal vesicle. The accessory gland flows in the post-vesicular region of the deferent duct. The testes and vesicles are both covered with a conjunctive capsule. Sexually mature individuals have all spermatogenesis stages in their follicles. Sperms are released from testes in bundles which are disorganized inside seminal vesicles.

Variações no sistema reprodutor entre os diferentes grupos de Hymenoptera oferecem caracteres que auxiliam nos estudos de comportamento e filogenia. O objetivo deste trabalho foi descrever o sistema reprodutor masculino de três espécies de Trypoxylon (Trypargilum). Para isso, os tecidos foram dissecados, fixados em glutaraldeído 2,5 por cento em tampão cacodilato de sódio 0,1 M, pH 7,2 e pós-fixados em tetróxido de ósmio a 1 por cento. O material foi desidratado e incluído para microscopias de luz e eletrônica de transmissão. As espécies possuem os sistemas reprodutores muito semelhantes, formados por um par de testículos, cada um com três folículos fusiformes, a partir dos quais emerge um ducto eferente que depois se juntam formando o ducto deferente. O ducto deferente termina no ducto ejaculatório. A primeira metade dos ductos deferentes é dilatada e diferenciada em uma região especializada no armazenamento de espermatozóides, a vesícula seminal. A glândula acessória desemboca na região pós-vesicular do ducto deferente. Testículos e vesículas seminais são envoltos por uma única cápsula conjuntiva. Indivíduos maduros sexualmente apresentam todos os estágios da espermatogênese em seus folículos. Os espermatozóides são liberados dos testículos em feixes, os quais estão desorganizados na vesícula seminal.

Revision of the South American species of Neralsia (Hymenoptera: Figitidae) with the description of eight new species

Neralsia is a genus of Figitid hymenopterans present in both the Neartic and the Neotropical regions. In this work, material from several museums (including all types of the South American species of Neralsia) was analyzed with light and electron microscopy. The South American species are studied as a whole, reviewing 26 previously cited species and describing eight new species. A key for their identification is included and the characters to differentiate species here considered are illustrated. Rev. Biol. Trop. 56 (2): 795-828. Epub 2008 June 30.

Neralsia es un género de himenópteros de distribución americana, presente tanto en la región Neártica como en la Neotropical. En este trabajo se estudian, usando microscopia de luz y electrónica, las especies sudamericanas en su conjunto, 26 anteriormente citadas y ocho especies nuevas, que se describen aquí formalmente. Se presenta, además, una clave para su identificación; y se ilustran los caracteres que permiten definir las especies consideradas en este estudio.

Structural and ultrastructural characterization of male reproductive tracts and spermatozoa in fig wasps of the genus Pegoscapus (Hymenoptera, Chalcidoidea).

The three Pegoscapus species present the same internal reproductive tract features comprising testes with a single testicular tubule, seminal vesicles, vasa deferentia, accessory glands and an ejaculatory duct. The seminal vesicle shows two morphologically distinct portions although they do not resemble the separate chambers found in other Chalcidoidea. The anterior portion of the seminal vesicle shows a prominent epithelium and stores the mature spermatozoa, while the posterior region is formed by a thicker muscular sheath that participates on ejaculation. The sexual maturation in Pegoscapus is achieved at emergence, when the testicular degeneration occurs. The spermatozoa of Pegoscapus reveal a basic structure similar to that of other Chalcidoidea. In Pegoscapus sp1. and Pegoscapus sp2. they present the same features, whereas Pegoscapus tonduzi comprises some different characteristics. It measures approximately 160 microm in Pegoscapus sp1. and Pegoscapus sp2., while in P. tonduzi the spermatozoa measure about 360 microm. The extracellular sheath thickness is another difference among the species. While Pegoscapus sp1. and Pegoscapus sp2. show a thick extracellular sheath, in P. tonduzi this sheath is very thin resulting in a large space intervening between the extracellular sheath and the nucleus. Despite these differences, the three species analyzed share some characteristics that allow the establishment of an identity to the spermatozoon of the genus Pegoscapus: the seminal vesicle not divided in chambers; the absence of acrosomal structures in the spermatozoa; the length of the extracellular sheath; the central microtubules being the firsts to terminate in the sequence of microtubular cutoff at the final axonemal portion.

Revision of the South American species of Neralsia (Hymenoptera: Figitidae) with the description of eight new species.

Neralsia is a genus of Figitid hymenopterans present in both the Neartic and the Neotropical regions. In this work, material from several museums (including all types of the South American species of Neralsia) was analyzed with light and electron microscopy. The South American species are studied as a whole, reviewing 26 previously cited species and describing eight new species. A key for their identification is included and the characters to differentiate species here considered are illustrated.

A comparative study of the proventricular structure in corbiculate apinae (Hymenoptera, Apidae).

The present study compares the proventricular structure, analyzed under scanning electronic microscope (SEM), among tribes of corbiculate Apinae. Fifty-one species of stingless bees (Meliponini), one species of honeybee (Apini), three species of bumblebees (Bombini) and seven species of orchid bees (Euglossini), were analyzed as in-group, and one species of sphecid wasp (Sphecidae) and two species of Halictidae bees, as out-groups. The proventricular bulb presents a basic morphology pattern similar to that of other Hymenoptera such as ants and wasps, being a symplesiomorphy for bees. The shape of proventricular folds constitutes a synapomorphy for Meliponini and an autapomorphy for Apini. The shape of hair-like projections of the cuticle that lines the proventriculus is a synapomorphy for Meliponini and Apini. These proventricular data corroborate the monophyly of the tribe Meliponini and the hypothesis that recognizes only one tribe for stingless bees. In addition, Meliponini+Apini constitutes a monophyletic group and Bombini+Euglossini another monophyletic group. The results confirm that internal morphology is a character that can be used in studies of the phylogeny in insects and the use of SEM as a powerful tool in these analyses.

Sensilla on the antenna and ovipositor of the parasitic wasps trichogramma galloi Zucchi and T. pretiosum Riley (Hym., Trichogrammatidae).

The understanding of the stimuli perceived by these parasitoids to accept and exploit a potential host may support the development of artificial hosts for their in vitro rearing. The most common structures involved in the host selection process in parasitoids are the antenna and the ovipositor. Sensilla present on these structures are involved in host recognition and acceptance, and the identification of which kind of stimulus (physical or chemical) is perceived by the sensilla could provide data about the host characteristics that elicit the parasitism behavior. The antenna and ovipositor of the wasps Trichogramma galloi and T. pretiosum were studied to identify their sensillar structure and their possible functions in the host selection process, as a support to the development of an artificial host for the in vitro rearing of these parasitoids. Seven sensilla and one setiform structure were present on the antenna of both parasitoid species, with a mixed, chemo-, or mechanoreception function. The ovipositor has four different sensilla and their role in the host evaluation is suggested.