Comparison of spermatheca morphology between reproductive and non-reproductive females in social wasps

Social wasps show an obvious evolution of the differentiation in behavior and external size between reproductive and non-reproductive females, with no clear differences in the Stenogastrinae, via overlap in the Polistinae, to clear differences in the Vespinae. In this study, we examined the morphological appearance of the spermatheca in representative species of these three subfamilies. The general anatomical organization of the spermatheca comprises a reservoir, a duct and two spermathecal glands, and is in line with its common structure in other social Hymenoptera. All examined wasp species have a spermathecal reservoir with uniform wall thickness, which is similar to the situation in the bees, but differentiates them from the ants. Within the wasps, the shape of the reservoir, the shape of the spermathecal glands and their attachment site to the spermatheca differs among the Stenogastrinae, Polistinae and Vespinae. The reservoir wall is thick in the Polistinae and Vespinae, while in the Stenogastrinae, it varies from thin in Parischnogaster to thick in Eustenogaster, with an intermediate situation in Liostenogaster. In all examined species, we found no differences in the spermathecal development between reproductive and non-reproductive wasps.     

Sociality and the Rate of rDNA Sequence Evolution in Wasps (Vespidae) and Honeybees (Apis)

Sequence data of mitochondrial 16S ribosomal DNA (mt-rDNA) and nuclear 28S ribosomal DNA (nuc-rDNA) were compared in two honeybee species (Apis mellifera and Apis dorsata) and a selection of 22 wasp species (Vespidae) with different levels of sociality. The averge substitution rates in mt-rDNA and nuc-rDNA were almost-equal in solitary species. In species with larger nests, however, the difference between the nuclear and the mitochondrial substitution rate significantly increased. The average substitution ratio, ψ (nucleotide substitutions in mt-rDNA/nucleotide substitutions in nuc-rDNA) was 1.48 ± 0.12 (SE) among the solitary Eumeninae, 3.70 ± 0.15 among five primitive social Stenogastrinae species, 3.24 ± 0.20 among five Polistinae species, 5.76 ± 0.33 among nine highly eusocial Vespinae, and 12.7 in the two Apis species. The high egg-laying rate and the effective population size skew between the sexes may contribute to the rise of the substitution ratio in the highly eusocial species. Drift and bottleneck effects in the mitochondrial DNA pool during speciation events as well as polyandry may further enhance this phenomenon. Received: 12 January 1998 / Accepted: 28 April 1998     

The origin and evolution of social life in the Stenogastrinae (Hymenoptera,Vespidae)

The Stenogastrinae are a subfamily of the Vespidae. The main difference between these and other social wasps (Polistinae and Vespinae) is a jelly-like substance that the Stenogastrinae secrete from the Dufour 's gland and use in many functions of their biology. It is suggested that this substance greatly contributed to the evolution of social life in these wasps by making it possible to nourish the brood with liquid food and store it in the nest, thus favoring also the evolution of the behavioral mechanisms which facilitated interactions between adults. Social organization of the colonies may have been kept at a low level through a basic system of continuous temporary helper replacement, while the evolution of large colonies was restrained, as well as by the poor quality of construction material, low egg-laying capacity and limited production of abdominal substance, imperfect social regulatory mechanisms, and the absence of defensive mechanisms of the colonies against large predators.     

Early lineages of Vespidae (Hymenoptera) in Cretaceous amber

Three wasp (Hymenoptera: Vespidae) fossils in Cretaceous amber (Late Albian) of northern Myanmar are described. Two are new species of the Mesozoic genus Curiosivespa (Rasnitsyn): C. zigrasi sp.n. and C. striata sp.n. The third species, Protovespa haxairei gen.n. et sp.n. , has a combination of features unique among Mesozoic Priorvespinae and the extant subfamilies. These well preserved fossils provide new morphological data for a cladistic analysis of the basal lineages of Vespidae. Results suggest that Euparagiinae is the sister group of all other Vespidae. The new genus Protovespa appears more closely related to extant Masarinae, Eumeninae and social wasps than to Priorvespinae. We assign it to a new subfamily: Protovespinae. Finally, fossil information combined with a phylogenetic tree shows that the main groups of Vespidae probably evolved during the Early Cretaceous. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1E7E4796‐6E70‐4D81‐BB34‐0FEEA765DC25 .     

Mitochondrial genome organization and phylogeny of two vespid wasps

We sequenced the entire mitochondrial genome of Abispa ephippium (Hymenoptera: Vespoidea: Vespidae: Eumeninae) and most of the mitochondrial genome of Polistes humilis synoecus (Hymenoptera: Vespoidea: Vespidae: Polistinae). The arrangement of genes differed between the two genomes and also differed slightly from that inferred to be ancestral for the Hymenoptera. The genome organization for both vespids is different from that of all other mitochondrial genomes previously reported. A number of tRNA gene rearrangements were identified that represent potential synapomorphies for a subset of the Vespidae. Analysis of all available hymenopteran mitochondrial genome sequences recovered an uncontroversial phylogeny, one consistent with analyses of other types of data.     

Sperm structure and ultrastructure in the Hymenoptera (Insecta)

A light and electron microscopical survey of spermatozoan gross morphology and ultrastructure in the Hymenoptera is presented. Details are provided for the first time for members of the families Xyelidae, Argidae, Tenthredinidae, Diprionidae, Cephidae, Figitidae, Proctotrupidae, Diaprii- dae, Heloridae, Eurytomidae, Leucospidae, Perilampidae, Torymidae, Braconidae, Dryinidae, Sphecidae, Pompilidae and Vespidae. Spermatozoan length ranged from 8 μ m in some Braconidae to 500 μm in one chalcidoid. Considerable variation in gross morphology and ultrastructure were observed between taxa. Several phylogenetically informative characters were noted. Very small spermatozoa characterized most of the non-cyclostome subfamilies of Braconidae; spirally twisted axoneme and mitochondrial derivatives occur in the Eulophidae, Eurytomidae and Pteromalidae; spermatozoa with virtually indistinguishable head (nucleus and acrosome) regions characterized the Vespinae and Polistinae. The presence of well-developed spermatodesmata in the vas deferens and seminal vesicle characterize the Symphyta and were largely absent from other groups though they are occasionally present in some bees.     

Diversity of wasps (Hymenoptera: Aculeata: Vespidae) along an altitudinal gradient of Atlantic Forest in Itatiaia National Park,Brazil

Surveying the diversity of stinging wasps (Hymenoptera: Aculeata) provides an important information base to assist in biodiversity conservation and the management of forest reserves, as wasps depend on and maintain the population balance of several other groups of insects. In accordance, this paper presents an altitudinal survey of wasps (Hymenoptera, Aculeata, Vespidae) in Itatiaia National Park, Brazil, which is a protected area covered by Atlantic Forest in a mountainous landscape, with altitudes ranging between 540 and 2791 metres above sea level. Six altitudinal zones were sampled with entomological net, and the abundance and diversity of the species were indicated by zones. Field sampling took 288 h of discontinuous activity, which was randomly conducted from December 2012 to December 2013. A total of 398 individuals belonging to 29 species and two subfamilies (Eumeninae and Polistinae) were sampled. Eight species are new records for the state of Rio de Janeiro. We found a monotonic decrease in wasp diversity in relation to altitude, and the number of captured individuals differed significantly between the low and high altitudes.     

Phylomitogenomics reveals mito-nuclear concordance in social wasps: The performance of mitochondrial markers and gene order for hymenopteran systematics

Mitochondrial (mtDNA) genes have served as widely utilised genetic loci for phylogenetic and phylogeographic studies of animals. However, the phylogenetic performance of many mtDNA genes has not been empirically evaluated across lineages within hymenopteran wasps. To address this question, we assembled and analysed mitogenomic data from social wasps, representing the four recognised tribes of Polistinae and all Epiponini genera. Additionally, we evaluated whether mtDNA gene order in Polistinae is congruent with its tribal classification. Using concatenation phylogenetic methods, we show phylogenetic congruence between mitogenomic and nuclear data. Statistically comparing the phylogenetic performance of individual mtDNA genes, we demonstrate that for social wasps the molecular markers COI, 16S, NAD5, and NAD2 perform best, while ATP6, COII, and 12S show the worst results. Finally, we verified that the tRNA cluster close to the noncoding region is a hotspot of genetic rearrangements in Vespidae and can be used as additional information for the systematics of this group. Together, these results indicate that mitogenomes contain robust phylogenetic signal to elucidate the evolutionary history of Vespidae. Moreover, our study identifies the best choice of mtDNA markers for systematic investigations of social wasps.     

The higher classification of the ant subfamily Leptanillinae (Hymenoptera: Formicidae)

Abstract. Until now the ant subfamily Leptanillinae has been closely linked with the army ant subfamilies Dorylinae and Ecitoninae, but on relatively tenuous evidence. The current phylogenetic study strongly indicates that this view is incorrect and that the leptanillines really constitute the sister-group of subfamily Ponerinae, and are at a consider- ably greater taxonomic distance from the Army Ant subfamilies. Three tribes are now recognized within the Leptanillinae (Leptanillini; Anomalornyrmini, new tribe; and Apomyrmini, transferred here from Ponerinae: Arnblyoponini), containing a total of eight genera with fewer than fifty species in all. The subfamily and its component tribes are diagnosed and discussed here, and a key to genera provided. New taxa described include Anomalomyrma Taylor gen.n., type-species A. taylori Bolton sp.n. and Protanilla Taylor gen.n., type-species P. rafflesi Taylor sp.n.     

The phylogeny of the social wasp subfamily Polistinae: evidence from microsatellite flanking sequences,mitochondrial COI sequence,and morphological characters

Background  

Social wasps in the subfamily Polistinae (Hymenoptera: Vespidae) have been important in studies of the evolution of sociality, kin selection, and within colony conflicts of interest. These studies have generally been conducted within species, because a resolved phylogeny among species is lacking. We used nuclear DNA microsatellite flanking sequences, mitochondrial COI sequence, and morphological characters to generate a phylogeny for the Polistinae (Hymenoptera) using 69 species.     

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.     

Phylogeny of the tachyporine group subfamilies and 'basal' lineages of the Aleocharinae (Coleoptera: Staphylinidae) based on larval and adult characteristics

Abstract. This paper reports the conclusions of studies into the phylogeny of tachyporine group subfamilies and the ‘basal’ lineages of the subfamily Aleocharinae (Coleoptera: Staphylinidae) based on both larval and adult morphological data (133 adult characters, twenty-seven larval characters). Representatives of forty species of the tachyporine group were used in the analysis, including representatives of the Aleocharinae, Trichophyinae, Habrocerinae, Phloeocharinae, Olisthaerinae, and Tachyporinae. The Aleocharinae included representatives of the tribes Gymnusini, Deinopsini, Mesoporini, the ‘subfamily’ Trichopseniinae, and representatives of nine major tribes in the ‘higher’ Aleocharinae (Athetini, Hoplandriini, Falagriini, Lomechisini, Oxypodini, Aleocharini, Myllaenini, Homalotini, and Hypocyphtini). Analyses were performed first with adult characters alone and then with both larval and adult characters in a simultaneous analysis. The analysis based on adult characters produced eighty-five equally parsimonious trees (length = 499, consistency index = 42; retention index = 69). In the consensus tree, the Tachyporinae are not monophyletic, and the sister-group relationship between the Trichophyinae + Habrocerinae and the Aleocharinae is not resolved. The Aleocharinae are monophyletic, but, among the ‘basal’ Aleocharinae, the relationships of Gymnusini + Deinopsini, the Mesoporini, and the Trichopseniinae are unresolved. The combined adult and larval data, using Tachinus as the outgroup, produced six equally parsimonious trees (tree length = 588; consistency index = 43; retention index = 69). The strict consensus tree of the combined larval and adult data supports the following conclusions: (1) larval characters substantially stabilize the tree; (2) the subfamily Tachyporinae is not supported to be monophyletic; (3) the subfamilies Trichophyinae and Habrocerinae are sister groups, and together they are sister to the Aleocharinae; (4) the ‘basal’ Aleocharinae are not a monophyletic group, but the ‘higher’ Aleocharinae are monophyletic; (5) the sister group of the remaining Aleocharinae is a lineage made up of genera currently in the tribes Gymnusini and Deinopsini; (6) within the Gymnusini–Deinopsini lineage, the monophyly of the Gymnusini is weakly supported, but the monophyly of the Deinopsini is strongly supported; (7) the subfamily Trichopseniinae is strongly supported to be a member of the ‘basal’ Aleocharinae; (8) the Myllaenini are resolved well within the ‘higher’ Aleocharinae; (9) strong support for the monophyly of some tribes of ‘higher’ Aleocharinae suggests that morphological characters provide substantial phylogenetic signal for analysis of higher-level phylogeny of the Aleocharinae in spite of the preliminary nature of the analysis at this taxonomic level.     

Insights into the phylogenetic relationships within Cixiidae (Hemiptera: Fulgoromorpha): cladistic analysis of a morphological dataset

Abstract.  According to the most recent classifications proposed, the planthopper family Cixiidae comprises three subfamilies, namely Borystheninae, Bothriocerinae and Cixiinae, the latter with 16 tribes. Here we examine morphological characters to present the first phylogenetic reconstructions within Cixiidae derived from a cladistic analysis. We scored 85 characters of the head, thorax, and male and female genitalia for 50 taxa representative of all cixiid subfamilies and tribes and for six outgroup taxa. Analyses were based on maximum parsimony – using both equally weighted and successive weighting procedures – and Bayesian inferences. The monophyly of most currently accepted tribes and subfamilies was investigated through Templeton statistical tests of alternative phylogenetic hypotheses. The cladistic analyses recover the monophyly of Cixiidae, the subfamily Bothriocerinae, and the tribes Pentastirini, Mnemosynini, and Eucarpiini. Successive weighting and Bayesian inference recover the monophyly of the tribe Gelastocephalini, but only Bayesian inference supports the monophyly of Semoniini. The relationships recovered support the groups [Stenophlepsini (Borystheninae + Bothriocerinae)] arising from the tribe Oecleini, and [Andini + Brixiidini + Brixiini (polyphyletic) + Bennini]. Templeton tests reject the alternative hypothesis of a monophyletic condition for the tribe Pintaliini as presently defined.     

The cladistics and higher classification of the Pimpliformes (Hymenoptera: Ichneumonidae)

The monophyly of the ichneumonid clade Pimpliformes is established and the phylogenetic relationships of the eight component subfamilies are resolved. The clade (Acaenitinae + (Diacritinae + (Cylloceriinae + (Diplazontinae + Orthocentrinae)))) is the sister-lineage to the clade (Pimplinae + (Rhyssinae + Poemeniinae)). The Nearctic genus Cressonia Dasch is transferred to the Diacritinae from the Orthocentrinae. Tribes are not recognized in the Acaenitinae as the Coleocentrini (sensu Townes, 1971) is paraphyletic with respect to the Acaenitini. The Cylloceriinae is recognized as comprising three genera, Cylloceria Schiødte, Allomacrus Förster and Sweaterella gen.n. The Orthocentrinae, including the Helictinae of authors, is shown to be monophyletic, but the latter is clearly shown to be paraphyletic if the Orthocentrus genus-group is excluded. The Pimplinae comprises four monophyletic tribes: the Delomeristini, consisting of Delomerista Förster and Atractogaster Kriechbaumer; the Perithoini trib.n., which includes only Perithous Holmgren (= Hybomischos Baltazar syn.n.); the Pimplini, which includes the Theronia genus-group as well as the Pimpla genus-group; and the Ephialtini, which includes the Polysphinctini syn.n., a monophyletic group that previously rendered the restricted Ephialtini paraphyletic. The tribe Delomeristini is the sister-group to the clade (Ephialtini + (Perithoini + Pimplini)). The subfamily Poemeniinae is recognized as comprising three tribes: the Pseudorhyssini (trib.n.) which includes the single Holarctic genus Pseudorhyssa Merrill; the Rodrigamini (trib.n.) which includes only the Costa Rican genus Rodrigama Gauld; and the Poemeniini. The tribe Pseudorhyssini is the sister-group to the clade (Rodrigamini + Poemeniini). The phylogenetic inter-relationships of the genera of Poemeniini are resolved. A new genus from South Africa, Guptella (gen.n.) is described, and Achorocephalus Kriechbaumer is shown to be a synonym of Eugalta Cameron (syn.n.). The evolution of biological traits within the Pimpliformes is discussed with reference to the elucidated phylogeny, and zoogeographic patterns are outlined.     

角马蜂贮精囊及精子的形态学观察

昆虫贮精囊和精子的形态多样性是重要的分类和系统发育分析特征之一, 然而在马蜂亚科乃至整个胡蜂科中却鲜有涉及。本文首次解剖了角马蜂Polistes chinensis antennalis Pérez的雄性生殖系统, 着重对其贮精囊的超微结构进行描述, 并简要报道了精子的外部形态。角马蜂的贮精囊由输精管亚前端膨大而成, 有一层发达的柱状上皮细胞贴在基底膜内壁： 细胞核位于柱状细胞基部, 上皮细胞端半部线粒体密集, 顶膜特化成微绒毛。角马蜂精子头长21.4 μm, 体长94 μm, 是已报道胡蜂科精子中长度最短、 相对头长最长的种类。研究结果为胡蜂科昆虫系统发育以及繁殖生理提供理论依据。     

Phylogenetic relationships of the Asian palm civets (Hemigalinae & Paradoxurinae, Viverridae, Carnivora)

The Viverridae (Mammalia, Carnivora), one of the least studied groups of carnivorans, include two subfamilies of Asian palm civets: Hemigalinae and Paradoxurinae. The relationships between and within these two subfamilies have never been thoroughly tested using an extensive molecular sample set. In this study, we gathered sequences of four genes (two mitochondrial: Cytochrome b and ND2 and two nuclear: beta-fibrinogen intron 7 and IRBP exon 1) for eight of the eleven extant species representing these two subfamilies. The results showed that: (1) the Asian palm civets (Hemigalinae and Paradoxurinae) have a single origin and form the sister-group of the (Genettinae+Viverrinae) clade, (2) the Hemigalinae (including the otter civet Cynogale bennettii) are monophyletic, (3) the Paradoxurinae are monophyletic and (4) the small-toothed palm civet (Arctogalidia trivirgata) is an early offshoot within the Paradoxurinae. Using a relaxed molecular clock analysis, the differentiation of the (Hemigalinae+Paradoxurinae) was inferred to occur in the Late Oligocene/Early Miocene.     

Subfamily Dichomeridinae (Lepidoptera, Gelechiidae): Phylogeny, classification, and position in the system of gelechiid moths

On the basis of comparative morphological analysis, taking into account the male genital musculature, a cladogram of the tribes and subfamilies of gelechiid moths is proposed. In accordance with the main branches of the cladogram the new system of the family Gelechiidae consisting of 5 subfamilies (Physoptilinae, Anomologinae, Gelechiinae, Anacampsinae, and Dichomeridinae) is developed. The subfamily Physoptilinae is considered as a sister-group to other gelechiids, the monophyly of the latter is supported by a complex of synapomorphies (female retinaculum with anteriorly directed row of scales located on the base of radial stalk; muscles m ₃ running across the longitudinal body axis, connecting the lateral arms of vinculum with margins of juxta; protractors of aedeagus m ₅ divided into two bunches m _5a and m _5b). The subfamilies Anacampsinae and Dichomeridinae are regarded as sistergroups. The monophyly of the subfamily Dichomeridinae is based on the complex of synapomorphies (presence of parategminal sclerites, which are the apodemes for muscles m ₄; presence of distinct ventral wall in tegumen; intrategminal position of the muscles m ₂). The subfamily Dichomeridinae is considered to consist of three tribes, Anarsiini, Chelariini and Dichomeridini, with 34 genera in total. The cladogram for the genera of the subfamily Dichomeridinae is proposed.     

Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea)

A comprehensive tribal‐level classification for the world’s subfamilies of Hesperiidae, the skipper butterflies, is proposed for the first time. Phylogenetic relationships between tribes and subfamilies are inferred using DNA sequence data from three gene regions (cytochrome oxidase subunit I‐subunit II, elongation factor‐1α and wingless). Monophyly of the family is strongly supported, as are some of the traditionally recognized subfamilies, with the following relationships: (Coeliadinae + (“Pyrginae” + (Heteropterinae + (Trapezitinae + Hesperiinae)))). The subfamily Pyrginae of contemporary authors was recovered as a paraphyletic grade of taxa. The formerly recognized subfamily Pyrrhopyginae, although monophyletic, is downgraded to a tribe of the “Pyrginae”. The former subfamily Megathyminae is an infra‐tribal group of the Hesperiinae. The Australian endemic Euschemon rafflesia is a hesperiid, possibly related to “Pyrginae” (Eudamini). Most of the traditionally recognized groups and subgroups of genera currently employed to partition the subfamilies of the Hesperiidae are not monophyletic. We recognize eight pyrgine and six hesperiine tribes, including the new tribe Moncini. © The Willi Hennig Society 2008.     

First evidence of wasp brood development inside active nests of a termite with the description of a previously unknown potter wasp species

Potter wasps (Vespidae: Eumeninae) are known to exhibit not only sophisticated preying strategies but also a remarkable ability to manipulate clay during nest building. Due to a mixture of plasticity in building behavior and flexibility in substrate preferences during nest building, the group has been reported nesting in a variety of places, including decaying nests abandoned by termite species. Yet, evidence of wasps nesting inside senescent termite mounds is poorly reported, and to date, accounts confirming their presence inside active colonies of termites are absent. Here, we address a novel intriguing association between two species from the Brazilian Cerrado: a previously unknown potter wasp (nest invader) and a termite species (nest builder). Besides scientifically describing Montezumia termitophila sp. nov. (Vespidae: Eumeninae), named after its association with the termite Constrictotermes cyphergaster (Silvestri, 1901) (Termitidae: Nasutitermitinae), we provide preliminary information about the new species'' bionomics by including (a) a hypothetical life cycle based on the evidence we collected and (b) a footage showing the first interaction between a recently ecloded wasp and a group of termites. In doing so, we attempt to provoke relevant discussions in the field and, perhaps, motivate further studies with the group. Finally, we describe a solution to efficiently detect and sample termitophilous species from termite nests, an intrinsic yet challenging task of any studies dealing with such a cryptic biological system.