After more than half a century: Dielis diabo (Hymenoptera: Scoliidae: Scoliinae), a new scoliid from the Neotropics.

A new species of Scoliidae, Dielis diabo n. sp., is described from material previously collected and misidentified as Dielis dorsata Fabricius, 1787 after almost 60 years since the last description of a neotropical scoliid. Detailed morphological descriptions of the female and male are presented, as well as differences between the new species and Dielis dorsata.

Behavioural evolution of Neotropical social wasps (Vespidae: Polistinae): the queen selection process.

All Epiponini wasps are polygynic, with multiple queens alternating over the colony cycle. There are several potential queens in the early stages of this cycle, but as it progresses, the number of queens is reduced. Because most individuals remain reproductively totipotent, there is great potential for conflicts over reproduction. Workers could have an advantage in controlling queen production because they are much more numerous than queens. Nevertheless, the queen selection process is little known for Epiponini. For this reason, we aimed to study the behaviour of queens and workers during queen selection in multiple species of Epiponini, integrate information from previous behavioural studies, and perform a comparative analysis to interpret changes evolutionarily. We conducted observations on nine species belonging to five genera: Brachygastra, Chartergellus, Metapolybia, Polybia and Protopolybia. Females were individually marked to make direct and video observations. Queen production was artificially induced. A total of 28 behaviours related to queen selection were identified. The most aggressive interactions between castes, such as bite and dart, were lost in the major lineages of Epiponini. Bending display I is an ancient behaviour used as the main dominance display. Behaviours exhibited by workers to test queen status arose in the common ancestor of the Epiponini and are not shared by other polistine wasps. Consequently, the act of workers testing queen status probably was present in the Epiponini ancestor. Ritualized test display and dominance behaviours are used in Epiponini as honest signals of the queen's reproductive potential instead of aggressive behaviours. Caste flexibility had already been suggested as the ground plan for Epiponini and is herein discussed as decisive for colony survival of swarm wasps, because it allows colonies to respond efficiently to different situations that may eventually arise.

Histology and ultrastructure of glands associated with the sting apparatus of the swarm-founding wasp Protopolybia exigua (Hymenoptera: Vespidae: Polistinae).

Social insects present a great diversity of exocrine glands, which are linked to fundamental roles of social life, and their morphological characterization represents the first step toward the knowledge of their function. We here describe the exocrine glands present in the sting apparatus of queens and workers of the wasp Protopolybia exigua. Histological analysis of serial sections under light microscopy revealed a glandular epithelium in the spiracular plate, the quadrate plate, and the gonostyli, while the last two in addition also contain clusters of class-3 cells. A big cluster of class-3 cells was also found in the oblong plate of some workers and queens. These findings reveal that a variety of glands, in addition to the common venom gland and Dufour gland, are present in association with the sting apparatus, of which three glands are novel reports for the exocrine repertoire of social insects. The position of the glands suggests a role in lubrication to provide mobility of the sting and venom ejection and/or participation in alarm behavior, while a function related to reproduction is also possible. The peculiar characteristics displayed by swarm-founding wasps are linked with exocrine gland secretions, making their investigation in Epiponini wasps a vast and interesting field.

Notes on Zethini (Hymenoptera, Vespidae, Eumeninae): complementary sex descriptions, synonymies and a new species of Zethus Fabricius.

This paper brings a series of contributions to the taxonomy of the Zethini: description of a new species, Zethus denticlypeus Lopes, sp. n.; description of the female of Z. anomalus Cooper, 1999; description of the males of Z. angustior Bohart Stange, 1965, Z. clypeolaris Bohart Stange, 1965, Z. satanicus Bohart Stange, 1965 and Argentozethus willinki Stange, 1979; synonymy of Discoelius argentinus Brèthes, 1905 under Z. dicomboda Spinola, 1851; and rearrangements for the species of Raphiglossoides Giordani Soika, 1936, with Raphiglossoides minutus (Gusenleitner, 2000) (= R. aethiopicus Giordani Soika, 1936) and R. gibbus (Gusenletiner, 2000) n. comb.

Taxonomic Review of Eumenes Latreille, 1802 (Hymenoptera, Vespidae, Eumeninae) from the New World.

The genus Eumenes Latreille, 1802, from the New World, is revised based on external morphology and male genitalia. New synonymy is proposed as follows: E. aureus Isely, 1917 = E. americanus de Saussure, 1852; E. bollii oregonensis Bequaert, 1838, and E. b. ehrenbergi Zavattari, 1912 = E. bollii Cresson, 1872; E. consobrinus pedalis Fox, 1894 = E. consobrinus de Saussure, 1855; E. crucifera bolliformis Viereck, 1908, E. c. nearcticus de Saussure, 1855 and E. c. stricklandi Bequaert, 1944 = E. crucifera Provancher, 1888; E. smithii belfragei Cresson, 1872 = E. smithii de Saussure, 1852; E. verticalis coloradensis Cresson, 1875, E. v. neoboreus Bequaert, 1944, E. v. tricinctus Isely, 1917 = E. verticalis Say, 1824. Eumenes flavitinctus Bohart, 1950, revised status, is no longer a subspecies of E. crucifera but elevated to specific level. Eumenes brunneus is elevated to the specific level and a new name is proposed because of homonymy, E. bequaerti Grandinete Carpenter nom. nov. An identification key for all the species of the genus from the New World is provided and the geographical distributions are updated.

On the dumping ground genus Scotaena Klug, 1810 (Hymenoptera: Tiphiidae: Thynninae): Phylogeny, taxonomy and geographic distribution.

The Neotropical thynnine genus Scotaena is revised and a cladistic analysis is presented. The analysis, conducted from 75 morphological characters of 31 terminal taxa, returned a single tree under equal weighting. The monophyly of Scotaena was not recovered. Three new genera and five new species are described: Kaysara gen. nov., Pseudoscotaena gen. nov. and Pampathynnus gen. nov., Scotaena reversa sp. nov., Kaysara laterolata sp. nov., Kaysara apiciconcava sp. nov., Kaysara marginoplicata sp. nov. and Kaysara levicrenata sp. nov. Three species are transferred to other genera as follows: Eucyrtothynnus rosenbergi (Turner, 1910) comb. nov., Glottynoides genisei Kimsey, 1991 comb. nov., Ornepetes clypearis Durán-Moya, 1941 comb. nov. Scotaena now comprises four species: S. trifasciata Klug, 1810; S. horni (Turner, 1927); S. vetusta Turner, 1909; and S. reversa. An identification key and geographical distribution maps for the studied species are also provided.

Phylogeographic analysis reveals high genetic structure with uniform phenotypes in the paper wasp Protonectarina sylveirae (Hymenoptera: Vespidae).

Swarm-founding wasps are endemic and common representatives of neotropical fauna and compose an interesting social tribe of vespids, presenting both complex social characteristics and uncommon traits for a eusocial group, such as the absence of castes with distinct morphology. The paper wasp Protonectarina sylveirae (Saussure) presents a broad distribution from Brazil, Argentina and Paraguay, occurring widespread in the Atlantic rainforest and arboreal Caatinga, being absent in the Amazon region. Given the peculiar distribution among swarm-founding wasps, an integrative approach to reconstruct the evolutionary history of P. sylveirae in a spatial-temporal framework was performed to investigate: the presence of genetic structure and its relationship with the geography, the evolution of distinct morphologic lineages and the possible historical event(s) in Neotropical region, which could explain the observed phylogeographic pattern. Individuals of P. sylveirae were obtained from populations of 16 areas throughout its distribution for DNA extraction and amplification of mitochondrial genes 12S, 16S and COI. Analysis of genetic diversity, construction of haplotype net, analysis of population structure and dating analysis of divergence time were performed. A morphometric analysis was also performed using 8 measures of the body of the adult (workers) to test if there are morphological distinction among populations. Thirty-five haplotypes were identified, most of them exclusively of a group and a high population structure was found. The possibility of genetic divergence because of isolation by distance was rejected. Morphological analysis pointed to a great uniformity in phenotypes, with only a small degree of differentiation between populations of south and the remaining. Divergence time analysis showed a Middle/Late Miocene origin, a period where an extensive marine ingression occurred in South America. Divergence of haplogroups began from the Plio/Pleistocene boundary and the last glacial maximum most likely modeled the current distribution of species, even though it was not the cause of genetic breaks.

Three new species of tribe Odynerini (Hymenoptera, Vespidae, Eumeninae) from the Neotropical Region.

Three new species from the Neotropical region are described: Ancistroceroides soikai Grandinete & Carpenter, sp. nov. (Costa Rica, Paraguay); Cephalastor huautla Grandinete & Carpenter, sp. nov. (Mexico), and Symmorphus centralis Carpenter & Grandinete, sp. nov. (Costa Rica).

Ontogenic Caste Differences in the Van der Vecht Organ of Primitively Eusocial Neotropical Paper Wasps.

Recent studies have reported incipient morphological caste dimorphism in the Van der Vecht organ size of some temperate Polistes paper wasps. Whether species other than the temperate ones show a similar pattern remains elusive. Here, we have studied some Neotropical Polistes species. By comparing females collected through the year, we showed caste related differences in the size of the Van der Vecht organ in P. ferreri (body size corrected Van der Vech organ size of queens = 0.45 ± 0.06, workers = 0.38 ± 0.07 mm2, p = 0.0021), P. versicolor (body size corrected Van der Vech organ size of queens = 0.54 ± 0.11, workers = 0.46 ± 0.09 mm2, p = 0.010), but not P. simillimus (body size corrected Van der Vech organ size of queens = 0.52 ± 0.05, workers = 0.49 ± 0.06 mm2, p = 0.238). Therefore, it seems that queens and workers of some Neotropical Polistes have diverged in their ontogenic trajectory of the Van der Vecht organ size, providing clear evidence for incipient morphological caste dimorphism. As Polistes are distributed mostly in the tropics, we propose that physical caste differences may be widespread in the genus. Also, we highlight that morphological divergence in the queen-worker phenotypes may have started through differential selection of body structures, like the Van der Vecht organ.

Additions to the checklist of Scoliidae, Sphecidae, Pompilidae and Vespidae of Peru, with notes on the endemic status of some species (Hymenoptera, Aculeata).

The first checklist of the Peruvian Hymenoptera listed 1169 species and subspecies of aculeate wasps, including 173 species of Pompilidae, seven of Scoliidae, 39 of Sphecidae and 403 of Vespidae. Herein are reported 32 species as new for Peru based mainly on the collection of the Natural History Museum, London. The loss of the endemic status of two species is also reported: Entypus peruvianus (Rohwer) (Pompilidae: Pepsinae) and Omicron ruficolle schunkei Giordani Soika (Vespidae: Eumeninae).

Cladistic analysis of self-grooming indicates a single origin of eusociality in corbiculate bees (Hymenoptera: Apidae).

Behavioural traits have been used extensively in recent years as an important character source for making phylogenetic inferences. The phylogenetic positions of the members of the Apini subtribe are increasingly being debated, and new characters must be examined. We analysed the presence and absence of certain behavioural patterns, as well as the sequences of some of these patterns, to generate 79 characters. Eleven species comprised the ingroup, and Xylocopini comprised the outgroup. Parsimony analysis showed that the most parsimonious tree was (Euglossina(Bombina(Apina+Meliponina))). This topology is consistent with most studies that use morphological data and the few that use behavioural data, which suggests that advanced eusociality arose only once in a common ancestor of the clade Apina plus Meliponina; however, this hypothesis is inconsistent with our molecular data. Thus we considered behavioural, molecular, and morphological data and recovered the same topology, in which eusociality has a single origin in corbiculate bees.

Scanning electron microscopy of male terminalia and its application to species recognition and phylogenetic reconstruction in the Drosophila saltans group.

The Drosophila saltans group consists of five subgroups and 21 species, most of which have been identified only by morphological aspects of the male terminalia revealed by drawings using a camera lucida and a bright-field microscope. However, several species in the group, mainly those included in the saltans subgroup, are difficult to differentiate using only these characteristics. In this study, we used scanning electron microscopy (SEM) to analyze 19 structures of the male terminalia in 10 species from the five saltans subgroups. Among these structures, nine could be identified only through SEM analysis. We aimed to find other characteristics useful for morphological recognition of these species and to use these characteristics for phylogenetic reconstruction. These morphological differences enabled us to effectively distinguish among sibling species. These findings confirmed the monophyly of this group as previously determined in evolutionary studies based on other markers. The single most parsimonious tree (CI = 87 and RI = 90) indicated that the cordata subgroup is the most basal lineage and the saltans subgroup is the most apical lineage, as shown in earlier studies based on morphological data. However, our findings differed somewhat from these studies with respect to the phylogenetic relationships of species in the saltans group indicating that this group is still a puzzle that remains to be deciphered.

Morphological and physiological variation between queens and workers of Protonectarina sylveirae (de Saussure) (Hymenoptera, Vespidae, Epiponini) / Variações morfológicas e fisiológicas entre rainhas e operárias de Protonectarina sylveirae (de Saussure) (Hymenoptera, Vespidae, Epiponini)

Morphological and physiological variation between queens and workers of Protonectarina sylveirae (de Saussure) (Hymenoptera, Vespidae, Epiponini). The Neotropical swarm-founding wasps, Epiponini, range from the absence of morphological differentiation between castes to highly distinct castes. We measured eight body parts of females of two colonies of Protonectarina sylveirae (de Saussure, 1854). ANOVA and Discriminant Analysis evidenced significant differences between castes, as previously observed by other authors for other species of Epiponini. However, some females previously categorized as queens, were actually workers, supported by our statistic analyses. These individuals showed intermediate morphological features between queens and workers, having distinct patterns of hairs and clypeal spots. The castes of P. sylveirae are distinct, however intermediate individuals may be found in colonies promoting social flexibility.

As vespas enxameadoras neotropicais, Epiponini, apresentam desde ausência de diferenciação morfológica entre as castas até castas altamente distintas. Medimos oito partes corporais das fêmeas de duas colônias de Protonectarina sylveirae (de Saussure, 1854). ANOVA e Análise Discriminante evidenciaram diferenças significativas entre as castas, como observado anteriormente por outros autores em outros Epiponini. No entanto, algumas fêmeas, previamente categorizadas como rainhas, eram na verdade operárias, com base nas análises estatísticas. Esses indivíduos apresentaram características morfológicas intermediárias entre rainhas e operárias, tendo padrões distintos de pilosidade e da marca do clípeo. As castas em P. sylveirae são distintas, no entanto indivíduos intermediários podem ser encontrados nas colônias promovendo uma flexibilidade social.

Morphological studies on castes of Protopolybia chartergoides (Hymenoptera, Vespidae, Epiponini) observed in colonies during male production stage

Social wasps are important for the study of social behavior evolution because their colonies present different degrees of caste differentiation, from morphologically similar to highly conspicuous, associated with the occasional presence of intermediate females, which bears developed ovaries but no insemination. In the Polistinae, depending on the taxon, such differentiation can be discreet or conspicuous. This work intended to study morphological and physiological differences between castes in Protopolybia chartergoides by using morphometric analyses associated with multivariate statistical analyses and physiological evaluations from females' ovarian development. Results evidence low morphological and physiological differences among the castes in P. chartergoides, indicating three groups of females: queens, workers and intermediates. In this way, it was possible to suggest that Protopolybia chartergoides presents post-imaginal caste differentiation (or a very subtle form of pre-imaginal determination).

Vespas sociais são importantes para o estudo da evolução do comportamento social devido suas colônias apresentarem diferentes graus de diferenciação de castas, de morfologicamente similares às altamente conspícuas, associadas com a presença de fêmeas intermediárias, as quais desenvolvem seus ovários, mas não estando inseminadas. Nos Polistinae, tais diferenciações podem ser discretas ou conspícuas. Este trabalho teve como objetivo estudar as diferenças morfológicas e fisiológicas entre as castas de Protopolybia chartergoides pelo uso de análises morfométricas associada com análises estatísticas multivariadas e avaliações fisiológicas do desenvolvimento ovariano das fêmeas. Os resultados mostraram baixa diferenciação morfológica e fisiológica entre as castas de P. chartergoides, indicando a presença de três grupos de fêmeas: rainhas, operárias e intermediárias. Assim, foi possível sugerir que Protopolybia chartergoides apresenta diferenciação de casta pós-imaginal (ou uma forma muito sutil de determinação pré-imaginal).

Non-ovipositing nurse workers with developed ovaries in trigona cilipes cilipes (Hymenoptera, Meliponini)

Numerous species of tropical stingless bees (Hymenoptera, Meliponini) have workers that lay eggs even inthe presence of a queen, i.e., queenright colonies. These worker eggs, which are laid during cell provisioningand oviposition process (POP), are eaten mainly by the queen. In this work, we studied POP in a colonyof Trigona cilipes in the queenright and queenless stages. POP behavior was observed on 67 occasions. Inaddition, the ovaries of different aged workers in both stages were analyzed. Although workers in both stagesof the colony possessed developed ovaries, they never oviposited, despite showing behavior similar to thatseen in species with workers which lay eggs. This form of sterility is outstanding among social insects andis comparable only to other meliponines that have fully sterile workers.