Marimbondos: systematics, biogeography, and evolution of social behaviour of neotropical swarm-founding wasps (Hymenoptera: Vespidae: Epiponini).

Neotropical swarm-founding wasps are divided into 19 genera in the tribe Epiponini (Vespidae, Polistinae). They display extensive variation in several colony-level traits that make them an attractive model system for reconstructing the evolution of social phenotypes, including caste dimorphism and nest architecture. Epiponini has been upheld as a solid monophyletic group in most phylogenetic analyses carried out so far, supported by molecular, morphological and behavioural data. Recent molecular studies, however, propose different relationships among the genera of swarm-founding wasps. This study is based on the most comprehensive epiponine sampling so far and was analyzed by combining morphological, nesting and molecular data. The resulting phylogenetic hypothesis shows many of the traditional clades but still impacts the way certain behavioural characters, such as nest structure and castes, evolved, and thus requires some re-interpretations. Angiopolybia as sister to the remaining Epiponini implies that nest envelopes and a casteless system are plesiomorphic in the tribe. Molecular dating points to an early tribal diversification during the Eocene (c. 55-38 Ma), with the major differentiation of current genera concentrated in the Oligocene/Miocene boundary.

Phylogenomic Evidence Overturns Current Conceptions of Social Evolution in Wasps (Vespidae).

The hypothesis that eusociality originated once in Vespidae has shaped interpretation of social evolution for decades and has driven the supposition that preimaginal morphophysiological differences between castes were absent at the outset of eusociality. Many researchers also consider casteless nest-sharing an antecedent to eusociality. Together, these ideas endorse a stepwise progression of social evolution in wasps (solitary → casteless nest-sharing → eusociality with rudimentary behavioral castes → eusociality with preimaginal caste-biasing (PCB) → morphologically differentiated castes). Here, we infer the phylogeny of Vespidae using sequence data generated via anchored hybrid enrichment from 378 loci across 136 vespid species and perform ancestral state reconstructions to test whether rudimentary and monomorphic castes characterized the initial stages of eusocial evolution. Our results reject the single origin of eusociality hypothesis, contest the supposition that eusociality emerged from a casteless nest-sharing ancestor, and suggest that eusociality in Polistinae + Vespinae began with castes having morphological differences. An abrupt appearance of castes with ontogenetically established morphophysiological differences conflicts with the current conception of stepwise social evolution and suggests that the climb up the ladder of sociality does not occur through sequential mutation. Phenotypic plasticity and standing genetic variation could explain how cooperative brood care evolved in concert with nest-sharing and how morphologically dissimilar castes arose without a rudimentary intermediate. Furthermore, PCB at the outset of eusociality implicates a subsocial route to eusociality in Polistinae + Vespinae, emphasizing the role of mother-daughter interactions and subfertility (i.e. the cost component of kin selection) in the origin of workers.

Phylogenomic analysis of yellowjackets and hornets (Hymenoptera: Vespidae, Vespinae).

The phylogenetic relationships among genera of the subfamily Vespinae (yellowjackets and hornets) remain unclear. Yellowjackets and hornets constitute one of the only two lineages of highly eusocial wasps, and the distribution of key behavioral traits correlates closely with the current classification of the group. The potential of the Vespinae to elucidate the evolution of social life, however, remains limited due to ambiguous genus-level relationships. Here, we address the relationships among genera within the Vespinae using transcriptomic (RNA-seq) data. We sequenced the transcriptomes of six vespid wasps, including three of the four genera recognized in the Vespinae, combined our data with publicly available transcriptomes, and assembled two matrices comprising 1,507 and 3,356 putative single-copy genes. The results of our phylogenomic analyses recover Dolichovespula as more closely related to Vespa than to Vespula, therefore challenging the prevailing hypothesis of yellowjacket (Vespula+Dolichovespula) monophyly. This suggests that traits such as large colony size and high paternity arose in the genus Vespula following its early divergence from the remaining vespine genera.

Phylogeny, landmark analysis and the use of wing venation to study the evolution of social wasps (Hymenoptera: Vespidae: Vespinae).

Wing venation provides useful characters with which to classify extant and fossil insects. Recently, quantification of its shape using landmarks has increased the potential of wing venation to distinguish taxa. However, the use of wing landmarks in phylogenetic analyses remains largely unexplored. Here, we tested landmark analysis under parsimony (LAUP) to include wing shape data in a phylogenetic analysis of hornets and yellow jackets. Using 68 morphological characters, nine genes and wing landmarks, we produced the first total-evidence phylogeny of Vespinae. We also tested the influence of LAUP parameters using simulated landmarks. Our data confirmed that optimization parameters, alignment method, landmark number and, under low optimization parameters, the initial orientation of aligned shapes can influence LAUP results. Furthermore, single landmark configurations never accurately reflected the topology used for data simulation, but results were significantly close when compared to random topologies. Thus, wing landmark configurations were unreliable phylogenetic characters when treated independently, but provided some useful insights when combined with other data. Our phylogeny corroborated the monophyly of most groups proposed on the basis of morphology and showed the fossil Palaeovespa is distantly related to extant genera. Unstable relationships among genera suggest that rapid radiations occurred in the early history of the Vespinae.

Phylogeny and historical biogeography of the paper wasp genus Polistes (Hymenoptera: Vespidae): implications for the overwintering hypothesis of social evolution.

The phylogeny of the paper wasp genus Polistes is investigated using morphological and behavioural characters, as well as molecular data from six genes (COI, 12S, 16S, 28S, H3, and EF1-α). The results are used to investigate the following evolutionary hypotheses about the genus: (i) that Polistes first evolved in Southeast Asia, (ii) that dispersal to the New World occurred only once, and (iii) that long-term monogyny evolved as an adaptation to overwintering in a temperate climate. Optimization of distribution records on the recovered tree does not allow unambiguous reconstruction of the ancestral area of Polistes. While the results indicate that Polistes dispersed into the New World from Asia, South America is recovered as the ancestral area for all New World Polistes: Nearctic species groups evolved multiple times from this South American stock. The final tree topology suggests strongly that the genus first arose in a tropical environment, refuting the idea of monogyny as an overwintering adaptation.

Phylogenetic relationships of yellowjackets inferred from nine loci (Hymenoptera: Vespidae, Vespinae, Vespula and Dolichovespula).

Eusociality has arisen repeatedly and independently in the history of insects, often leading to evolutionary success and ecological dominance. Eusocial wasps of the genera Vespula and Dolichovespula, or yellowjackets, have developed advanced social traits in a relatively small number of species. The origin of traits such as effective paternity and colony size has been interpreted with reference to an established phylogenetic hypothesis that is based on phenotypic data, while the application of molecular evidence to phylogenetic analysis within yellowjackets has been limited. Here, we investigate the evolutionary history of yellowjackets on the basis of mitochondrial and nuclear markers (nuclear: 28S, EF1α, Pol II, and wg; mitochondrial: 12S, 16S, COI, COII, and Cytb). We use these data to test the monophyly of yellowjackets and species groups, and resolve species-level relationships within each genus using parsimony and Bayesian inference. Our results indicate that a yellowjacket clade is either weakly supported (parsimony) or rejected (Bayesian inference). However, the monophyly of each yellowjacket genus as well as species groups are strongly supported and concordant between methods. Our results agree with previous studies regarding the monophyly of the Vespula vulgaris group and the sister relationship between the V. rufa and V. squamosa groups. This suggests convergence of large colony size and high effective paternity in the vulgaris group and V. squamosa, or a single origin of both traits in the most recent common ancestor of all Vespula species and their evolutionary reversal in the rufa group.

The higher-level phylogenetic relationships of the Eumeninae (Insecta, Hymenoptera, Vespidae), with emphasis on Eumenes sensu lato.

Cladistic analyses were carried out to infer the phylogenetic relationships among taxa that were originally part of the large genus Eumenes. Terminals belonging to other eumenine lineages were also included, as well as terminals from other vespid subfamilies. Analyses under equal weights and implied weights were carried out, and better results were obtained with the latter. The results corroborated the monophyly of Eumeninae, and recovered Zethini sensu lato as the sister-lineage to the remaining eumenines. Eumenes sensu lato as originally recognized is paraphyletic relative to Odynerus sensu lato. A natural classification at the tribal level congruent with the phylogenetic results may be proposed, and the names Zethini, Odynerini, and Eumenini are already available. This is the most comprehensive phylogeny of the Eumeninae to date. A new generic synonymy is Alfieria Giordani Soika, 1934 = Delta de Saussure, 1855.

A review of the genus Paraleptomenes Giordani Soika, 1970 (Hymenoptera: Vespidae: Eumeninae: Odynerini) from the Indian subcontinent,with the description of a new species from the eastern Himalayas.

The genus Paraleptomenes Giordani Soika, 1970 is reviewed for the Indian subcontinent. A new species Paraleptomenes darugiriensis Kumar, Carpenter & Sharma, sp. nov. is described. The male of P. rufoniger Giordani Soika, 1994 is described for the first time. The distribution records of P. humbertianus (de Saussure, 1867), P. miniatus mephitis (Cameron, 1901), P. miniatus miniatus (de Saussure, 1855), and P. rufoniger Giordani Soika, 1994 in the Indian states are augmented. A key to species of the Indian subcontinent and a world checklist of species are provided.

A new paper wasp from Late Eocene of France (Hymenoptera: Vespidae: Polistinae).

The new vespid genus and species Palaeopolistes jattioti gen. et. sp. nov. is described from the Late Eocene of Monteils (Gard, France). The new taxon has clear features of the Polistinae but its tribal assignment is uncertain.

Taxonomic account and phylogenetic relationships of the species of the Neotropical social wasp genus Chartergus Lepeletier, 1836 (Vespidae: Polistinae: Epiponini).

The Neotropical social wasp genus Chartergus is reviewed and comparative diagnoses, images of type material, and an updated identification key for species are presented. To investigate the phylogenetic relationships within the genus, we used a combined dataset of morphological characters, nest architecture traits, and molecular data. The results revealed the following relationships among the Chartergus species: (C. globiventris + (C. artifex + C. metanotalis)).

Phylogenomics of Tetraopes longhorn beetles unravels their evolutionary history and biogeographic origins.

Tetraopes longhorn beetles are known for their resistance to milkweed plant toxins and their coevolutionary dynamics with milkweed plants (Asclepias). This association is considered a textbook example of coevolution, in which each species of Tetraopes is specialized to feed on one or a few species of Asclepias. A major challenge to investigating coevolutionary hypotheses and conducting molecular ecology studies lies in the limited understanding of the evolutionary history and biogeographical patterns of Tetraopes. By integrating genomic, morphological, paleontological, and geographical data, we present a robust phylogeny of Tetraopes and their relatives, using three inference methods with varying subsets of data, encompassing 2-12 thousand UCE loci. We elucidate the diversification patterns of Tetraopes species across major biogeographical regions and their colonization of the American continent. Our findings suggest that the genus originated in Central America approximately 21 million years ago during the Miocene and diversified from the Mid-Miocene to the Pleistocene. These events coincided with intense geological activity in Central America. Additionally, independent colonization events in North America occurred from the Late Miocene to the early Pleistocene, potentially contributing to the early diversification of the group. Our data suggest that a common ancestor of Tetraopini migrated into North America, likely facilitated by North Atlantic land bridges, while closely related tribes diverged in Asia and Europe during the Paleocene. Establishing a robust and densely sampled phylogeny of Tetraopes beetles provides a foundation for investigating micro- and macroevolutionary phenomena, including clinal variation, coevolution, and detoxification mechanisms in this ecologically important group.

A taxonomic review of the genus Antodynerus de Saussure, 1855 (Hymenoptera: Vespidae: Eumeninae) from the Indian subcontinent.

The genus Antodynerus de Saussure, 1855 is reviewed from the Indian subcontinent. Three species with one additional subspecies are present, namely Antodynerus flavescens flavescens (Fabricius, 1775), A. f. karachiensis Giordani Soika, 1970, A. limbatus (de Saussure, 1852), and A. punctatipennis (de Saussure, 1853). The parasitic association of strepsipteran insects and the symbiotic association of mites are reported for the first time in the genus Antodynerus. The distribution of A. f. flavescens (Fabricius) and A. limbatus (de Saussure) in the Indian states is augmented. The latter species is newly recorded from China (Tibet). A key to species of the Indian subcontinent is provided.

Well, what about intraspecific variation? Taxonomic and phylogenetic characters in the genus Synoeca de Saussure (Hymenoptera, Vespidae).

Cely and Sarmiento (2011) took issue with the cladistic analysis of relationships among species of the genus Synoeca by Andena et al. (2009a), and presented a reanalysis. They claimed that intraspecific variation in the genus is meaningful, and proper consideration yields a conclusion different from that of Andena et al. Both their critique and reanalysis are vitiated by numerous errors, as is shown in the present paper.

A Catalogue of the Pacific Eumeninae (Hymenoptera: Vespidae).

A catalogue of Eumeninae occurring in Oceania is presented, the first in more than a century.

Revision of the Afrotropical species of the genus Anterhynchium de Saussure (Hymenoptera: Vespidae: Eumeninae).

A taxonomic study on the Afrotropical species in the genus Anterhynchium de Saussure, 1863 is presented. Some subspecies are elevated to specific rank (Anterhynchium argenteopilosellum (Giordani Soika, 1937) stat. nov., A. bugandanum Giordani Soika, 1987 stat. nov., A. cariosum Giordani Soika, 1987 stat. nov., A. denticulatum (Mocsáry, 1903) stat. nov., A. sulphureomaculatum (von Schulthess, 1928) stat. nov.). New synonymies are proposed for Anterhynchium fallax (de Saussure, 1852) (=Rhynchium luctuosum Gerstäcker 1857 syn. nov.), Anterhynchium grandidieri (de Saussure, 1890) (= Epiodynerus grandidieri limbatulus Giordani Soika, 1991 syn. nov.), Anterhynchium grayi (de Saussure, 1855) (= Synagris inermis var. atrata Mocsáry, 1903 syn. nov.; = Rhynchium grayi var. sumptuosum Gribodo, 1895 syn. nov.), Anterhynchium mephisto (Gribodo, 1892) (= Anterhynchium beninum Gusenleitner, 2018 syn. nov.; = Odynerus rufoniger Bequaert, 1918 syn. nov.; = Synagris uncata Tullgren, 1904 syn. nov.); Anterhynchium synagroide (de Saussure, 1852) (=Rhynchium synagroides var. alpha von Schulthess, 1923 syn. nov.; Rhynchium synagroides var. beta von Schulthess, 1923 syn. nov.), Anterhynchium andreanum (de Saussure, 1890) (= Odynerus andreanus discolor Giordani Soika, 1941 syn. nov.); Anterhynchium synagroide gamma (von Schulthess, 1924) is considered as a doubtful synonym of A. argenteopilosellum syn. nov. The nominal subgenus is redefined and divided into the following species-groups: grayi species-group, mephisto species-group, osborni species-group and synagroide species-group. The Malagasy species are transferred from Epiodynerus Giordani Soika to the nominal subgenus but considered as incertae sedis in it. Odynerus natalensis de Saussure, 1855 and Anterhynchium uniforme Gusenleitner, 2012 are transferred to Antodynerus de Saussure, 1855 (Antodynerus natalensis comb. nov. and Antodynerus uniforme comb. nov.), Rhynchium histrionicum Gerstäcker, 1857 is transferred to Euodynerus Dalla Torre, 1904 (Euodynerus (Euodynerus) histrionicus comb. nov.), Rhynchium holomelas André, 1895 is transferred to Pseudagris de Saussure, 1863 (Pseudagris holomelas comb. nov.; = Rhynchium junodianum von Schulthess, 1899 syn. nov.; = Synagris aterrima Maidl, 1914 syn. nov.), Anterhynchium obscurum Gusenleitner, 2007 is transferred to Tricarinodynerus Giordani Soika, 1952 (Tricarinodynerus obscurus comb. nov.). Anterhynchium dahomeyicus Gusenleitner, 2018 is a junior synonym of Aethiopicodynerus punctiventris (Gusenleitner, 2002) syn. nov. Lectotypes for Odynerus andreanus de Saussure, 1890 and Rhynchium sulphureomaculatum von Schulthess, 1928 are designated. A key to the species is provided.

Divergence time analysis of the Neotropical wasp genus Protopolybia Ducke, 1905 (Vespidae, Polistinae, Epiponini) using a Multilocus Phylogenetic Approach.

The genus Protopolybia Ducke was analyzed using a multilocus phylogenetic approach, which confirmed its monophyly. In contrast with the arrangements derived solely from morphological data, however, this analysis divides the genus into only two major clades. The study also presents divergence times for the origin and diversification of Protopolybia, which are related to possible vicariant events occurring in the Amazon biome.

Taxonomic notes on the genus Charterginus Fox, 1898 (Vespidae: Polistinae: Epiponini) with images of the type specimens and new occurrence records.

Charterginus Fox, 1898 is a neotropical genus of swarm-founding social wasps consisting of six species. Despite its ecological significance, there are limited studies on various aspects of these wasps. To address this, this study aims to provide a comprehensive comparative diagnosis for all Charterginus species, accompanied by high-quality images of the type material. Moreover, a distribution map incorporating both previous and newly recorded occurrences is presented, shedding light on the geographical range of these species. Through these efforts, we aim to enhance the knowledge and facilitate future research on Charterginus, ultimately contributing to a better understanding of swarm-founding social wasps.

On homology.

Homology in cladistics is reviewed. The definition of important terms is explicated in historical context. Homology is not synonymous with synapomorphy: it includes symplesiomorphy, and Hennig clearly included both plesiomorphy and synapomorphy as types of homology. Homoplasy is error, in coding, and is analogous to residual error in simple regression. If parallelism and convergence are to be distinguished, homoplasy would be evidence of the former and analogy evidence of the latter. We discuss whether there is a difference between molecular homology and morphological homology, character state homology, nested homology (additive characters), and serial homology. We conclude by proposing a global definition of homology. ©The Will Henning Society 2011.

Phylogenetic relationships among superfamilies of Hymenoptera.

The first comprehensive analysis of higher-level phylogeny of the order Hymenoptera is presented. The analysis includes representatives of all extant superfamilies, scored for 392 morphological characters, and sequence data for four loci (18S, 28S, COI and EF-1α). Including three outgroup taxa, 111 terminals were analyzed. Relationships within symphytans (sawflies) and Apocrita are mostly resolved. Well supported relationships include: Xyeloidea is monophyletic, Cephoidea is the sister group of Siricoidea + [Xiphydrioidea + (Orussoidea + Apocrita)]; Anaxyelidae is included in the Siricoidea, and together they are the sister group of Xiphydrioidea + (Orussoidea + Apocrita); Orussoidea is the sister group of Apocrita, Apocrita is monophyletic; Evanioidea is monophyletic; Aculeata is the sister group of Evanioidea; Proctotrupomorpha is monophyletic; Ichneumonoidea is the sister group of Proctotrupomorpha; Platygastroidea is sister group to Cynipoidea, and together they are sister group to the remaining Proctotrupomorpha; Proctotrupoidea s. str. is monophyletic; Mymarommatoidea is the sister group of Chalcidoidea; Mymarommatoidea + Chalcidoidea + Diaprioidea is monophyletic. Weakly supported relationships include: Stephanoidea is the sister group of the remaining Apocrita; Diaprioidea is monophyletic; Ceraphronoidea is the sister group of Megalyroidea, which together form the sister group of [Trigonaloidea (Aculeata + Evanioidea)]. Aside from paraphyly of Vespoidea within Aculeata, all currently recognized superfamilies are supported as monophyletic. The diapriid subfamily Ismarinae is raised to family status, Ismaridae stat. nov. © The Will Henning Society 2011.

When attempts at robbing prey turn fatal.

Because group-hunting arboreal ants spread-eagle insect prey for a long time before retrieving them, these prey can be coveted by predatory flying insects. Yet, attempting to rob these prey is risky if the ant species is also an effective predator. Here, we show that trying to rob prey from Azteca andreae workers is a fatal error as 268 out of 276 potential cleptobionts (97.1 %) were captured in turn. The ant workers hunt in a group and use the "Velcro®" principle to cling firmly to the leaves of their host tree, permitting them to capture very large prey. Exceptions were one social wasp, plus some Trigona spp. workers and flies that landed directly on the prey and were able to take off immediately when attacked. We conclude that in this situation, previously captured prey attract potential cleptobionts that are captured in turn in most of the cases.