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Phylogenetic studies addressing relationships among chrysidid wasps have been limited. There are no hypotheses proposed for the Neotropical lineages of Chrysidini other than the classic cladogram published in the 1990s by Kimsey and Bohart. Herein we present a cladistic analysis based on 64 morphological characters coded for 54 species of Chrysidini, 32 of them being Ipsiura and 22 representing Caenochrysis, Chrysis, Exochrysis, Gaullea, Neochrysis, and Pleurochrysis. The species of Ipsiura were recovered as monophyletic and as the sister clade of Neochrysis in all most parsimonious trees. We discuss the high plasticity of some morphological characters as evidenced by their high homoplasy in the phylogenetic results, and we clarify the main morphological changes inferred on the phylogenetic tree for this genus. The effects of the inferred homoplasy were evaluated under an implied weighting cladistic analysis, and from a probabilistic perspective with Bayesian inference. Those alternative strategies did not alter the general conclusions about the monophyly of Ipsiura or the generic relationships in Chrysidini (changes were noticed in the species-level relationships within certain parts of Ipsiura, where low branch support was common across all approaches). Among the species groups proposed by Linsenmaier (1985), only the marginalis group was recovered as monophyletic. We also evaluated the convoluted biogeographic history of the group. The resulting historical reconstructions indicate a complicated scenario of diversification of these wasps in the Neotropics, and a close association with forested biomes is discussed.

Stingless bees (tribe Meliponini) are remarkable for their characteristically large social colonies, their capacity to produce honey and other useful products, and their morphological and behavioral diversity. They have a disjunct pan-tropical distribution, primarily occurring in warm and humid environments in the Neotropical, Afrotropical, and Indo-Australasian regions. Even though phylogenetic hypotheses have been proposed for Meliponini based on morphology and molecular data, many questions are still unsolved regarding the evolutionary relationships and systematics of the tribe. In this contribution, we present a large phylogenomic dataset comprising over 2500 ultra-conserved element (UCE) loci sequenced for 153 species of Meliponini, representing all known genera of stingless bees. The genera Camargoia, Paratrigonoides, Plectoplebeia, Cleptotrigona, Ebaiotrigona, Papuatrigona, Pariotrigona, Platytrigona, and Sahulotrigona were included in molecular phylogenetic analyses for the first time. Concatenated and species-tree analyses were performed using different partitioning strategies and summary methods. We performed gene-genealogy interrogation (GGI) on several recalcitrant nodes to resolve discordances among recovered tree topologies. Results were mostly consistent among analyses, recovering three main lineages of Meliponini congruent with the biogeographic domains to which they are associated. Within major clades, discordances were found in relation to previous works. The genus Frieseomelitta was recovered as paraphyletic in relation to Trichotrigona, and the genus Lepidotrigona was revealed to be composed of two independent lineages. Even though concatenated and weighted ASTRAL analyses were mostly effective in recovering the relationships favored by GGI, they retrieved different results in relation to the phylogenetic placements of Oxytrigona and Cephalotrigona. The most favored hypothesis in GGI analyses was not found in any other analyses, being more congruent with morphological evidence and highlighting the relevance of exploring the support given to alternative hypotheses through topological tests. Recent advances in our capacity to generate molecular sequences from old specimens using modern sequencing methods allowed for unparalleled sampling across genera, yielding a backbone for the phylogenetic relationships of stingless bees, which will further investigations into their systematics and evolution., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Although the knowledge of the skeletal morphology of bees has progressed enormously, a corresponding advance has not happened for the muscular system. Most of the knowledge about bee musculature was generated over 50 years ago, well before the digital revolution for anatomical imaging, including the application of microcomputed tomography. This technique, in particular, has made it possible to dissect small insects digitally, document anatomy efficiently and in detail, and visualize these data three dimensionally. In this study, we document the skeletomuscular system of a cuckoo bee, Thyreus albomaculatus and, with that, we provide a 3D atlas of bee skeletomuscular anatomy. The results obtained for Thyreus are compared with representatives of two other bee families (Andrenidae and Halictidae), to evaluate the generality of our morphological conclusions. Besides documenting 199 specific muscles in terms of origin, insertion, and structure, we update the interpretation of complex homologies in the maxillolabial complex of bee mouthparts. We also clarify the complicated 3D structure of the cephalic endoskeleton, identifying the tentorial, hypostomal, and postgenal structures and their connecting regions. We describe the anatomy of the medial elevator muscles of the head, precisely identifying their origins and insertions as well as their homologs in other groups of Hymenoptera. We reject the hypothesis that the synapomorphic propodeal triangle of Apoidea is homologous with the metapostnotum, and instead recognize that this is a modification of the third phragma. We recognize two previously undocumented metasomal muscle groups in bees, clarifying the serial skeletomusculature of the metasoma and revealing shortcomings of Snodgrass' "internal-external" terminological system for the abdomen. Finally, we elucidate the muscular structure of the sting apparatus, resolving previously unclear interpretations. The work conducted herein not only provides new insights into bee morphology but also represents a source for future phenomic research on Hymenoptera., (© 2024 The Author(s). Journal of Morphology published by Wiley Periodicals LLC.)

The shift to a pollen diet and the evolution of more highly organized societies, i.e., eusocial, were key milestones in bee diversification over their evolutionary history, culminating in a high dependence on feeding broods with a large variety of floral resources. Here, we hypothesized that obligatory eusocial bees have a wider diet diversity than their relatives with solitary lifestyles, and this would be related to colony size. To test both hypotheses, we surveyed diet breadth data (palynological analysis) based on the Shannon-Wiener index (H') for 85 bee taxa. We also obtained colony size for 47 eusocial bee species. These data were examined using phylogenetic comparative methods. The results support the generalist strategy as a derived trait for the bee taxa evaluated here. The dietary diversity of eusocial bees (H': 2.1, on average) was 67.5% higher than that of noneusocial bees (H': 1.21, on average). There was, however, no relationship between diet breadth and colony size, indicating that smaller colonies can harvest a pollen variety as diverse as larger colonies. Taken together, these results provide new insights into the impact of lifestyle on the diversity of collected pollen. Furthermore, this work sheds light on an advantage of living in more highly structured societies irrespective of the size of the colony., (© The Author(s) 2024. Published by Oxford University Press on behalf of Entomological Society of America.)

Fernando A. Silveira had the unique combination of being a sagacious scientist and a remarkable human being. Throughout his career, he made significant contributions to understanding bee diversity and keenly spread this scientific information to the academic community at large and beyond the university walls. His rich character, warm heart, strong voice and laughter are missed by those privileged to be Fernandos students, friends, mentors, and family. In this volume, we honor Fernando A. Silveira, who prematurely passed away at the age of 62, leaving three sons, his wife, and numerous friends.

The genus Actenosigynes includes two species, A. fulvoniger (Michener, 1989) and A. mantiqueirensis Silveira, 2009, both oligolectic on flowers of Blumenbachia (Loasaceae) in southern Brazil. We describe a third species, Actenosigynes silveirai Siriani-Oliveira, sp. n., and provide additional evidence to the suspected narrow host-plant specificity between bees of this genus and Loasaceae. This new species was only recorded to collect resources on flowers of Aosa, a genus closely related to Blumenbachia in the subfamily Loasoideae. We illustrate female and male specimens of the three species to offer a complete summary of the morphological variation within this modestly sized genus of Neopasiphaeinae, including photographs of male genitalia and associated metasomal sterna. Moreover, we provide an identification key for the three species of Actenosigynes and the first phylogenetic and dating estimate for these taxa. The genus diversified in southern South America during the Miocene-Pliocene, following a more ancient divergence associated with the orogenic events that separated its sister-genus, Torocolletes, west of the Andes. We dedicate this newly described species to Fernando A. Silveira for his contributions to research on Brazilian bee taxonomy and biology.

Competition is an important biological filter that can define crucial features of species' natural history, like survival and reproduction success. We evaluated in the Brazilian tropical savanna whether two sympatric and congenereric species, Qualea multiflora Mart. and Q. parviflora Mart. (Vochysiaceae), compete for pollinator services, testing whether there is a better competitor or whether plants present any anti-competitive mechanism. Additionally, we investigated the breeding system, pollinators, and flowering phenology of both species. The results showed that Q. multiflora and Q. parviflora are dependent on pollinators for fruit formation, as they exhibited a self-incompatible and non-agamospermic breeding system. These plants shared the same guild of pollinators, which was formed by bees and hummingbirds, and an overlap in the flower visitation time was observed. Each plant species had different pollinator attraction strategies: Q. multiflora invested in floral resource quality, while Q. parviflora invested in resource quantity. The blooming time showed a temporal flowering partition, with highly sequential flowering and no overlap. Qualea parviflora bloomed intensely from September to October, while Q. multiflora bloomed from November to January, with the flowering peak occurring in December. The two Qualea species have morphologically similar flowers, are sympatric, and share the same pollinator community, with overlapping foraging activity during the day. However, they do not compete for pollinator services as they exhibit an anti-competitive mechanism mediated by temporal flowering partition.

Bees are the most significant pollinators of flowering plants. This partnership began ca. 120 million years ago, but the uncertainty of how and when bees spread across the planet has greatly obscured investigations of this key mutualism. We present a novel analysis of bee biogeography using extensive new genomic and fossil data to demonstrate that bees originated in Western Gondwana (Africa and South America). Bees likely originated in the Early Cretaceous, shortly before the breakup of Western Gondwana, and the early evolution of any major bee lineage is associated with either the South American or African land masses. Subsequently, bees colonized northern continents via a complex history of vicariance and dispersal. The notable early absences from large landmasses, particularly in Australia and India, have important implications for understanding the assembly of local floras and diverse modes of pollination. How bees spread around the world from their hypothesized Southern Hemisphere origin parallels the histories of numerous flowering plant clades, providing an essential step to studying the evolution of angiosperm pollination syndromes in space and time., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 Elsevier Inc. All rights reserved.)

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., (© 2023 Willi Hennig Society.)

The diversity of the Elampini cuckoo wasps in northeastern Brazil is reviewed. Three new species are described: Hedychrum oxente Lucena & Zanella sp. nov., Holopyga lunae Lucena sp. nov., and Muesebeckidium clemensi Lucena & Zanella sp. nov. A lectotype is designated for Holopyga piliventris Ducke, 1907 and herein illustrated. Elampus aequinoctialis Ducke, 1901 is restored as a valid species (previously synonymized with Elampus gayi Spinola, 1851) and diagnosed. Holophris huberi (Ducke, 1901) and Muesebeckidium clemensi sp. nov. represent the first records of both genera to northeastern Brazil. New records along with up-to-date distributional maps are discussed, and an identification key to the species of Holopyga from northeastern Brazil is provided. The total diversity of Elampini recorded for northeastern Brazil is now represented by the following eight species: Elampus aequinoctialis Ducke, 1901, Exallopyga guatemalensis (Cameron, 1888), Hedychrum oxente Lucena & Zanella sp. nov., Holophris huberi (Ducke, 1901), Holopyga iheringi du Buysson, 1901, H. lunae Lucena sp. nov., H. piliventris Ducke, 1907, and Muesebeckidium clemensi Lucena & Zanella sp. nov.

Ribosomal DNA (rDNA) loci are essential for cellular metabolism due to their participation in ribosome biogenesis. Although these genes have been widely cytogenetically mapped, the evolutionary mechanisms behind their variability in number and chromosomal location remain elusive, even in well-known biological groups, such as ants, bees and wasps (Insecta: Hymenoptera). To address this question in Hymenoptera and therefore advance the understanding of rDNA evolution in insects in general, we integrated molecular cytogenetic data, a phylogenomic framework, model-based predictions and genome sequencing. Hence, we assessed the main evolutionary trends shaping the chromosomal distribution of rDNA loci in Hymenoptera. We noticed the conservation of one site of rDNA per haploid genome, suggesting that a single 45S rDNA locus is the putative ancestral pattern for aculeate Hymenoptera. Moreover, our results highlighted a nonrandom distribution of rDNA in Hymenoptera karyotypes, as well as a lineage-specific preferential location. The proximal location of rDNA is favoured in species with multiple loci and in the two families of Hymenoptera that show the highest range of chromosome numbers: Formicidae and Vespidae. We propose that chromosome fissions have played a crucial role in the distribution pattern of rDNA loci through the evolutionary diversification of Hymenoptera. Moreover, our genomic analysis of two species, one with a single locus of rDNA and one with multiple loci, supported that loci multiplication is followed by sequence divergence. Our results provide detailed information about the number and chromosomal position of rDNA in Hymenoptera and, therefore, broaden our knowledge regarding rDNA evolutionary dynamics in insects., (© 2021 European Society for Evolutionary Biology.)

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., (© The Willi Hennig Society 2020.)

Incongruence among phylogenetic results has become a common occurrence in analyses of genome-scale data sets. Incongruence originates from uncertainty in underlying evolutionary processes (e.g., incomplete lineage sorting) and from difficulties in determining the best analytical approaches for each situation. To overcome these difficulties, more studies are needed that identify incongruences and demonstrate practical ways to confidently resolve them. Here, we present results of a phylogenomic study based on the analysis 197 taxa and 2,526 ultraconserved element (UCE) loci. We investigate evolutionary relationships of Eucerinae, a diverse subfamily of apid bees (relatives of honey bees and bumble bees) with >1,200 species. We sampled representatives of all tribes within the group and >80% of genera, including two mysterious South American genera, Chilimalopsis and Teratognatha. Initial analysis of the UCE data revealed two conflicting hypotheses for relationships among tribes. To resolve the incongruence, we tested concatenation and species tree approaches and used a variety of additional strategies including locus filtering, partitioned gene-trees searches, and gene-based topological tests. We show that within-locus partitioning improves gene tree and subsequent species-tree estimation, and that this approach, confidently resolves the incongruence observed in our data set. After exploring our proposed analytical strategy on eucerine bees, we validated its efficacy to resolve hard phylogenetic problems by implementing it on a published UCE data set of Adephaga (Insecta: Coleoptera). Our results provide a robust phylogenetic hypothesis for Eucerinae and demonstrate a practical strategy for resolving incongruence in other phylogenomic data sets., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.)

It is widely recognized that different regions of a genome often have different evolutionary histories and that ignoring this variation when estimating phylogenies can be misleading. However, the extent to which this is also true for morphological data is still largely unknown. Discordance among morphological traits might plausibly arise due to either variable convergent selection pressures or else phenomena such as hemiplasy. Here, we investigate patterns of discordance among 282 morphological characters, which we scored for 50 bee species particularly targeting corbiculate bees, a group that includes the well-known eusocial honeybees and bumblebees. As a starting point for selecting the most meaningful partitions in the data, we grouped characters as morphological modules, highly integrated trait complexes that as a result of developmental constraints or coordinated selection we expect to share an evolutionary history and trajectory. In order to assess conflict and coherence across and within these morphological modules, we used recently developed approaches for computing Bayesian phylogenetic information allied with model comparisons using Bayes factors. We found that despite considerable conflict among morphological complexes, accounting for among-character and among-partition rate variation with individual gamma distributions, rate multipliers, and linked branch lengths can lead to coherent phylogenetic inference using morphological data. We suggest that evaluating information content and dissonance among partitions is a useful step in estimating phylogenies from morphological data, just as it is with molecular data. Furthermore, we argue that adopting emerging approaches for investigating dissonance in genomic datasets may provide new insights into the integration and evolution of anatomical complexes. [Apidae; entropy; morphological modules; phenotypic integration; phylogenetic information.]., (© The Author(s) 2020. Published by Oxford University Press, on behalf of the Society of Systematic Biologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

The present catalogue lists the insect types (Hexapoda) deposited at Coleção Entomológica "Prof. J.M.F.Camargo" (RPSP), Universidade de São Paulo, Ribeirão Preto, Brazil. This collection is known as one of the most significant depositories of stingless bees (Hymenoptera: Apidae: Meliponini) for the Brazilian and Neotropical faunas, largely because of the legacy of Professor João Franco Maria de Camargo (in memoriam) and co-workers. The Meliponini constitute the most emblematic portion of the collection, comprising 3,335 type-specimens (66 holotypes, 3,254 paratypes, and 15 paralectotypes), which represent 132 species-names. In addition to stingless bees, 495 type specimens of other Hymenoptera (23 holotypes and 472 paratypes) are deposited at RPSP representing 71 hymenopteran species-names: Agaonidae (16 names), Andrenidae (3 names), non-Meliponini Apidae (21 names), Chrysididae (2 names), Halictidae (16 names), Colletidae (5 names), Megachilidae (5 names), Melittidae (1 name), Pteromalidae (1 name), and Vespidae (1 name); the only other insect order currently represented by types at RPSP is Diptera (Drosophilidae: 2 names). Altogether, there currently are 3,832 type-specimens deposited at RPSP. Label data and additional information from the RPSP registers, literature sources, and taxonomic remarks are given, including photographs of name bearing types and representatives of other type specimens of the species herein considered.

The pharyngeal plate is a morphological complex with extensive anatomical variation among bees and, therefore, potential as a source of phylogenetic information. The pharyngeal plate of bees is divided into four morphologically distinct regions: sitophore, hypopharyngeal lobe, pharyngeal rods, and median oral plate. In this work we illustrate and document in detail for the first time the pharyngeal plate of 43 bee species, providing descriptions of the morphological variation and contrasting these findings with representatives of apoid wasps (Crabronidae and Sphecidae). We evaluate and discuss the potential of this structure as a rich source of morphological information in the context of bee phylogeny and any research potentially impacted by comparative morphological data. The shape of the hypopharyngeal lobe is highly variable among suprageneric taxa of bees and can be readily employed to characterise taxa at various levels. We argue that the global patterns in the variation of the pharyngeal plate can provide information for phylogenetic inference within bees and constructed and coded 10 characters that encompass the most noticeable morphological differences discussed herein., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The main sources of food for stingless bees are the nectar and pollen harvested from flowers, whereas one important kind of nesting material (i.e. wax) is produced by their own abdominal glands. Stingless bees can, nonetheless, obtain alternative resources of food and wax from exudates released by sap-sucking insects as honeydew and waxy cover, respectively. To date, there are no comprehensive studies investigating how diversified and structured the network interactions between stingless bees and sap-sucking insects are. Here, we conducted a survey of the data on relationship between stingless bees and sap-sucking insects to evaluate: (1) which resources are collected by which stingless bee species; (2) how diverse the interaction network is, using species degree and specialisation index as a proxy; and if (3) there would be any phylogenetic signal in the species degree and specialisation indices. Our findings demonstrate that approximately 21 stingless bee species like Trigona spp. and Oxytrigona spp. have been observed interacting with 11 sap-sucking species, among which Aethalion reticulatum is the main partner. From ca. 50 records, Brazil is the country with most observations (n = 38) of this type of ecological interaction. We found also that stingless bees harvest fivefold more honeydew than waxy covers on sap-sucking insects. However, we did not find any phylogenetic signal for the occurrence of this interaction, considering species degree and specialisation indices, suggesting that both traits apparently evolved independently among stingless bee species. We suggest that specific ecological demands may drive this opportunistic behaviour exhibited by stingless bees, because major sources of food are obtained from flowers and these bees produce their own wax.

Two increasingly popular approaches to reconstruct the Tree of Life involve whole transcriptome sequencing and the target capture of ultraconserved elements (UCEs). Both methods can be used to generate large, multigene datasets for analysis of phylogenetic relationships in non-model organisms. While targeted exon sequencing across divergent lineages is now a standard method, it is still not clear if UCE data can be readily combined with published transcriptomes. In this study, we evaluate the combination of UCEs and transcriptomes in a single analysis using genome-, transcriptome-, and UCE data for 79 bees in the largest and most biologically diverse bee family, Apidae. Using existing tools, we first developed a workflow to assemble phylogenomic data from different sources and produced two large nucleotide matrices of combined data. We then reconstructed the phylogeny of the Apidae using concatenation- and coalescent-based methods, and critically evaluated the resulting phylogenies in the context of previously published genetic, genomic, and morphological data sets. Our estimated phylogenetic trees are robustly supported and largely congruent with previous molecular hypotheses, from deep nodes to shallow species-level phylogenies. Moreover, the combined approach allows us to resolve controversial nodes of the apid Tree of Life, by clarifying the relationships among the genera of orchid bees (Euglossini) and the monophyly of the Centridini. Additionally, we present novel phylogenetic evidence supporting the monophyly of the diverse clade of cleptoparasitic Apidae and the placement of two enigmatic, oil-collecting genera (Ctenoplectra and Tetrapedia). Lastly, we propose a revised classification of the family Apidae that reflects our improved understanding of apid higher-level relationships., (Copyright © 2018 The Authors. Published by Elsevier Inc. All rights reserved.)

Despite the (TTAGG) n telomeric repeat supposed being the ancestral DNA motif of telomeres in insects, it was repeatedly lost within some insect orders. Notably, parasitoid hymenopterans and the social wasp Metapolybia decorata (Gribodo) lack the (TTAGG) n sequence, but in other representatives of Hymenoptera, this motif was noticed, such as different ant species and the honeybee. These findings raise the question of whether the insect telomeric repeat is or not phylogenetically predominant in Hymenoptera. Thus, we evaluated the occurrence of both the (TTAGG) n sequence and the vertebrate telomere sequence (TTAGGG) n using dot-blotting hybridization in 25 aculeate species of Hymenoptera. Our results revealed the absence of (TTAGG) n sequence in all tested species, elevating the number of hymenopteran families lacking this telomeric sequence to 13 out of the 15 tested families so far. The (TTAGGG) n was not observed in any tested species. Based on our data and compiled information, we suggest that the (TTAGG) n sequence was putatively lost in the ancestor of Apocrita with at least two subsequent independent regains (in Formicidae and Apidae).