DOI: 10.3897/asp.83.e166986

Authors: Gideon Pisanty, Rémi Santerre, Teresa Martin, Sophie Cardinal, Thomas J. Wood

Abstract: Andrena (Micrandrena) Ashmead is the largest subgenus of Andrena Fabricius 1775, with more than 150 species distributed throughout the northern hemisphere. The subgenus has gained notoriety as one of the most difficult taxonomic groups of bees, and remains poorly studied across large parts of the Palaearctic region. Based on extensive collecting in Israel, Lebanon and Cyprus, together with efficient DNA barcoding and and re-examination of old museum material, we present a thorough revision of the species of A. (Micrandrena) occuring in the region of the Levant (Israel, the West Bank, Jordan, Lebanon and Syria) and the island of Cyprus, including a detailed identification key, and a summary of our knowledge of the distribution, phenology and foraging biology of each taxon. Our study enumerates forty-two species of Andrena (Micrandrena) from the Levant and Cyprus, including four species new to the region, and nine species new to science: Andrena aphroditae Pisanty sp. nov., A. alshaykh Pisanty sp. nov., A. aspera Pisanty & Wood sp. nov., A. chananaea Pisanty & Wood sp. nov., A. friedmani Pisanty sp. nov., A. hebraica Pisanty & Wood sp. nov., A. kugleri Pisanty sp. nov., A. libanica Wood sp. nov., and A. phoenicia Pisanty sp. nov. We describe the males of A. calandra Warncke, 1975 and A. lindbergella Pittioni, 1950 for the first time. We additionally synonymise Andrena dargia Warncke, 1965 with A. minutula (Kirby, 1802) syn. nov., and recognise two previously subspecific names as valid species: Andrena leptura Warncke, 1974 stat. nov. and A. povolnyi, 1974 Warncke stat. nov.

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DOI: 10.3897/asp.83.e151385

Authors: Bernardo F. Santos, Santiago Bordera

Abstract: Abstract The phylogeny of the Old World genus Stenarella Szépligeti is investigated using data from genomic ultraconserved elements. Thirteen species from the four biogeographic regions where the genus occurs were included in the analyses. The resulting trees corroborate the monophyly of Stenarella and show substantial biogeographic structure within the genus, with three major clades corresponding to species occurring in the Afrotropical, Palearctic and Oriental–Australasian regions. The Afrotropical component of the genus is represented by fifteen species, while outside of the Afrotropics there seem to be only three species, each with very wide geographic distribution. The taxonomic limits of the genus are reviewed, and an updated diagnosis is provided. A total of eighteen species are recognized as valid, of which twelve are described as new: S. aurea sp. nov., S. bugalana sp. nov., S. delicata sp. nov., S. favilla sp. nov., S. fenestralis sp. nov., S. hopkinsi sp. nov., S. katanga sp. nov., S. natalina sp. nov., S. nigriscuta sp. nov., S. nigromaculata sp. nov., S. saaksjarvi sp. nov., and S. vannoorti sp. nov. An updated identification key to the species of the genus is provided, as well as diagnoses, redescriptions, and new distribution records for the previously known species.

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DOI: 10.3897/asp.83.e153226

Authors: Emma Kärrnäs, Christer Hansson, Niklas Wahlberg

Abstract: Abstract Despite their ecological and economic importance as key regulators of insect populations worldwide, eulophid wasps remain one of the taxonomically most neglected groups of insects. As these minute parasitoid wasps exhibit an astonishing species diversity while being challenging to identify, relying on morphological characters alone has long been problematic. By using large-scale barcoding and molecular species delimitation analyses of 909 specimens, of which 641 were of European origin, we reveal a remarkable diversity of cryptic species in the two eulophid wasp genera Oomyzus Rondani and Quadrastichus Girault (Eulophidae, Tetrastichinae) present in Europe. We also present 40 new country records for 17 species and greatly improve the existing DNA barcode libraries of these two groups. Using the four different species delimitation methods ABGD, ASAP, GMYC and mPTP, the number of potentially undescribed European species detected in this study ranges from 20 to 34, of which 16 to 24 are cryptic. These results suggest that not even half of the European Oomyzus and Quadrastichus species are currently known, thus highlighting the need for further taxonomic work. However, the results also highlight issues with molecular species delimitations based on barcode data, such as incongruence across different analysis methods and potential barcode sharing between closely related species. Therefore, the results also challenge the validity of relying on a single molecular marker when detecting and subsequently describing new species, an increasing practise within modern taxonomy termed “turbo-taxonomy”.

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DOI: 10.3897/asp.83.e155906

Authors: Sixun Ge, Tai Gao, Yizhou Liu, Jiale Li, Juan Shi, Lili Ren, Hongliang Shi

Abstract: Abstract Sirex wood wasps (Hymenoptera: Siricidae) constitute a group of economically significant xylophagous pests that cause substantial damage to forestry globally. Herein, we present a systematic revision of the Chinese Sirex species. Through phylogenetic analyses, some taxonomic ambiguities were resolved. Sirex chimera sp. nov. is described (type locality: Chifeng, Inner Mongolia). Two new synonyms are proposed: S. rufiabdominis Xiao & Wu, 1983 and S. sinicus Maa, 1949 are synonymized with S. nitobei. S. dux (Semenov, 1921) stat. rev. is confirmed as a distinct species rather than a subspecies of S. juvencus or S. cyaneus, with the male described for the first time. Re-examination confirms that previous Chinese records of S. imperialis represent misidentifications of S. nitobei. The Chinese Sirex fauna is revised to comprise nine valid species: S. chimera sp. nov., S. dux, S. ermak, S. juvencus, S. nitobei, S. noctilio, S. piceus, S. tianshanicus, and S. vates. Comprehensive re-descriptions and a key to species are provided.

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DOI: 10.3897/asp.82.e106734

Authors: André L. Martins, Gabriel A. R. Melo

Abstract: Thaumatodryininae is a small subfamily of Dryinidae, known to attack nymphs of auchenorrhynchous Flatidae (Hemiptera). Only one genus is recognized, Thaumatodryinus Perkins, with 35 species including fossil and extant taxa. Currently, the oldest record for the genus is from Baltic amber. Here, we present the first record of Thaumatodryininae from mid-Cretaceous Burmese amber with the description of †Thaumatorrhinos athrix gen. et sp. nov., derived from the first phylogeny for this subfamily based on morphological characters. The placement of †Thaumatorrhinos gen. nov. in Thaumatodryininae and the phylogenetic relationships of this subfamily within Dryinidae are discussed.

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DOI: 10.3897/asp.82.e111148

Authors: Manuel Brazidec, Lars Vilhelmsen, Brendon E. Boudinot, Adrian Richter, Jörg U. Hammel, Evgeny E. Perkovsky, Yong Fan, Zhen Wang, Qiong Wu, Bo Wang, Vincent Perrichot

Abstract: The Megalyridae are a small family of parasitoid wasps comprising eight extant genera (71 species) and six extinct genera (13 species). Here, we report eight new species from Late Cretaceous Kachin (Myanmar) and Taimyr (Russia) ambers; the family is recorded for the first time from the latter. †Cretolyra noijebumensis gen. et sp. nov., †Cretolyra shawi gen. et sp. nov., †Genkyhag innebula gen. et sp. nov., †Megacoxa chandrahrasa gen. et sp. nov., †Megacoxa janzeni gen. et sp. nov., and †Megacoxa synchrotron gen. et sp. nov., are described from late Albian – early Cenomanian Kachin amber; †Kamyristi exfrigore gen. et sp. nov. and †Kamyristi yantardakhensis gen. et sp. nov. from Taimyr amber (Baikura, late Albian – early Cenomanian Ognevka Formation and Yantardakh, Santonian Kheta Formation, respectively). Phylogenetic analyses of the family are presented and the classification of the Megalyridae is revised accordingly. A new tribe, †Megalavini trib. nov., is erected to accommodate †Cretolyra gen. nov., †Genkyhag gen. nov., †Megacoxa gen. nov. together with †Megalava Perrichot, 2009; †Megalavini trib. nov. is characterized by the vein M fully pigmented and veins Rs+M and M+Cu aligned, the crenulate mesometapectal sulcus, and the pronotal spiracle not posteriorly surrounded by cuticle. This spiracular condition was previously only observed in †Megazar Perrichot, 2009, thus the subfamily †Megazarinae Perrichot, 2009 stat. nov. is proposed for the clade (†Megalavini + †Megazar), as sister to the remaining Megalyridae. The latter is defined as the Megalyrinae, with †Kamyristi gen. nov. being retrieved as sister to all other genera except Rigel Shaw, 1987 under parsimony analyses. To align phylogeny with classification, three new tribes are erected (†Kamyristini trib. nov., †Megallicini trib. nov., and †Valaaini trib. nov.) and the †Cretodinapsini is synonymized under Megalyrini. A grouping [Rigel + Megalyridia] is supported under Bayesian analyses, which is the only specific conflict with the parsimony analyses, suggesting some degree of caution with respect to the internal relationships of the Megalyrinae. Finally, a revised key to the Megalyridae genera is provided.

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DOI: 10.3897/asp.81.e104810

Authors: Zachary H. Griebenow, Adrian Richter, Thomas van de Kamp, Evan P. Economo, Ziv E. Lieberman

Abstract: The male genitalia of the Insecta are famed for structural and functional diversity. Variation in this anatomical region shows ample phylogenetic signal, and this variation has proven indispensable for classification across the insects at multiple taxonomic ranks. However, in the ants (Hymenoptera: Formicidae) the male genital phenotype is ancillary to the morphology of the worker caste for systematic purposes. Ants of the enigmatic subfamily Leptanillinae are an exception, as males are easier to collect than workers. Ongoing systematic revision of the Leptanillinae must therefore rely upon the male phenotype – particularly the spectacular morphological profusion of the male genitalia. To thoroughly illuminate this anatomical region and aid comparative morphological research on ant male genitalia, we present a comparative morphological study of the male genitalia in nine exemplar lineages spanning the Leptanillinae, plus three outgroups representing other major clades of the Formicidae. We use micro-computed tomography (micro-CT) to generate 3D volumetric reconstructions of male genital skeletomusculature in these specimens. Our descriptions use new muscular terminology compatible with topographic main-group systems for the rest of the pterygote soma, and applicable to all Hymenoptera. We find that male genitalia in the Leptanillinae show an overall trend towards skeletomuscular simplification, with muscular reduction in some cases being unprecedented in ants, or even hymenopterans in general. In several lineages of the Leptanillinae we describe derivations of the male genitalia that are bizarre and unparalleled among the Hymenoptera. We conclude by discussing the functional implications of the often-extreme morphologies here observed.

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DOI: 10.3897/asp.81.e107579

Authors: Si-Xun Ge, Zhuo-Heng Jiang, Li-Li Ren, Cornelis van Achterberg, Jiang-Li Tan

Abstract: The family Stephanidae (Hymenoptera) constitutes a unique group within the Apocrita, playing a pivotal role in the evolution of parasitoid wasps. Although the phylogeny of Stephanidae has been previously inferred, it remains at a low resolution when considering both extinct and extant genera, as well as the enigmatic extinct genus †Electrostephanus. Here, we undertake a revision of Stephanidae extinct, presenting descriptions of new specimens from late Cretaceous Burmese amber and early Eocene Baltic amber. Combining all extant and extinct genera, the phylogeny of Stephanidae was analyzed, incorporating 57 species within 21 genera based on 64 morphological characters. We apply both under maximum parsimony with equal weighting and implied weighting methods, with four species representing early Apocrita as outgroups. Divergence times are estimated by utilizing extinct taxa as calibration points. A new basal subfamily of stephanid wasp, †Lagenostephaninae subf. nov. was established, encompassing †Lagenostephanus and the newly described genera †Tumidistephanus gen. nov and †Neurastephanus gen. nov. The genus †Electrostephanus is redefined, with two species assigned under distinct genera, †Neurastephanus gen. nov. and †Aphanostephanus gen. nov.. We discuss some of the putative morphological synapomorphies of evolutionary significance within the phylogenetic framework. Our results complement several characteristics of great taxonomic importance for Stephanidae and provide new insights into the early evolution of the family.

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DOI: 10.3897/asp.81.e96737

Authors: Manuel Brazidec, Frédéric Legendre, Vincent Perrichot

Abstract: The Lancepyrinae are an extinct subfamily of Bethylidae known exclusively from Cretaceous amber deposits of Lebanon, Spain, Taimyr and Myanmar. In this study, we describe and illustrate four new genera and five new species of lancepyrine wasps from the Albian of Hkamti and late Albian-early Cenomanian of Kachin (Myanmar): Azepyris delamarrei gen. et sp. nov., Burmapyris ohmkuhnlei sp. nov., Gwesped groehni gen. et sp. nov., Paralanceis chotardi gen. et sp. nov. and Yunbayin rossei gen. et sp. nov. These taxa not only highlight the taxonomic diversity of the Lancepyrinae during the mid-Cretaceous but they also reveal the morphological disparity of the subfamily. To establish the phylogenetic relationships of these fossils and to check the monophyly of the Lancepyrinae, we add them to a pre-existing morphological matrix and perform a cladistic analysis. We retrieve the subfamily as poorly supported yet monophyletic, with the newly described taxa deeply nested in it. A key to the genera of Lancepyrinae is provided. Finally, we erect the subfamily Cretabythinae subfam. nov. for the genera Cretabythus Evans, 1973, Holopsenelliscus Engel, 2019 and Megalopsenella Jouault et al., 2020 as no taxonomic treatment has been provided for these taxa after the transfer of Holopsenella Engel et al., 2016 as Aculeata incertae sedis.

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DOI: 10.3897/asp.80.e84428

Authors: Enrico Schifani, Antonio Alicata, Mattia Menchetti, Lech Borowiec, Brian L. Fisher, Celal Karaman, Kadri Kiran, Wala Oueslati, Sebastian Salata, Rumsaïs Blatrix

Abstract: The West-Palearctic region is a diversity hotspot for the ant genus Aphaenogaster. Species in this region are characterized by high morphological variation, which has led to their subdivisioninto different infrageneric groups. The very first classification in three subgenera, dated 1915, was gradually replaced by eight species-groups. To probe the evolutionary consistency of these species-groups, we sequenced 46 species from all eight species-groups and biogeographic sectors of the region, using one mitochondrial (COI) and six nuclear markers (EPICs), and interpreted the results by integrating qualitative morphology. Our results demonstrate the non-monophyly of all formerly recognized subgenera and species-groups, except for the crocea group. We use the phylogeny and morphological characters to propose a new classification of six monophyletic species-groups (crocea, gibbosa, graeca, pallida, sardoa, subterranea). The pallida, subterranea and sardoa (formerly testaceopilosa) groups attain monophyletic status by reassigning a few taxa. The gibbosa group is to be considered exclusively Western-Mediterranean until further assessments of similar Eastern species. The new graeca group is established by including former members of the splendida and subterranea groups, while the polyphyletic cecconii, obsidiana, and splendida groups are dismissed. Notably, the first is not part of the tropical Deromyrma clade as previously thought, while at least two independent clades which require further investigation are composed of species from both the cecconii and splendida groups, suggesting repeated morphological convergences based on similar ecological adaptations. Finally, A. cardenai is confirmed to be a significantly divergent lineage. In addition, three Aphaenogaster species are moved to different genera: Messor asmaae (Sharaf, 2018) comb. nov., Messor isekram (Bernard, 1977) comb. nov., and Pheidole sarae (Sharaf, 2018) comb. nov. Further studies should address the evolutionary relationships between the clades recovered in this study.

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DOI: 10.3897/asp.80.e71943

Authors: Anderson Lepeco, Rodrigo Barbosa Gonçalves

Abstract: Augochlora Smith, with 127 valid species, is the most widespread genus of Augochlorini bees, ranging from Argentina to southern Canada, including the Caribbean islands. The genus is divided into three subgenera, Augochlora s. str., Oxystoglossella Eickwort, and the fossil Electraugochlora Engel. The extant subgenera were traditionally diagnosed by their nesting substrate, social behavior and morphology. However, accumulating evidence suggests that these features are not reliable for their separation, especially with the discovery of an enigmatic species sharing characteristics from both subgenera. Our objective is to provide a phylogenetic hypothesis to evaluate the monophyly of the extant subgenera and to place a new species, Augochlora (Augochlora) intermedia sp. nov. For this purpose, we compiled 110 unordered characters for 40 species of Augochlora plus seven outgroup species and analyzed under parsimony and Bayesian inference. Topologies were very similar under both frameworks, allowing us to consistently characterize a few major lineages. Our results demonstrate that the extant subgenera correspond to monophyletic groups and the new species is sister group to remaining Augochlora s. str. species. Both subgenera are widespread in the Western Hemisphere, with species groups differing in range and distributional patterns. Our interpretation is that Augochlora arose in South America, subsequently colonizing Mesoamerica, the Caribbean and North America several times. Facultative social behavior can be found in both subgenera and in most lineages, indicating that the exclusive solitary behavior found in Augochlora pura is an exception. Based on morphological clues we interpret that the habit of nesting out of the soil arose once in Augochlora s. str.

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DOI: 10.3897/asp.80.e71492

Authors: Paulo Cézar Salgado Barroso, Rodolpho Santos Telles Menezes, Marcio Luiz de Oliveira, Alexandre Somavilla

Abstract: For the Neotropical genus Angiopolybia Araujo 1946, several phenotypic forms were previously described, however, they have not been studied within an integrative taxonomic framework. Here, we used molecular data (variation of two mitochondrial genetic markers with molecular species delimitation methods) and morphology (adult morphology, male genitalia, and scanning electron microscopy images) to test the number of species within Angiopolybia. Specifically, we investigated the taxonomic validity of the morphological variants A. pallens dark morph, A. paraensis morph paraensis, A. paraensis morph ruficornis, and A. paraensis morph obscurior. Moreover, we reviewed the taxonomy and geographic distribution of the genus. Our results of morphological and molecular analyses are compatible with the current classification of Angiopolybia, and we did not find reasons to propose the morphological variants of A. pallens and A. paraensis as valid species. Additionally, we reassess the spatial range of the four Angiopolybia species and provide refined maps of their geographical distributions.

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DOI: 10.3897/asp.79.e66978

Authors: Philip S. Ward, Brendon E. Boudinot

Abstract: Camponotus and Colobopsis are widely distributed and species-rich genera in the ant tribe Camponotini. Molecular phylogenetic studies demonstrate that they are not sister taxa, but several lineages within each genus have converged to a remarkable degree, confounding the taxonomy of these ants. Based on multiple lines of evidence, including worker and male morphology, we demonstrate that: (1) three species of “Camponotus” belonging to the subgenus Myrmotemnus, including its type species, are in fact members of the genus Colobopsis; (2) four species previously assigned to Colobopsis belong to the subgenus Myrmamblys of Camponotus; and (3) three Nearctic taxa recently placed in Colobopsis are members of the genus Camponotus and closely related to Camponotus clarithorax. These taxonomic findings yield the following new or revived combinations: Colobopsis moeschi (comb. nov.), Colobopsis moeschi lygaea (comb. nov.), Colobopsis nutans (comb. nov.), Colobopsis nutans cleliae (comb. nov.), and Colobopsis reichenspergeri (comb. nov.); Camponotus apostemata (comb. nov.), Camponotus aurelianus (comb. rev.), Camponotus cavibregma (comb. nov.), Camponotus horrens (comb. rev.), Camponotus politae (comb. rev.), Camponotus trajanus (comb. rev.), and Camponotus yogi (comb. rev.). A further consequence is the following generic synonymy (senior synonym listed first): Colobopsis = Myrmotemnus syn. nov., and Camponotus = Dolophra syn. rev. At the species level, we argue that Camponotus apostemata and Camponotus cavibregma are junior synonyms (syn. nov.) of Camponotus yogi, and Camponotus quercicola is a junior synonym (syn. nov.) of Ca. laevigatus. Taxonomic comments are also provided on some members of the Camponotus reticulatus group, with Camponotus adustus (stat. nov.) and Ca. leucodiscus (stat. rev.) being recognized as distinct species rather than subspecies of Ca. bellus. A male-based diagnosis of the Camponotini is provided, and differences between the males of Colobopsis and Camponotus are documented and illustrated for the first time. This study reveals new character systems of potential value to the systematics of these ants, including features of the male genitalia, and emphasizes the value of reciprocal illumination between phylogenomics and critical morphological analysis.

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DOI: 10.3897/asp.79.e64304

Authors: Alexandre Somavilla, Paulo C. S. Barroso, Marcos Aragão, Sidnei Mateus, Rodolpho S. T. Menezes

Abstract: Phenotypic characters are traditionally the main information for species discrimination in taxonomic studies of invertebrates. However, the presence of inter- and intraspecific polymorphism makes it difficult to identify species in many groups such as Neotropical social wasps. Herein, we examined different sources of biological information such as adult morphology, male genitalia, nest architecture, and genetic data applying an integrative taxonomic approach to study pinned museum specimens belonging to the social wasp genus Pseudopolybia de Saussure, 1863. Based on multiple independent lines of evidence, we described a new Neotropical swarm-founding social wasp, Pseudopolybia cryptica sp. n. Moreover, we proposed a phylogenetic hypothesis for Pseudopolybia including this new species. Our taxonomic findings applying an integrative approach reinforce that the social wasp diversity in the Neotropics may be underestimated due to morphological similarity.

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