JHR 85: 119-133 (2021) ge, JOURNAL OF eet nnscnn in doi: 10.3897/jhr.85.68658 RESEARCH ARTICLE () Hymenopter a 4 https://jhr.pensoft.net The Inarasional Society of ymenopeeriss, RESEARCH A new genus and species of Pristocerinae (Hymenoptera, Bethylidae) from upper Eocene Baltic amber with a review of conspecific association from insect fossils Carly Melissa Tribull', Madeline V. Pankowski*, Wesley Dondoni Colombo? | Farmingdale State College (State University of New York), Biology Department, Farmingdale NY, USA 2 Rockville, Maryland, USA 3 Universidade Federal do Espirito Santo, Departamento de Ciéncias Biolégicas, Vitoria ES, Brazil Corresponding author: Carly Melissa Tribull (cmtribull@gmail.com) Academic editor: Michael Ohl | Received 14 May 2021 | Accepted 15 June 2021 | Published 31 August 2021 http://zoobank.org/9D5 BOOCA-75F0-4E 1F-B847-11D490F769CD Citation: Tribull CM, Pankowski MV, Colombo WD (2021) A new genus and species of Pristocerinae (Hymenoptera, Bethylidae) from upper Eocene Baltic amber with a review of conspecific association from insect fossils. Journal of Hymenoptera Research 85: 119-133. https://doi.org/10.3897/jhr.85.68658 Abstract A new extinct genus and species of Pristocerinae, tArcheonesia eocena Tribull, Pankowski & Colombo, gen. et. sp. nov., are described from upper Eocene Baltic amber from the Yantarny amber mine in the Kaliningrad region, Russia. Descriptions, remarks, illustrations, and comparisons to all extinct and extant Pristocerinae are provided. tArcheonesia is described as a new genus because neither the male nor the female can be placed in any previously described genera, although the female is most similar to Acrenesia and the male is most similar to Cleistepyris. Rare for Bethylidae, and Hymenoptera in general, this fossil contains both a male and female specimen that we are describing as conspecifics. A brief review from the paleoentomological literature is provided to describe how insect fossils containing evidence of reproduc- tive behavior have been used to associate conspecifics in extinct species. Keywords Amber, Bethylidae, Eocene, fossil, Pristocerinae Copyright Carly Melissa Tribull et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 120 Carly Melissa Tribull et al. / Journal of Hymenoptera Research 85: 119-133 (2021) Introduction Bethylidae are a family of parasitoid wasps within the aculeate superfamily Chrysi- doidea that have a cosmopolitan distribution and are known for attacking lepidopteran and coleopteran hosts, including agricultural pests like the navel orangeworm, pink bollworm, and coffee berry borer (Gordh et al. 1983; Abraham et al. 1990; Azevedo et al. 2018). Currently, the family contains nearly 3,000 species in about 100 genera within five extant subfamilies (Bethylinae, Epyrinae, Mesitiinae, Pristocerinae, and Scleroderminae) and four extinct subfamilies (fElektroepyrinae, {Lancepyrinae, tHol- opsenellinae, and *Protopristocerinae) (Azevedo et al. 2018; Colombo et al. 2020a, b). There are about 90 flat wasps fossil species described, with the oldest known extinct Bethylidae coming from early Cretaceous amber deposits (Engel et al. 2016). How- ever, the greatest number of fossil bethylids comes from the Eocene, specifically Oise, Rovno, and Baltic amber deposits (Colombo et al. 2020a, 2021). With over 1,000 species in 26 genera, Pristocerinae are the most speciose subfam- ily within Bethylidae and are found worldwide (Azevedo et al. 2018; Colombo et al. 2020a, 2021). The subfamily is known for its remarkable sexual dimorphism, with males possessing robust bodies, wings, and conspicuous eyes and ocelli while females are wingless, lack ocelli, and have eyes that are extremely reduced or missing (Alencar et al. 2018). The vast majority of Pristocerinae species are known from ‘males-only’ or ‘females-only and conspecific associations between males and females are rare, typi- cally the result of collecting specimens copulating or rearing them from the same host (Azevedo et al. 2016; Alencar et al. 2018; Chen and Azevedo 2020). There are 15 extinct species of Pristocerinae in 10 genera, with the oldest species, }Foenobethylus electriphilus (Cockerell) known from mid-Cretaceous Burmese amber (Cockerell 1920; Faliéres and Nel 2019, 2020; Colombo et al. 2020a). Of these species, 11 are known from males only, three are known from females only, and only one spe- cies, Pristocera skwarrae (Brues), was described from a fossil that contained both a male and female. Additionally, Brues (1933) does not include any illustrations of the species and it was presumed to be lost during World War Hf (Colombo et al. 2021). About 50% of extinct Bethylidae come from Baltic amber deposits dated to the late Eocene, including four of the 15 extinct Pristocerinae from the genera Pseud- isobrachium Kieffer, Cleistepyris Kieffer, and Pristocera Klug (Colombo and Azevedo 2019; Colombo et al. 2021). Here we present a new extinct genus of Pristocerinae from Baltic amber, tArcheonesia Tribull, Pankowski & Colombo, gen. nov. with the type species TA. eocena Tribull, Pankowski & Colombo, sp. nov. Like the lost Pristocera skwarrae, this amber fossil shows both a male and female together, granting us the rare opportunity to describe both sexes for a new extinct bethylid genus. Materials and methods The specimens are embedded in Baltic amber sourced from the Yantarny mine in the Kaliningrad region. The piece was acquired from, trimmed, and polished by Marius A new genus and species of Pristocerinae 121 Veta and has clear dorsal and ventral views, although the lateral views are obscured by bubbles. The type material is deposited in the American Museum of Natural His- tory, USA (AMNH, curator: David Grimaldi) with the specimen catalogue number AMNH_IZC 00361788. The specimens were studied with an Olympus SZX-10 stereomicroscope and pho- tomicrographs were acquired with a DP27 digital camera, using Olympus’s Cellsens software. Multiple Z-stacks were compiled using Helicon Focus. The drawings were scanned and vectorized into Adobe Illustrator CS6 version 23.0.3, and images were edited and combined into a single plate using Adobe Photoshop CC. Terminology for the integument and sculpturing follows Harris (1979) and gen- eral terms follow Lanes et al. (2020). Systematic palaeontology Family Bethylidae Haliday, 1839 Subfamily Pristocerinae Mocsary, 1881 Genus }Archeonesia Tribull, Pankowski & Colombo, gen. nov. http://zoobank.org/6764DE3C-095F-4067-B96D-79706FF609C5 Type-species. {A. cocena Tribull, Pankowski & Colombo, sp. nov. by original designation. Description. Male (Figs 1-4). Head, pronotum, mesoscutum, metapectal-prop- odeal complex, petiole, antenna, and metasoma dark castaneous to black; wings hya- line. Head as long as wide and subquadrate, not globoid in lateral view. Clypeus with triangular median lobe, visible dorsally, lateral lobe reduced. Median clypeal carina delimited, lower than frons. Flagellomere longer than wide, with first flagellomeres larger than distal ones; flagellar pubescence erected; pedicel shorter than flagellomere I, apex dilated. Eye located touching mandibular base, glabrous, bulging. Frons weakly coriaceous, punctures large and sparse. Frontal line not visible. Ocellus large, salient. Frontal angle of ocellar triangle in obtuse angle. Anterior ocellus posterior to supra- ocular line. Occipital carina present. Dorsal pronotal area wider than long, weakly coriaceous, punctures small and sparse. Metafurcal pit oval. Posterior mesofurcal pit crown-shaped. Notaulus present, large, converging posteriorly, smooth. Parapsidal sig- num shorter than notauli. Forewing with three cells closed (C, R, 1Cu), distal flexion line visible, 2r-rs&Rs vein tubular, long, well pigmented, angled, not converging pos- teriorly to anterior margin, R1 vein tubular, long. Pterostigma enlarged, lanceolate. Mesoscutum-mesoscutellar sulcus present, posterior margin strongly incurved medi- ally. Metanotum well-developed medially. Metapectal-propodeal disc not visible. Mes- otibia without spines. Metasoma polished. Ninth abdominal segment with margin weakly incurved, undivided. Female (Figs 1-3, 5). Head, pronotum, mesoscutum, metapectal-propodeal com- plex, petiole, antenna, and metasoma light to dark brown. Head longer than wide and rectangular, not globoid in lateral view. Clypeus with median lobe trapezoidal, 122 Carly Melissa Tribull et al. / Journal of Hymenoptera Research 85: 119-133 (2021) ee al se @ , a “oe a an o Figure |. {Archeonesia eocena Tribull, Pankowski & Colombo, gen. et. sp. nov. A dorsal habitus of male holotype and female allotype B ventral habitus of male holotype and female allotype. Scale bars: 1 mm (A, B). x S35 CX x OOO PO > os