A new epyrine wasp (Hymenoptera: Bethylidae) from the late Eocene of Europe
Abstract
The aculeate family Bethylidae is represented in Baltic amber by about 40 species. We here describe and figure a new species in the subfamily Epyrinae from this late Eocene deposit: †Gracilepyris carsteni sp. nov., based on a complete male specimen. This is the first fossil species of Gracilepyris Colombo, Tribull & Azevedo, 2022, a recently discovered genus, comprising five extant species from the northern Neotropical, Nearctic, and Palaearctic regions. Gracilepyris is characteristic of the Epyris complex, a group of genera possessing individualized foveae at the mesoscuto-mesoscutellar suture. The finding of †Gracilepyris carsteni sp. nov. in Baltic amber is consistent with the estimated Late Cretaceous–Paleogene radiation of this derived group. Finally, we revive the name Risepyris dubius (Kieffer, 1904) for an extant European bethylid species which was unnecessarily renamed Holepyris neodubius Azevedo, 2018; we also propose the new name †Risepyris bruesi Brazidec nom. nov. for the Eocene fossil †Risepyris dubius (Brues, 1933), which was objectively invalid as a junior homonym.
References
Aleksandrova G.N. & Zaprozhets N.I. 2008. Palynological characteristics of Upper Cretaceous and Paleogene deposits on the east of the Sambian Peninsula (Kaliningrad region), Part. 2. Stratigraphy and Geological Correlation 16 (5): 528–539. https://doi.org/10.1134/S0869593808050067
Amante M., Russo A., Schöller M. & Steidle J.L.M. 2018. Olfactory host location and host preference of Holepyris sylvanidis (Brèthes) (Hymenoptera: Bethylidae), a parasitoid of Tribolium confusum Jacquelin du Val and T. castaneum (Herbst) (Coleoptera: Tenebrionidae). Journal of Stored Products Research 78: 105–109. https://doi.org/10.1016/j.jspr.2018.06.008
Azevedo C.O. & Azar D. 2012. A new fossil subfamily of Bethylidae (Hymenoptera) from the Early Cretaceous Lebanese amber and its phylogenetic position. Zoologia 29 (3): 210–218. https://doi.org/10.1590/S1984-46702012000300004
Azevedo C.O., Alencar I.D.C.C., Ramos M.S., Barbosa D.N., Colombo W.D., Vargas J.M. & Lim J. 2018. Global guide of the flat wasps (Hymenoptera, Bethylidae). Zootaxa 4489 (1): 1–294. https://doi.org/10.11646/zootaxa.4489.1.1
Barbosa D.N. & Melo G.A.R. 2023. Revision of Bethylinae from Dominican amber, with description of a new genus (Hymenoptera, Bethylidae). Journal of Hymenoptera Research 96: 167–180. https://doi.org/10.3897/jhr.96.100862
Barbosa D.N., Perkovsky E.E. & Azevedo C.O. 2013. Two new species of Laelius Ashmead (Hymenoptera, Bethylidae) from Upper Eocene Rovno amber. Neues Jahrbuch für Geologie und Paläontologie 267 (1): 67–73. https://doi.org/10.1127/0077-7749/2012/0297
Batchelor T.P., Hardy I.C.W. & Barrera J.F. 2006. Interactions among bethylid parasitoid species attacking the coffee berry borer, Hypothenemus hampei (Coleoptera: Scolytidae). Biological Control 36 (1): 106–118. https://doi.org/10.1016/j.biocontrol.2005.07.015
Brazidec M. & Perrichot V. 2022a. The first flat wasps (Hymenoptera: Bethylidae) from Miocene Mexican amber. Journal of South American Earth Sciences 116: e103838. https://doi.org/10.1016/j.jsames.2022.103838
Brazidec M. & Perrichot V. 2022b. A fossil flat wasp (Hymenoptera: Bethylidae) from the early Eocene Green River Formation suggests past cosmopolitan distribution of the genus Eupsenella Westwood, 1874. Palaeoentomology 5 (4): 378–384. https://doi.org/10.11646/palaeoentomology.5.4.10
Brazidec M. & Perrichot V. 2023. The first fossil Parascleroderma (Hymenoptera: Bethylidae): a new species in mid-Miocene Zhangpu amber. Palaeoworld 32 (4): 709–715. https://doi.org/10.1016/j.palwor.2022.01.009
Brazidec M., Lohrmann V. & Perrichot V. 2024. The first Cretaceous epyrine wasp (Hymenoptera: Bethylidae): a new genus and species from Early Cenomanian Kachin amber. Insects 15 (5): 318. https://doi.org/10.3390/insects15050318
Brock R.E., Cini A. & Sumner S. 2021. Ecosystem services provided by aculeate wasps. Biological Reviews 96 (4): 1645–1675. https://doi.org/10.1111/brv.12719
Brues C.T. 1910. The parasitic Hymenoptera of the Tertiary of Florissant, Colorado. Bulletin of the Museum of Comparative Zoology 54: 1–125.
Brues C.T. 1923. Some new fossil parasitic Hymenoptera from Baltic Amber. Proceedings of the American Academy of Arts and Sciences 58 (8): 327–346. https://doi.org/10.2307/20025999
Brues C.T. 1933. The parasitic Hymenoptera of the Baltic Amber. Part I. Bernstein-Forschungen 3: 4–172.
Brues C.T. 1939. New Oligocene Braconidae and Bethylidae from Baltic Amber. Annals of the Entomological Society of America 32 (2): 251–263. https://doi.org/10.1093/aesa/32.2.251
Çan I. 2022. The Turkish Bethylidae (Hymenoptera, Chrysidoidea) fauna, with the new records of Epyrinae and Scleroderminae. Zootaxa 5169 (5): 447–456. https://doi.org/10.11646/zootaxa.5169.5.3
Clauer N., Huggett J.M. & Hillier S. 2005. How reliable is the K-Ar Glauconite chronometer? A case study of Eocene sediments from the Isle of Wight. Clay Minerals 40: 167–176. https://doi.org/10.1180/0009855054020163
Cockerell T.D.A. 1921. Fossil arthropods in the British Museum, V. Oligocene Hymenoptera from the Isle of Wight. Annals and Magazine of Natural History 7 (37): 1–25. https://doi.org/10.1080/00222932108632485
Colombo W.D. & Azevedo C.O. 2021. Synopsis of the fossil Scleroderminae (Hymenoptera, Bethylidae) with description of a new genus and four new species from Baltic amber. Historical Biology 33 (5): 630–638. https://doi.org/10.1080/08912963.2019.1650275
Colombo W.D. & Azevedo C.O. 2023. Discovery of a new Afrotropical genus of flat wasps Ifrika gen. n. (Hymenoptera: Bethylidae). Journal of Natural History 57 (29–32): 1472–1494. https://doi.org/10.1080/00222933.2023.2258605
Colombo W.D., Perkovsky E.E., Waichert C. & Azevedo, C.O. 2021a. Synopsis of the fossil flat wasps Epyrinae (Hymenoptera, Bethylidae), with description of three new genera and ten new species. Journal of Systematic Palaeontology 19 (1): 29–89. https://doi.org/10.1080/14772019.2021.1882593
Colombo W.D., Gobbi F.T., Perkovsky E.E. & Azevedo C.O. 2021b. Synopsis of the fossil Pristocerinae (Hymenoptera, Bethylidae), with description of two new genera and six species from Burmese, Taimyr, Baltic and Rovno ambers. Historical Biology 33 (9): 1736–1752. https://doi.org/10.1080/08912963.2020.1733551
Colombo W.D., Tribull C.M., Waichert C. & Azevedo C.O. 2022. Integrative taxonomy solves taxonomic impasses: a case study from Epyrinae (Hymenoptera, Bethylidae). Systematic Entomology 47 (3): 504–529. https://doi.org/10.1111/syen.12544
Dlussky G.M. & Rasnitsyn A.P. 2009. Ants (Insecta: Vespida: Formicidae) in the upper Eocene amber of central and Eastern Europe. Paleontological Journal 43: 1024–1042. https://doi.org/10.1134/S0031030109090056
Engel M.S. 2001. A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of the American Museum of Natural History 259: 1–192. https://doi.org/fjzbzf
Engel M.S. 2019. A holopsenelline wasp in mid-Cretaceous amber from Myanmar (Hymenoptera: Bethylidae). Palaeoentomology 2 (2): 199–204. https://doi.org/10.11646/palaeoentomology.2.2.10
Evans H.E. 1964. A synopsis of the American Bethylidae (Hymenoptera, Aculeata). Bulletin of the Museum of Comparative Zoology 132: 1–222. Available from https://www.biodiversitylibrary.org/page/4315704 [accessed 7 Apr. 2026].
Falières E. & Nel A. 2018. The oldest representative of the extant genus Epyris in the Lowermost Eocene amber of France (Hymenoptera: Bethylidae: Epyrinae). Palaeoentomology 1 (1) : 32–36. https://doi.org/10.11646/palaeoentomology.1.1.3
Harris R.A. 1979. A glossary of surface sculpturing. Occasional Papers in Entomology 28: 1–31. https://doi.org/10.5281/zenodo.26215
Honsberger D.N., Magnacca K.N., Lorenzo-Elarco H. & Wright M.G. 2024. Life history of two new species of Prorops (Hymenoptera, Bethylidae) ectoparasitic on adult Hypothenemus eruditus beetles (Curculionidae, Scolytinae) in Hawai’i. Journal of Hymenoptera Research 97: 1221–1256. https://doi.org/10.3897/jhr.97.138113
Iakovleva A.I. 2023. Regional Paleogene stratigraphic scheme of Kaliningrad Oblast: state of art, problems and prospects for improvement. Stratigraphy and Geological Correlation 31 (Suppl. 1): S86–S110. https://doi.org/10.1134/S0869593824010064
Iakovleva A.I., Aleksandrova G.N. & Mychko E. 2022. Late Eocene (Priabonian) dinoflagellate cysts from Primorsky Quarry, southeast Baltic coast, Kaliningrad Oblast, Russia. Palynology 46 (2): 1–40. https://doi.org/10.1080/01916122.2021.1980743
Ito R., Tokushige N. & Mita T. 2022. Discovery of the genus Muellerella Saussure, 1892 (Hymenoptera: Bethylidae: Epyrinae) in Japan. Journal of Insect Biodiversity 34 (2): 42–44. https://doi.org/10.12976/jib/2022.34.2.3
Jouault C., Huang D. & Azevedo C.O. 2026. New flat wasps (Hymenoptera: Bethylidae) from the middle Cretaceous Kachin amber of Myanmar. Palaeoworld 35 (3): e201067. https://doi.org/10.1016/j.palwor.2026.201067
Kieffer J.J. 1904. Descriptions de nouveaux Dryinidae et Bethylidae du Musée Civique de Gênes. Annali del Museo Civico di Storia Naturale di Genova 41: 351–412. Available from https://www.biodiversitylibrary.org/page/7930943 [accessed 7 Apr. 2026].
Lanes G.O., Kawada R., Azevedo C.O. & Brothers D.J. 2020. Revisited morphology applied for systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa 4752 (1): 1–127. https://doi.org/10.11646/zootaxa.4752.1.1
Martynova K.V., Perkovsky E.E., Olmi M. & Vasilenko D.V. 2019. New records of Upper Eocene chrysidoid wasps from basins of Styr and Stokhod rivers (Rovno amber). Paleontological Journal 53: 998–1023. https://doi.org/10.1134/S0031030119100125
Meidani Z., Khayrandish M., Azevedo C.O., Madjdzadeh S.M. & Moghaddam M.G. 2024. A peered look at Gracilepyris (Hymenoptera, Bethylidae). Journal of Asia-Pacific Entomology 27 (1): 102177. https://doi.org/10.1016/j.aspen.2023.102177
Ohl M. 1995. Eine neue Bethylidae der Gattung Lytopsenella Kieffer 1911 aus dem baltischen Bernstein (Hymenoptera: Chrysidoidea). Paläontologische Zeitschrift 69: 409–416. https://doi.org/10.1007/BF02987803
Ortega-Blanco J. & Engel M.S. 2013. Bethylidae from Early Cretaceous Spanish amber (Hymenoptera: Chrysidoidea). Journal of the Kansas Entomological Society 86 (3): 264–276. https://doi.org/10.2317/JKES130312.1
Penney D.M. & Jepson J.E. 2014. Fossil Insects: An Introduction to Palaeoentomology. Siri Scientific Press, Manchester.
Perkovsky E.E., Rasnitsyn A.P., Vlaskyn A.P., Taraschuk M.V., 2007. A comparative analysis of the Baltic and Rovno amber arthropod faunas: representative samples. African Invertebrates 48 (1): 229–245. Available from https://hdl.handle.net/10520/EJC84578 [accessed 7 Apr. 2026].
Polaszek A., Almandhari T., Fusu L., Al-Khatri S.A.H., Al Naabi S., Al Shidi R.H., Russel S. & Hardy I.C.W. 2019. Goniozus omanensis (Hymenoptera: Bethylidae) an important parasitoid of the lesser date moth Batrachedra amydraula Meyrick (Lepidoptera: Batrachedridae) in Oman. PLoS ONE 14: e0223761. https://doi.org/10.1371/journal.pone.0223761
Ramos M.S., Perkovsky E.E., Rasnitsyn A.P. & Azevedo C.O. 2014. Revision of Bethylinae fossils (Hymenoptera: Bethylidae) from Baltic, Rovno and Oise amber, with comments on the Tertiary fauna of the subfamily. Neues Jahrbuch für Geologie und Paläontologie 271 (2): 203–228. https://doi.org/10.1127/0077-7749/2014/0385
Ritzkowski S. 1997. K-Ar-Altersbestimmungen der bernsteinführenden Sedimente des Samlandes (Paläogen, Bezirk Kaliningrad). Metalla (Sonderheft) 66: 19–23.
Santos B.F., Colombo W.D., Murray E.A., Brady S.G. & Azevedo C.O. 2024. Insights from the first phylogenomic analysis of flat wasps (Hymenoptera, Bethylidae) reveal two new subfamilies. Cladistics 40 (5): 510–525. https://doi.org/10.1111/cla.12594
Sendel N. 1742. Historia succinorum corpora aliena involventium et naturae opera pictorum et caelatorum ex Augustorum I et II cimeliis. Dresdae conditis aeri isculptorum. Gleditsch, Leipzig. https://doi.org/10.5962/bhl.title.150129
Seyfullah L.J., Beimforde C., Dal Corso J., Perrichot V., Rikkinen J. & Schmidt A.R. 2018. Production and preservation of resins – past and present. Biological Reviews 93 (3): 1684–1714. https://doi.org/10.1111/brv.12414
Singh S., Colombo W.D., Shreevihar S., Faisal M., Pandey V.P. & Tribull C.M., 2021. Rediscovery of Kurian’s types of Hymenoptera at Forest Research Institute, Dehradun, India with notes on Indian bethylid fauna. Zootaxa 5019 (1): 1–90. https://doi.org/10.11646/zootaxa.5019.1.1
Sorg M. 1988. Zur Phylogenie und Systematik der Bethylidae (Insecta, Hymenoptera, Chrysidoidea). Geologisches Institut der Universität zu Köln Sonderveröffentlichungen 63: 1–146.
Standke G. 2008. Bitterfelder Bernstein gleich Baltischer Bernstein? – Eine geologische Raum-Zeit-Betrachtung und genetische Schlußfolgerungen. Exkursionsführer und Veröffentlichungen der deutschen Gesellschaft für Geowissenschaften 236: 11–33.
Staude H.S., Maclean M., Mecenero S., Pretorius R.J., Oberprieler R.G., van Noort S., Sharp A., Sharp I., Balona J., Bradley S., Brink M., Morton A.S., Botha M.J., Collins S.C., Grobler Q., Edge D.A., Williams M.C. & Sihvonen P. 2020. An overview of Lepidoptera-host-parasitoid associations for southern Africa, including an illustrated report on 2 370 African Lepidoptera-host and 119 parasitoid-Lepidoptera associations. Metamorphosis 31 (3): 1–380. https://doi.org/10.4314/met.v31i3.1
Tribull C.M., Pankowski M.V. & Colombo W.D. 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
van Lenteren J.C. 2011. The state of commercial augmentative biological control: plenty of natural enemies, but a frustrating lack of uptake. BioControl 57: 1–20. https://doi.org/10.1007/s10526-011-9395-1
Weistchat W. & Wichard W. 2010. Baltic amber. In: Penney D. (ed.) Biodiversity of Fossils in Amber from the Major World Deposit: 80–115. Siri Scientific Press, Manchester.
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