Abstract
Eight species of Brasema Cameron, 1884 (Hymenoptera, Eupelmidae) associated with galls of Cynipini (Hymenoptera, Cynipidae) in North America are keyed and illustrated through macrophotography. The gender of Brasema is here treated as neuter rather than feminine and the reasons for this are given. Brasema mexicanum Cazorla-Vila & Pujade-Villar sp. nov. and Brasema obscurum Cazorla-Vila & Pujade-Villar sp. nov. are described from México emerging from different cynipid oak galls belonging to Quercus and Lobatae sections. Males and females of these new species are illustrated through macrophotography and scanning electron micrographs. The new species are supported as new by morphological and molecular evidence. New oak gall wasp hosts are recorded for B. auratum (Ashmead, 1886), B. dryophantae (Ashmead, 1886), B. macrocarpae (Ashmead, 1888) and B. speciosum (Girault, 1916). Distribution of B. macrocarpae and B. speciosum are expanded in USA and México, and B. flavovariegatum (Ashmead, 1888) and B. gemmarii (Ashmead, 1886) in the USA. A phylogenetic analysis of COI was carried out with 7 sequences of the two new Mexican species plus 64 sequences available from BOLD belonging to 21 identified and unidentified morphospecies to obtain the first Brasema phylogeny.
References
- Altschul, S.F., Gish, W., Miller, W., Myers, E.W. & Lipman, D.J. (1990) Basic local alignment search tool. Journal of Molecular Biology, 215 (3), 403–410. https://doi.org/10.1016/S0022-2836(05)80360-2
- Ashmead, W.H. (1886) Studies on the North-American Chalcididae with descriptions of new species from Florida (Paper No. 7). Transactions of the American Entomological Society and Proceedings of the Entomological Section of the Academy of Natural Sciences, 13 (2), 125–135. https://doi.org/10.2307/25076470
- Ashmead, W.H. (1888) Descriptions of some unknown parasitic Hymenoptera in the collection of the Kansas State Agricultural College, received from Prof. E.A. Popenoe. Bulletin of the Kansas State Agricultural College, 3, Appendix + i–viii.
- Balduf, W.V. (1926) On the bionomics of some Hymenoptera from a bur oak cynipid gall. The Canadian Entomologist, LVIII (7), 157–164. https://doi.org/10.4039/Ent58157-7
- Bird, J.P., Melika, G., Nicholls, J.A., Stone, G.N. & Buss, E.A. (2013) Life history, natural enemies, and management of Disholcaspis quercusvirens (Hymenoptera: Cynipidae) on Live Oak trees. Journal of Economic Entomology, 106 (4), 1747–1756. https://doi.org/10.1603/EC12206
- Bouček, Z. (1977) A faunistic review of the Yugoslavian Chalcidoidea (Parasitic Hymenoptera). Acta Entomologica Jugoslavica, 13, 1–145.
- Bridwell, J.C. (1897) A List of Kansas Hymenoptera. In: Smyth, B.B. (The Librarian) (Ed.), Transactions of the thirtieth and thirty-first annual meetings of the Kansas Academy of Science. Vol. XVI. W.Y. Morgan, Topeka, Kansas, pp. 203–211. https://doi.org/10.2307/3623715
- Burks, B.D. (1967) The North American species of Anastatus Motschulsky (Hymenoptera, Eupelmidae). Transactions of the American Entomological Society, 93 (4), 423–432.
- Burks, B.D. (1979) Family Eupelmidae. In: Krombein, K.V., Hurd, P.D. Jr., Smith, D.R. & Burks, B.D. (Eds.), Catalog of Hymenoptera in America North of Mexico. Vol. 1. Smithsonian Institute Press, Washington, pp. 878–889.
- Burks, R.A., Gibson, G.A.P. & La Salle, J. (2005) Nomenclatural changes in Neotropical Eulophidae, Eupelmidae and Torymidae (Hymenoptera: Chalcidoidea) relating to parasitoids of Cecidoses eremita (Lepidoptera: Cecidosidae). Zootaxa, 1082 (1), 45–55. https://doi.org/10.11646/zootaxa.1082.1.4
- Burks, R.A., Mitroiu, M.-D., Fusu, L., Heraty, J.M., Janšta, P., Heydon, S., Papilloud, N.D.-S., Peters, R.S., Tselikh, E.V., Woolley, J.B., Van-Noort, S., Baur, H., Cruaud, A., Darling, C., Haas, M., Hanson, P., Krogmann, L. & Rasplus, J. (2022) From hell’s heart I stab at thee! A determined approach towards a monophyletic Pteromalidae and reclassification of Chalcidoidea (Hymenoptera). Journal of Hymenoptera Research, 94, 13–88. https://doi.org/10.3897/jhr.94.94263
- Burks, R.A., Gibson, G.A.P. & Heraty, J.M. (2025) 5. External morphology of adult Chalcidoidea. In: Heraty, J.M & Woolley, J.B. (Eds.), Chalcidoidea of the World. CAB International, Wallingford, pp. 69–110. https://doi.org/10.1079/9781800623545.0005
- Coronado-Blanco, J.M., Ruíz Cancino, E. & Myartseva, S.N. (2005) Chalcidoidea (Hymenoptera) de Taumalipas, México (excepto Encyrtidae). In: Barrientos-Lozano, L., Correa-Sandoval, A., Horta-Vega, J.V. & Jiménez, J.G. (Eds.), Biodiversidad Tamaulipeca, 1, pp. 156–160.
- Cresson, E.T. (1887) Synopsis of the families and genera of the Hymenoptera of America, North of México, together with a catalogue of the descibed species, and bibliography. Transactions of the American Entomological Society Supplement, 14, 1–351.
- Cruaud, A., Rasplus, J.Y., Zhang, J., Burks, R., Delavare, G., Fusu, L., Gumovsky, A., Huber, J.T., Janšta, P., Mitroiu, M.D., Noyes, J.S., Van Noort, S., Baker, A., Böhmová, J., Baur, H., Blaimer, B.B., Brady, S.G., Bubeníková, K., Chartois, M., Copeland, R.S., Papilloud, N.D.S., Molin, A.D., Dominguez, C., Gebiola, M., Guerrieri, E., Kresslein, R.L., Krogmann, L., Lemmon, E., Murray, E.A., Nidelet, S., Nieves-Aldrey, J.L., Perry, R.P., Peters, R.S., Polaszek, A., Sauné, L., Torréns, J., Triapitsyn, S., Tselikh, E.V., Yoder, M., Lemmon, A.R., Woolley, J.B. & Heraty, J.M. (2024) The Chalcidoidea bush of life: evolutionary history of massive radiation of minute wasps. Cladistics, 40, 34–63. https://doi.org/10.1111/cla.12561
- Dalla Torre, C.G. (1898) Catalogus Hymenopterorum Hucusque Descriptorum Systematicus et Synonymicus. Vol V: Chalcididae et Proctotrupidae. Guilelmi Engelmann, Leipzig, 598 pp.
- Forbes, A.A., Hall, M.C., Lund, J., Hood, G.R., Izen, R., Egan, S.P. & Ott, J.R. (2016) Parasitoids, Hyperparasitoids, and Inquilines associated with the sexual and asexual generations of the gall former, Belonocnema treatae (Hymenoptera: Cynipidae). Annals of the Entomological Society of America, 109 (1), 49–63. https://doi.org/10.1093/aesa/sav112
- Fusu, L. (2013) A revision of the Palearctic species of Reikosiella (Hirticauda) (Hymenoptera, Eupelmidae). Zootaxa, 3636 (1), 1–34. https://doi.org/10.11646/zootaxa.3636.1.1
- Fusu, L. & Gibson, G.A.P. (2025) 29. Eupelmidae. In: Heraty, J.M & Woolley, J.B. (Eds.), Chalcidoidea of the World. CAB International, Wallingford, Oxfordshire, pp. 380–396. https://doi.org/10.1079/9781800623545.0029
- Gibson, G.A.P. (1986) Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist, 118 (7), 691–728. https://doi.org/10.4039/Ent118691-7
- Gibson, G.A.P. (1995) Parasitic Wasps of the Subfamily Eupelminae: Classification and Revision of World Genera (Hymenoptera: Chalcidoidea: Eupelmidae). Memoirs on Entomology, International, 5, i–v + 1.
- Gibson, G.A.P. (1997) Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera). NRC Monograph Publishing Program No. 40392. NRC Research Press, Ottawa, 794 pp. [Gibson, G.A.P., Huber, J.T. & Woolley, J.B. (Eds.)]
- Gibson, G.A.P. (2011) The species of Eupelmus (Eupelmus) Dalman and Eupelmus (Episondelia) Girault (Hymenoptera: Eupelmidae) in North America North of México. Zootaxa, 2951 (1), 1–97. https://doi.org/10.11646/zootaxa.2951.1.1
- Gibson, G.A.P. & Fusu, L. (2016) Revision of the Palearctic species of Eupelmus (Eupelmus) Dalman (Hymenoptera: Chalcidoidea: Eupelmidae). Zootaxa, 4081 (1), 1–331. http://doi.org/10.11646/zootaxa.4081.1.1
- Girault, A.A. (1916) Descriptions of and observations on some chalcidoid Hymenoptera. The Canadian Entomologist, 48 (7), 242–246. https://doi.org/10.4039/Ent48242-7
- Glick, A.P. (1939) The distribution of Insects, spiders and mites in the air. Technical Bulletin of the United States Department of Agriculture, 673, 1–150. https://doi.org/10.22004/ag.econ.168268
- González-Hernández, A. (2004) Actualización del catálogo de autoridades taxonómicas: Hymenoptera-Parasítica. Departamento de Zoología de Invertebrados, Facultad de Ciencias Biológicas, Universidad Autónoma de Nuevo León. Proyecto BE008. Base de datos SNIB-CONABIO, México, D.F., 15 pp.
- Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: Improving the Ultrafast Bootstrap Approximation. Molecular Biology and Evolution, 35 (2), 518–522. https://doi.org/10.1093/molbev/msx281
- International Commission of Zoological Nomenclature (1999) International Code of Zoological Nomenclature. 4th Edition. The International Trust for Zoological Nomenclature, London, 306 pp. [https://www.iczn.org/the-code/the-code-online/]
- Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K., Von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature methods, 14 (6), 587–589. https://doi.org/10.1038/nmeth.4285
- Kamura, T., Stecher, G. & Kumar, S. (2021) MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution, 38 (7), 3022–3027. https://doi.org/10.1093/molbev/msab120
- Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M.D., Von Haeseler, A. & Lanfear, R. (2020) IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution, 37 (5), 1530–1534. https://doi.org/10.1093/molbev/msaa015
- Peck, O. (1951) Superfamily Chalcidoidea. In: Muesebeck, C.F.W., Krombein, K.V. & Townes, H.K. (Eds.), Hymenoptera of America North of Mexico: Synoptic Catalog. U.S. Department of Agriculture, Agriculture Monographs No. 2. United States Government Printing Office, Washington, D.C., pp. 410–594.
- Peck, O. (1963) A catalogue of the Nearctic Chalcidoidea (Insecta; Hymenoptera). Canadian Entomologist Supplement, 30, 1–1092. https://doi.org/10.4039/entm9530fv
- Ruíz Cancino, E., Coronado Blanco, J.M., Myartseva, S.N. & Luna Salas, J.F. (2004) Adenda a Chalcidoidea (Hymenoptera). In: Bousquets, J.L., Morrone, J.J., Ordóñez, O.Y. & Fernández, I.V. (Eds.), Biodiversidad, Taxonomía y Biogeografía de artrópodos de México: Hacia una síntesis de su conocimiento, IV, pp. 725–734.
- Schmiedeknecht, O. (1909) Hymenoptera, Fam. Chalcididae. In: Verteneuil, V. & Desmet, L. (Eds.), Genera Insectorum. Fascicle: XCVII. Wytsman, P., Bruxelles, pp. 1–550.
- Schulz, W.A. (1906) Spolia hymenopterologica. Junfermannsche Buchhanflung, Paderborn, 335 pp. https://doi.org/10.5962/bhl.title.59757
- UCD Community (2023) Universal Chalcidoidea Database Website. Available from: https://ucd.chalcid.org (accessed 31 May 2025)
- Ward, A.K., Busbee, R.W., Chen, R.A., Davis, C.K., Driscoe, A.L., Egan, S.P., Goldberg, B.A.R., Hood, G.R., Jones, D.G., Kranz, A.J., Meadely-Dunphy, S.A., Milks, A.K., Ott, J.R., Prior, K.M., Sheikh, S.I., Shzu, S., Weinersmith, K.L., Zhang, L., Zhang, Y.M. & Forbes, A.A. (2022) The arthropod associates of 155 North American Cynipid Oak Galls. Zoological Studies, 61, e57. https://doi.org/10.1101/2022.04.26.489445
- Weinersmith, K.L., Forbes, A.A., Ward, A.K., Brandão-Dias, P.F., Zhang, Y.M. & Egan, S.P. (2020) Arthropod community associated with the asexual generation of Bassettia pallida (Hymenoptera: Cynipidae). Annals of the Entomological Society of America, 113 (5), 373–388. https://doi.org/10.1093/aesa/saaa009
