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Published: 2021-02-26

Re-description and systematic re-appraisal of the genus Kokkocynips Pujade-Villar & Melika, (Hymenoptera: Cynipidae: Cynipini), including new combinations of Nearctic species and the description of a new species from Panama

Museo Nacional de Ciencias Naturales (CSIC), Departamento de Biodiversidad y Biología Evolutiva, C/ José Gutiérrez Abascal 2, ES-28006 Madrid, Spain.
CSIRO, Australian National Insect Collection, Clunies Ross Street, Acton, ACT 2601, Australia.
Department of Biological Sciences, The George Washington University, SEH, Suite 6300, 800 22nd Street, NW, Washington, DC 20052-0001, USA. Institute of Evolutionary Biology, University of Edinburgh, King’s Buildings, Charlotte Auerbach Road, Edinburgh EH9 3JT, Scotland, U.K.
Plant Health and Molecular Biology Laboratory, National Food Chain Safety Office, Directorate of Plant Protection, Soil Conserva
Institute of Evolutionary Biology, University of Edinburgh, King’s Buildings, Charlotte Auerbach Road, Edinburgh EH9 3JT, Scotland, U.K.
Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Animal, Avda. Diagonal 645, 08028-Barcelona (Spain).
Systematic Entomology Laboratory, ARS-USDA c/o National Museum of Natural History, Smithsonian Institution, 10th & Constitution Ave NW, Washington, DC 20013, USA.
Cátedras CONACYT‑Instituto de Investigaciones sobre los Recursos Naturales, Universidad Michoacana de San Nicolás de Hidalgo,
Programa de Maestría en Entomología y Departamento de Ciencias Ambientales, Facultad de Ciencias Naturales, Exactas y Tecnología Universidad de Panamá. C. P. 0824.
Hymenoptera oak gall wasps key Dryocosmus Callirhytis Chiriquí Quercus Lobatae red oaks


The monophyly and taxonomic validity of some currently accepted genera of gall wasps in the Cynipini (Hymenoptera: Cynipidae) are being challenged by recent systematic studies. Here we used morphological and molecular data to re-describe and revise the taxonomic limits of the monotypic genus Kokkocynips Pujade-Villar & Melika, previously recorded only from Mexico.

                We describe a new species from Panama, Kokkocynips panamensis Medianero & Nieves-Aldrey, reared from galls on Quercus salicifolia Neé, and add new records from Mexico for the type species of the genus K. doctorrosae Pujade-Villar. Six Nearctic species, Dryocosmus rileyi (Ashmead, 1896), D. imbricariae (Ashmead, 1896), D. coxii (Basset, 1881), D. deciduus (Beutenmueller, 1913), Callirhytis difficilis (Ashmead, 1887) and C. attractans (Kinsey, 1922) are transferred to Kokkocynips. Species of Kokkocynips are associated only with red oaks (Quercus section Lobatae (Fagaceae)) and are distributed in the Nearctic and Neotropical regions, from Canada through Mexico and Panama. Taxonomic limits of Kokkocynips are discussed in light of the molecular phylogenetic relationships of the studied species. Diagnostic characters, gall description, distribution, and biological data of Kokkocynips species are given, including a key for the identification of the asexual generations of seven species.



  1. Ashmead, W.H. (1885) A Bibliographical and Synonymical Catalogue of the North American Cynipidae, with Description of New Species. Transactions of the American Entomological Society, 12, 291–304.

    Ashmead, W.H. (1887) On the cynipidous galls of Florida, with descriptions of new species and synopses of the described species of North America. Transactions of the American Entomological Society, 14, 125–158.

    Ashmead, W.H. (1896) Descriptions of new cynipidous galls and gall-wasps in the United States National Museum. Proceedings of the United States National Museum, 19, 113–136.

    Bassett, H.F. (1881) New species of Cynipidae. The Canadian Entomologist, 13, 51–112.

    Bassett, H.F. (1890) New species of North American Cynipidae. Transations of the American Entomological Society, 26, 310–336.

    Beutenmueller, W. (1913) Description of a new Gallfly (Andricus decidua) (Hymenoptera, Cynipidae). Insecutor Inscitiae Mestruus, 1, 131–132.

    Blaimer, B.B., Gotzek, D., Brady, S.G. & Buffington, M.L. (2020) Comprehensive phylogenomic analyses re-write the evolution of parasitism within cynipoid wasps. BMC Evolutionary Biology, 20 (155), 1–22.

    Breedlove, D. (2001) Fagaceae. In: Stevens, W., Ulloa, U., Pool, A. & Montiel, O. (Eds.), Flora de Nicaragua. Monographs in Systematic Botany. Missouri Botanical Garden, St. Louis, Missouri, pp. 1076–1084.

    Burger, W. (1977) Fagaceae. In: Burger, W. (Eds.), Flora Costaricensis. Fieldiana: Botany. Field Museum of Natural History, Chicago, Illinois, pp. 59–82.

    Burks, B.D. (1979) Superfamily Cynipoidea. In: Krombein, K., Hurd, P., Smith, D. & Burks, B. (Eds.), Catalog of Hymenoptera in America of North of Mexico. Vol. 1. Symphyta and Apocrita. Smithsonian Inst. Press, Washington, D.C., pp. 1045–1107.

    Buffington, M.L., Forshage, M., Liljeblad, J., Tang, C.-T. & van Noort, S. (2020) World Cynipoidea (Hymenoptera): A key to higher-level group. Insect Systematics and Diversity, 4, 1–69.

    Cerasa, G., Lo Verde, G., Caleca, V., Massa, B, Nicholls, J.A. & Melika, G. (2018) Description of Dryocosmus destefanii new species (Hymenoptera: Cynipidae: Cynipini) from Quercus suber L. in Italy. Zootaxa, 4370 (5), 535–548.

    Cerasa, G., Lo Verde, G., Caleca, V., Massa, B, Nicholls, J.A. & Melika, G. (2020) Description of the sexual generation of Dryocosmus destefanii (Hymenoptera: Cynipidae: Cynipini) and disclosure of its life cycle. Zootaxa, 4742 (2), 359–374.

    Csóka, G., Stone, G.N. & Melika, G. (2005) Biology, ecology, and evolution of gall-inducing Cynipidae. In: Raman, A., Schaefer, C.W. & Withers, T.M. (Eds.), Biology, ecology, and evolution of gall-inducing arthropods. Science Publishers, Inc., Enfield, New Hamsphire, pp. 573–642.

    Cuesta-Porta, V., Arnedo, M.A., Cibrián-Tovar, D., Barrera-Ruiz, U.M., García-Martiñón, R.D., Equihua-Martínez, A., Estrada-Venegas, E.G., Clark-Tapia, R., Romero-Range, S. & Pujade-Villar, J. (2020) A new genus of oak gall wasp, Striatoandricus Pujade-Villar (Hymenoptera: Cynipidae: Cynipini) from America with descriptions of two new Mexican species. Zoological Studies, 59 (8), 1–21.

    Dalla Torre, K.W. & Kieffer, J.J. (1910) Cynipidae. In: Das Tierreich. Vol. 24. Friedlander & Sohn, Berlin, pp. 1–891.

    Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772.

    Dailey, D. C. (1969) Synonymy of Dryocosmus attractans (Kinsey) and Callirhytis uvellae Weld (Hymenoptera: Cynipidae). The Pan-Pacific Entomologist, 45 (2), 132–134.

    Fang, Z., Tang, C.-T., Nicholls, J.A., Zhu, Y., Xiong, T., Hearn, J., Sinclair, F., Melika, G., Nieves-Aldrey, J.L., Csóka, G., Mikolajczak, K.M., Stone, G.N. & Fang, S. (2020) A New Genus of Oak Gallwasp, Heocynips Fang, Nieves-Aldrey, and Melika (Hymenoptera: Cynipidae: Cynipini), from China. Proceedings of the Entomological Society of Washington, 122 (4), 787–804.

    Giraud, J.E. (1859) Signalements de quelques espèces nouvelles de Cynipides et de leurs Galles. Verhandlungen des Zoologisch-Botanischen Vereins in Wien, 9, 337–374.

    Harris, R.A. (1979) A glossary of surface sculpturing. State of California, Department of Food and Agriculture. Occasional Papers in Entomology, 28, 1–31.

    Katoh, S. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.

    Kinsey, A.C. (1922a) Studies of some new and described Cynipidae (Hymenoptera). Indiana University Studies, 9 (53), 1–141.

    Kinsey, A.C. (1922b) New Pacific Coast Cynipidae (Hymenoptera). Bulletin of the American Museum of Natural History, 46, 279–296.

    Liljeblad, J., Ronquist, F., Nieves-Aldrey, J. L., Fontal-Cazalla, F., Ros-Farre, P., Gaitros, D. & Pujade-Villar, J. (2008) A fully web-illustrated morphological phylogenetic study of relationships among oak gall wasps and their closest relatives (Hymenoptera: Cynipidae). Zootaxa, 1796 (1), 1–73.

    Mayr, G. (1902) Ueber Nordamerichaniske Cynipiden. Verdhanlungen der k.k. zoologische-botanischen Gesellschaft in Wien, 52, 287–290.

    Medianero, E. & Nieves-Aldrey, J.L. (2011) Primer estudio de las avispas de las agallas de la república de Panamá, incluyendo una lista actualizada de los cinípidos neotropicales (Hymenoptera, Cynipoidea, Cynipidae). Boletín de la Sociedad Entomólogica Aragonesa (S.E.A.), 48, 89–104.

    Medianero, E. & Nieves-Aldrey, J.L. (2013) Barucynips panamensis, a new genus and species of oak gallwasps (Hymenoptera, Cynipidae, Cynipini) from Panama, and description of one new species of Coffeikokkos. ZooKeys, 277, 25–46.

    Melika, G. & Abrahamson, W.G. (2002) Review of the World Genera of Oak Cynipid Wasps (Hymenoptera: Cynipidae, Cynipini). In: Melika, G. & Thuróczy, Cs. (Eds.), Parasitic Wasps: Evolution, Systematics, Biodiversity and Biological Control. Agroinform, Budapest, pp. 150–190.

    Melika, G. (2006) Gall wasps of Ukraine cynipidae. Vestnik Zoologii, Supplement 21 (1–2), 1–300 + 301–644.

    Melika, G., Pujade-Villar, J., Abe, Y., Tang, C.T., Nicholls, J.A., Wachi, N., Ide, T., Yang, M.M., Pénzes, Zs., Csóka, Gy. & Stone, G.N. (2010) Palaearctic oak gallwasps galling oaks (Quercus) in the section Cerris: re-appraisal of generic limits, with descriptions of new genera and species (Hymenoptera: Cynipidae: Cynipini). Zootaxa, 2470 (1), 1–79.

    Melika, G., Tang, C.-T., Sinclair, F., Yang, M.-M., Lohse, K., Hearn, J., Nicholls, J.A. & Stone, G.N. (2013) A new genus of oak gallwasp, Cyclocynips Melika, Tang & Sinclair (Hymenoptera: Cynipidae: Cynipini), with descriptions of two new species from Taiwan. Zootaxa, 3630 (3), 534–548.

    Nicholls, J.A. & Pujade-Villar, J. (2020) Re-instatement of the species name Callirhytis erythrosoma (Dettmer, 1933), with comments on other Callirhytis species (Hymenoptera: Cynipidae). Butlletiì de la Institucioì Catalana d’HistoÌria Natural, 84, 31–34.

    Nicholls, J.A., Melika, G. & Stone, G.N. (2017) Sweet tetra-trophic interactions: multiple evolution of nectar secretion, a defensive extended phenotype in cynipid gallwasps. American Naturalist, 189, 67–77.

    Nicholls, J.A., Melika, G., DeMartini, J. & Stone, G.N. (2018a) New species of Dryocosmus Giraud gallwasps from California (Hymenoptera: Cynipidae: Cynipini) galling Chrysolepis Hjelq. (Fagaceae). Zootaxa, 4532 (3), 407–433.

    Nicholls J.A., Stone, G.N. & Melika, G. (2018b) A new genus of oak gallwasp, Protobalandricus Melika, Nicholls and Stone (Hymenoptera: Cynipidae: Cynipini) from California. Zootaxa, 4472 (1), 141–152.

    Nieves-Aldrey, J.L. (1992) Revision de las especies Europeas del genero Callirhytis Förster (Hymenoptera, Cynipidae). Graellsia, 48, 171–183.

    Nieves-Aldrey, J.L. (2001) Hymenoptera, Cynipidae. In: Ramos, M., Alba-Tercedor, J., Bellés-i-Ros, X., Gosálbez-i-Noguera, J., Guerra-Sierra, A., Macpherson-Mayol, E., Martín-Piera, F., Serrano-Marino, J. & Templado-González, J. (Eds.), Fauna Ibérica. Museo Nacional de Ciencias Naturales, CSIC, Madrid, pp. 1–636.

    Nieves-Aldrey, J.L, Liljeblad, J., Hernández Nieves, M., Grez, A. & Nylander, J.A.A. (2009) Revision and phylogenetics of the genus Paraulax Kieffer (Hymenoptera, Cynipidae) with biological notes and description of a new tribe, a new genus, and five new species. Zootaxa, 2200 (1), 1–40.

    Pénzes, Z., Tang, C.-T., Stone, G.N., Nicholls, J.A., Schwéger, S., Bozsó, M. & Melika, G. (2018) Current status of the oak gallwasp (Hymenoptera: Cynipidae: Cynipini) fauna of the Eastern Palaearctic and Oriental Regions. Zootaxa, 4433 (2), 245–289.

    Pujade, J. & Arnedo, M.A. (1998) Morfologia i evolució de la genitàlia masculina dels Cynipinae. Sessió Conjunta d’Entomologia de la ICHN-SCL, 9 (1995), 115–135.

    Pujade-Villar, J., Hanson, P., Medinas, C.A., Torres, M. & Melika, G. (2012a) A new genus of oak gallwasps, Zapatella Pujade-Villar and Melika, gen. n., with a description of two new species from the Neotropics (Hymenoptera, Cynipidae, Cynipini). Zookeys, 210, 75–104.

    Pujade-Villar, J., Hanson, P. & Melika, G. (2012b) A new genus of oak gallwasp, Coffeikokkos Pujade-Villar and Melika, gen. n., with a description of a new species from Costa Rica (Hymenoptera, Cynipidae). Zookeys, 168, 19–29.

    Pujade-Villar J, Romero-Rangel S, Chagoyán-Garcia C, Equihua Martinez A, Estrada-Venegas EG, Melika G. (2010) A new genus of oak gallwasps, Kinseyella Pujade-Villar and Melika, with a description of a new species from Mexico (Hymenoptera: Cynipidae: Cynipini). Zootaxa, 2335 (1), 16–28.

    Pujade-Villar, J., Cibrián-Tovar, D., Cibrián-Llanderal, V.D., Equihua-Martínez, A., Estrada-Venegas, E.G., Serrano-Muñoz, M. & Lomeli-Flores, J.R. (2014) A new genus of oak gallwasp, Melikaiella Pujade-Villar (Hymenoptera: Cynipidae: Cynipini), from the Nearctic region. Dugesiana, 21 (1), 1–29.

    Pujade-Villar, J., Equihua-Martínez, A., Estrada-Venegas, E.G. & Melika, G. (2013) A new genus of oak gallwasp, Kokkocynips Pujade-Villar & Melika gen. n., with a description of a new species from Mexico (Hymenoptera, Cynipidae). Acta Zoológica, Mexican, New Series, 29 (1), 209–218.

    Ronquist, F. & Nordlander, G. (1989) Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera, Ibaliidae). Entomologica Scandinavica, Supplement 33, 1–60.

    Ronquist, F. (1995) Phylogeny and early evolution of the Cynipoidea (Hymenoptera). Systematic Entomology, 20, 309–335.

    Ronquist, F., Nieves-Aldrey, J.L., Buffington, M.L., Liu, Z., Liljeblad, J. & Nylander, J.A. (2015) Phylogeny, evolution and classification of gall wasps: the plot thickens. PLoS One, 10, e0123301.

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MRBAYES 3.2: Efficient Bayesian phylogenetic inference and model selection across a large model space. Systematic Biology, 61, 539–542.

    Stone, G.N., Atkinson, R.J., Rokas, A., Nieves-Aldrey, J.-L., Melika, G., Ács, Z., Csóka, G., Hayward, A., Bailey, R., Buckee, C. & McVean, G.A.T. (2008) Evidence for widespread cryptic sexual generations in apparently asexual Andricus gallwasps. Molecular Ecology, 17, 652–665.

    Stone, G.N., Hernandez-Lopez, A., Nicholls, J.A., di Pierro, E., Pujade-Villar, J., Melika, G. & Cook, J.M. (2009) Extreme host plant conservatism during at least 20 million years of host plant pursuit by oak gallwasps. Evolution, 63, 854–869.

    Swofford, D.L. (1998) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Sinauer Associates, Sunderland, Massachusetts. [program]

    Tang, C.T., Melika, G., Nicholls, J. A., Yang, M.M. & Stone, G.N. (2011) A new genus of oak gallwasps, Cycloneuroterus Melika & Tang, with the description of five new species from Taiwan (Hymenoptera: Cynipidae: Cynipini). Zootaxa, 3008 (1), 33–62.

    Tang, C.T., Mikó, I., Nicholls, J.A., Schwéger, S., Yang, M-M., Stone, G.N., Sinclair, Bozsó, M., Melika, G. & Pénzes, Z. (2016) New Dryocosmus Giraud species associated with Cyclobalanopsis and non-Quercus host plants from the Eastern Palaearctic (Hymenoptera, Cynipidae, Cynipini). Journal of Hymenoptera Research, 53, 77–163.

    van Noort, S., Stone, G.N., Whitehead, V.B. & Nieves-Aldrey, J.-L. (2007) Biology and redescription of Rhoophilus loewi (Cynipidae: Cynipoidea: Hymenoptera), with evolutionary implications on the inquilinism in gall wasps. Biological Journal of the Linnaean Society, 90, 153–172.

    Wachi, N., Ide, T. & Abe, Y (2011) A new inquiline species of Saphonecrus (Hymenoptera: Cynipidae: Synergini) associated with cecidomyiid galls on oak trees in Japan

    Annals of the Entomological Society of America, 104 (3), 369–373.

    Weld, L. (1926a) Notes on cynipid wasps, with descriptions of new North American species. Proceedings of the United States National Museum, 61 (18), 1–29.

    Weld, L. (1926b) Field notes on gall-inhabiting cynipid wasps with description of new species. Proceedings of the United States National Museum, 68 (10), 1–131.

    Weld, L.H. (1944) New American cynipids from galls. Proceedings of the United States National Museum, 95, 1–24.

    Weld, L.H. (1951) Superfamily Cynipoidea. In: Muesebeck, C.F.W., Krombein, K.V. & Townes, H.K. (Eds.), Hymenoptera of America North of Mexico. Agricultural Monograph No. 2. United States Government Printing Office, Washington, D.C., pp.594–654.

    Weld, L.H. (1952) Cynipoidea (Hym.) 19051950, being a supplement to the Dalla Torre and Kieffer monograph, the Cynipidae in Das Tierreich, Lieferung 24, 1910, and bringing the systematic literature of the world up to date, including keys to families and subfamilies and lists of new generic, specific and variety names. Privately printed, Ann Arbor, Michigan, 351 pp.

    Weld, L.H. (1959) Cynipid galls of the Eastern United States. Privately printed, Ann Arbor, Michigan, 124 pp.