Abstract
Four species of Dryocosmus cynipid gallwasps are now known to induce galls on Chrysolepis in California and Oregon. Two new species, Dryocosmus demartinii Melika, Nicholls & Stone and Dryocosmus juliae Melika, Nicholls & Stone are described. Males of the sexual generation of D. rileypokei plus adults of the asexual generation of this species are both described for the first time. A detailed description of D. castanopsidis is given for the first time. In addition to descriptions we provide diagnoses and information on biology and host associations for all four species. All the newly described taxa are supported by morphological and molecular data.
References
Ács, Z., Melika, G., Pénzes, Z., Pujade-Villar, J. & Stone, G.N. (2007) The phylogenetic relationships between Dryocosmus, Chilaspis and allied genera of oak gallwasps (Hymenoptera, Cynipidae: Cynipini). Systematic Entomology, 32, 70–80.
https://doi.org/10.1111/j.1365-3113.2006.00351.x
Azizkhani, E., Rasoulian, G.-R., Kharazi-Pardel, A., Tavakoli, M., Sadeghi, S.E., Melika, G., Stone, G.N. & Atkinson, R. (2006) New species of oak gall wasps from Zagross Mountains of Iran (Hymenoptera: Cynipidae: Cynipini). Folia Entomologica Hungarica, 67, 161–197.
Beutenmueller, W. (1917) Descriptions of new Cynipidae. Canadian Entomologist, 49, 345–349.
https://doi.org/10.4039/Ent49345-10
Bouckaert, R., Heled, J., Kühnert, D., Vaughan, T., Wu, C.-H., Xie, D., Suchard, M.A., Rambaut, A. & Drummond, A.J. (2014) BEAST 2: a software platform for Bayesian evolutionary analysis. PLoS Computational Biology, 10, e1003537.
https://doi.org/10.1371/journal.pcbi.1003537
Buffington, M.L. & Morita, S.I. (2009) Not all oak gall wasps gall oaks: the description of Dryocosmus rileypokei, a new, apostate species of Cynipini from California. Proceedings of the Entomological Society of Washington, 111 (1), 244–253.
https://doi.org/10.4289/0013-8797-111.1.244
Burks, B.D. (1979) Superfamily Cynipoidea. 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. Symphyta and Apocrita. Smithsonian Institution Press, Washington, D.C., pp. 1045–1107.
Campbell, B.C., Steffen-Campbell, J.D. & Werren, J.H. (1993) Phylogeny of the Nasonia species complex (Hymenoptera: Pteromalidae) inferred from an internal transcribed spacer (ITS2) and 28S rDNA sequences. Insect Molecular Biology, 2, 225–237.
https://doi.org/10.1111/j.1365-2583.1994.tb00142.x
Cerasa, G., LoVerde, 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.
https://doi.org/10.11646/zootaxa.4370.5.5
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 Hampshire, pp. 569–636.
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772.
https://doi.org/10.1038/nmeth.2109
Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797.
https://doi.org/10.1093/nar/gkh340
Gene Codes Corporation (1995) Sequencher. Version 3.0. Ann Arbor, Michigan. [software]
Harris, R. (1979) A glossary of surface sculpturing. State of California, Department of Food and Agriculture, Occasional Papers in Entomology, 28, 1–31.
Hymenoptera Online (HOL). (2018) Available from: https://hol.osu.edu (accessed 13 November 2018)
Ide, T. & Abe, Y. (2015) A new species of Dryocosmus Giraud (Hymenoptera: Cynipidae: Cynipini) in Japan and Korea—first record of Eastern Palearctic Dryocosmus species showing alternation of generations on section Cerris oaks. Proceedings of the Entomological Society of Washington, 117 (4), 467–480.
https://doi.org/10.4289/0013-8797.117.4.467
Ide, T., Wachi, N. & Abe, Y. (2013) Description of two new species of Dryocosmus (Hymenoptera: Cynipidae: Cynipini) inducing galls on the strictly Asian subgenus Cyclobalanopsis of the genus Quercus, with a key to species of Dryocosmus in East Asia. Annals of the Entomological Society of America, 106, 18–25.
https://doi.org/10.1603/AN12050
Kinsey, A.C. (1922) Studies of some new and described Cynipidae (Hymenoptera). Indiana University Studies, 9 (53), 1–141.
Liljeblad, J., Ronquist, F., Nieves-Aldrey, J.-L., Fontal-Cazalla, F., Ros-Farré, 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, 1–73.
Melika, G. (2006) Gall Wasps of Ukraine. Cynipidae. Vestnik zoologii, Supplement 21 (1–2), 1–300 + 301–644.
Melika, G. & Abrahamson, W.G. (2002) Review of the world genera of oak cynipid wasps (Hymenoptera: Cynipidae: Cynipini). In: Melika, G. & Thuróczy, C. (Eds.), Parasitic wasps: evolution, systematics, biodiversity and biological control. Agroinform, Budapest, pp. 150–190.
Melika, G., Pujade-Villar, J., Abe, Y., Tang, C.-T., Nicholls, J., Wachi, N., Ide, T., Yang, M.M., Pénzes, Z., Csóka, G. & 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, 1–79.
Melika, G., Tang, C.-T., Nicholls, J.A., Yang, M.-M. & Stone, G.N. (2011) Four new species of Dryocosmus gallwasps from Taiwan (Hymenoptera: Cynipidae, Cynipini). ISRN Zoology, 2011, 725180.
https://doi.org/10.5402/2011/725180
Nicholls, J.A., Challis, R.J., Mutun, S. & Stone, G.N. (2012) Mitochondrial barcodes are diagnostic of shared refugia but not species in hybridising oak gallwasps. Molecular Ecology, 21, 4051–4062.
https://doi.org/10.1111/j.1365-294X.2012.05683.x
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.
https://doi.org/10.1086/689399
Nicholls, J.A., Melika, G., DeMartini, J.D. & Stone, G.N. (2018) A new species of Andricus Hartig gallwasp from California (Hymenoptera: Cynipidae: Cynipini) galling Notholithocarpus (Fagaceae). Stuttgarter Beiträge zur Naturkunde A, Biologie. [in press]
Pang, Y., Su, C.-Y. & Zhu, D.-H. (2018) Discovery of a new species of Dryocosmus Giraud, 1859 (Hymenoptera: Cynipidae: Cynipini) on Castanopsis tibetana from China. Zootaxa, 4403 (1), 99–110.
https://doi.org/10.11646/zootaxa.4403.1.5
Pénzes, Z., Tang, C.-T., Stone, G.N., Nicholls, J.A., Schwéger, S., Bozsó, M. & Melika, G. (2008) Current status of the oak gallwasp (Hymenoptera: Cynipidae: Cynipini) fauna of the Eastern Palaearctic and Oriental Regions. Zootaxa, 4433 (2), 245–289.
https://doi.org/10.11646/zootaxa.4433.2.2
Ronquist, F., Nieves-Aldrey, J.-L., Buffington, M.L., Liu, Zh., Liljeblad, J. & Nylander, J.A.A. (2015) Phylogeny, Evolution and Classification of Gall Wasps: The Plot Thickens. PLoS ONE, 10, e0123301.
https://doi.org/10.1371/journal.pone.0123301
Ronquist, F. & Nordlander, G. (1989) Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, 33 (Supplement), 1–60.
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 purely asexual Andricus gallwasps. Molecular Ecology, 17, 652–665.
https://doi.org/10.1111/j.1365-294X.2007.03573.x
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.
https://doi.org/10.1111/j.1558-5646.2008.00604.x
Stone, G.N., White, S.C., Csóka, G., Melika, G., Mutun, S., Pénzes, Z., Sadeghi, S.E., Schönrogge, K., Tavakoli, M. & Nicholls, J.A. (2017) Tournament ABC analysis of the Western Palaearctic population history of an oak gallwasp, Synergus umbraculus. Molecular Ecology, 26, 6685–6703.
https://doi.org/10.1111/mec.14372
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, 33–62.
Tang, C.-T., Mikó, I., Nicholls, J.A., Schwéger, S., Yang, M.-M., Stone, G.N., Sinclair, F., 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, 77–162.
https://doi.org/10.3897/jhr.53.9890
Tavakoli, M., Melika, G., Sadeghi, S.E., Pénzes, Z., Assareh, M.A., Atkinson, R., Bechtold, M., Mikó, I., Zargaran, M.R., Aligolizade, D., Barimani, H., Bihari, P., Pirozi, F., Fülöp, D., Somogyi, K., Challis, R., Preuss, S., Nicholls, J. & Stone, G.N. (2008) New species of oak gallwaps from Iran (Hymenoptera: Cynipidae: Cynipini). Zootaxa, 1699, 1–64.
Weld, L.H. (1957) Cynipid galls of the Pacific slope. Privately printed, Ann Arbor, MI, 124 pp.
Zhu, D.-H., Liu, Z., Lu, P.-F., Yang, X.-H., Su, C.-Y. & Liu, P. (2015) New Gall Wasp Species Attacking Chestnut Trees: Dryocosmus zhuili n. sp. (Hymenoptera: Cynipidae) on Castanea henryi from Southeastern China. Journal of InsectScience, 15 (1), 156.
https://doi.org/10.1093/jisesa/iev118