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Type: Article
Published: 2023-05-26
Page range: 210-232
Abstract views: 381
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The experimental life cycle closure of Andricus truncicolus (Giraud, 1859) (Hymenoptera: Cynipidae: Cynipini) and taxonomic description of its sexual generation

Museo Civico di Lentate Sul Seveso; Via D. Aureggi 25; 20823 Lentate Sul Seveso
Department of Agricultural; Food and Forest Sciences (SAAF); University of Palermo; Viale delle Scienze Bd. 5A; 90128 Palermo
Department of Agricultural; Food and Forest Sciences (SAAF); University of Palermo; Viale delle Scienze Bd. 5A; 90128 Palermo
Department of Agricultural; Food and Forest Sciences (SAAF); University of Palermo; Viale delle Scienze Bd. 5A; 90128 Palermo
Hymenoptera oak gallwasp heteroecic life cycle biology sexual generation morphology distribution Quercus


The life cycle of Andricus truncicolus (Giraud, 1869), a species previously known only from its asexual generation, is closed. Laboratory rearing and field sampling have revealed the sexual generation of this species, which develops in a hypertrophic degeneration of the terminal or lateral shoot buds appearing as a muddled agglomerate of leaves on Quercus, section Cerris oaks. We describe for the first time the newly discovered sexual generation of A. truncicolus and provide illustrations and diagnosis for adults and galls of both generations, further observations on biology and information on the species distribution.



  1. Askew, R.R. (1984) The Biology of gall wasps. In: Ananthakrishnan, T.N. (Ed.), Biology of gall insects. Edward Arnold, London, pp. 223–271.
  2. Avtzis, D.N., Melika G., Matošević, D. & Coyle, D.R. (2018) The Asian chestnut gall wasp Dryocosmus kuriphilus: a global invader and a successful case of classical biological control. Journal of Pest Science, 92 (1), 107–115. DOI:
  3. Benson, R.B. (1953) Revision of Nomenclature. In: Marsden-Jones, E.M. (Ed), A study of the life-cycle of Adleria kollari Hartig, the Marble or Devonshire gall. Transactions of the Royal entomological Society of London, 104 (7), 195–222. DOI:
  4. 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.
  5. DOI:
  6. 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.
  7. DOI:
  8. Cerasa, G., Sottile, S., Massa, B. & Lo Verde, G. (2021). Rediscovery of Cerroneuroterus apenninus (Trotter, 1923) (Hymenoptera: Cynipidae: Cynipini) a valid species from Italy with description and neotype designation. Zootaxa, 4941 (3), 399–414. DOI:
  9. 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. 573–642.
  10. Dalla Torre, K.W. & Kieffer, J.J. (1910) Cynipidae. Das Tierreich. Vol. 24. Friedlander & Sohn, Berlin, 891 pp. DOI:
  11. De Stefani T. (1901) Contribuzione all’entomocecidiologia della flora sicula. Nuovo Giornale Botanico Italiano. Memorie della Società Botanica Italiana, 8, 440–556.
  12. Doutt, R.L. (1959) Heterogony in Dryocosmus (Hymenoptera: Cynipidae). Annals of the Entomological Society of America, 52, 69–74. DOI:
  13. Eady, R.D. (1968) Some illustrations of microsculpture in the Hymenoptera. Proceedings of the Royal Entomological Society of London, 43, 66–72. DOI:
  14. Fergusson, N.D.M. (1995) The cynipoid families. In: Hanson, P.E. & Gauld, I.D. (Eds.), The Hymenoptera of Costa Rica. Oxford University Press, Oxford, New York and Tokyo, pp. 247–265.
  15. Folliot, R. (1964) Contribution a l’étude de la biologie des cynipides gallicoles (Hymenoptera, Cynipoidea). Annales des Sciences Naturelles Zoologie, 12 (6), 407–564.
  16. Folliot, R., Ros-Farré, P., Bellido, D. & Pujade-Villar, J. (2004) Alternation of generations in Andricus corruptrix (Schlechtendal): Comments on and description of a new sexual form (Hymenoptera: Cynipidae). Contributions to Zoology, 73 (4), 263–270. DOI:
  17. Folliot, R. & Pujade-Villar, J. (2006) Males of Andricus hystrix Trotter, a new sexual form of Cynipidae (Hymenoptera). Boletín Sociedad Entomológica Aragonesa, 38, 157–160.
  18. Garbin, L., Díaz, N.B. & Pujade-Villar, J. (2008) Experimental study of the reproductive cycle of Plagiotrochus amenti Kieffer, 1901 (Hymenoptera, Cynipoidea, Cynipidae), with comments on its taxonomy. Boletín de la Asociación española de Entomología, 32, 341–349.
  19. 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.
  20. Graeffe, E. (1907) Beitrag zur Kenntnis der gallenbewohnenden Cynipinen der Umgebung Triests. Bollettino della Società Adriatica di Scienze Naturali in Trieste, 23, 1–61
  21. Hadley, A. (2011) Combine ZP. Available from: (accessed 12 February 2015)
  22. Harris, R. (1979) A glossary of surface sculpturing. State of California, Department of Food and Agriculture, Occasional Papers in Entomology, 28, 1–31.
  23. ICZN [International Commission on Zoological Nomenclature] (1999) International Code of Zoological Nomenclature. 4th Edition. The International Trust for Zoological Nomenclature: London, 305 pp. Available from: (accessed 31 May 2021)
  24. Ionescu, A.M. (1957) Fauna Republicii Populare Romine. Insecta. Cynipinae. 9 (2). Academiei Republicii Populare Romane Press, Bucuresti, 246 pp.
  25. Katılmış, Y. & Kıyak, S. (2008) Checklist of Cynipidae of Turkey with a new genus record. Journal of Natural History, 42 (31–32), 2161–2167. DOI:
  26. Kieffer, J.J. (1897-1901) Les Cynipides. In: André, E. (Eds.), Species des Hyménoptères d’Europe & d’Algérie. Tome Septième. Froment-Dubosclard, Paris, pp. 1–687.
  27. Kwast, E. (2012) A contribution to the fauna of Cynipidae (Insecta, Hymenoptera, Cynipidae) of Croatia with a description of an asexual female of Andricus korlevici (Kieffer, 1902) nov. Comb. Natura Croatica, 21, 223–245.
  28. Liljeblad, J. & Ronquist, F. (1998) A phylogenetic analysis of higher-level gall wasp relationships (Hymenoptera: Cynipidae). Systematic Entomology, 23, 229–252. DOI:
  29. Massa, B., Cerasa, G. & Pagliano, G. (2021). Insecta Hymenoptera Cynipidae. In: Bologna, M.A., Zapparoli, M., Oliverio, M., Minelli, A., Bonato, L., Cianferoni, F. & Stoch, F. (Eds.), Checklist of the Italian Fauna. Version 1.0. Last update 31 May 2021. Available from: (accessed 20 January 2023)
  30. Mayr, G. (1882) Die europäischen Arten der gallenbewohnenden Cynipiden. Jahresberichte der Communal-Oberrealschule im I. Bezirke, Wien, 21, 1–44.
  31. Marković, Č. (2022) Survey of Cynipid Gall Wasps (Hymenoptera, Cynipidae) in Serbia. Journal of the Entomological Research Society, 24 (2), 177–193. DOI:
  32. Melika, G. (2006) Gall Wasps of Ukraine. Cynipidae. Vestnik zoologii, Supplement 21 (1–2), 1–300, 301–644.
  33. Melika, G., Pujade-Villar, J., Abe, Y., Tang, C.T., Nicholls, J.A., 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. DOI:
  34. Mutun, S., Dinç, S., Bozsó, M., Melika, G. (2014) Four new species of Andricus Hartig oak gall wasp from Turkey (Hymenoptera: Cynipidae, Cynipini). Zootaxa, 3760 (2), 241–259. DOI:
  35. Nicholls, J.A., Melika, G., DeMartini, J. & Stone, G.N. (2018) New species of Dryocosmus Giraud gallwasps from California (Hymenoptera: Cynipidae: Cynipini) galling Chrysolepis Hjelmq. (Fagaceae). Zootaxa, 4532 (3), 407–433. DOI:
  36. Polidori, C. & Nieves-Aldrey, J.L. (2014) Diverse filters to sense: great variability of antennal morphology and sensillae equipment in gall-wasps (Hymenoptera: Cynipidae). PloS ONE, 9 (7), e101843.
  37. DOI:
  38. Pujade-Villar, J., Bellido, D., Segu, G. & Melika, G. (2001) Current state of knowledge of heterogony in Cynipidae (Hymenoptera: Cynipoidea). Sessió Conjunta d’Entomologia, 11, 87–107.
  39. Pujade-Villar, J., Folliot, R. & Bellido, D. (2003) The life cycle of Andricus hispanicus (Hartig, 1856) n. stat., a sibling species of A. kollari (Hartig, 1843) (Hymenoptera: Cynipidae). Butlletí de la Institució Catalana d’Història Natural, 71, 83–95. Available from: (accessed 31 May 2021)
  40. Pujade-Villar, J., Tavakoli, M., Melika, G. & Ferrer-Suay, M. (2015) Andricus synophri (Hymenoptera: Cynipidae), a new species of oak gallwasp from Iran. Journal of Insect Biodiversity and Systematics, 1 (1), 1–10.
  41. Rohwer, S.A. & Fagan, M.M. (1917) The Type-species of the Genera of the Cynipoidea, or the Gall Wasps and parasitic Cynipoids. Proceedings of the United States Natural History Museum, 53 (2208), 357–380. DOI:
  42. Ronquist, F. & Nordlander, G. (1989) Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, supplement 33, 1–60.
  43. Seltmann, K.C., Yoder, M.J., Mikó, I., Forshage, M., Bertone, M.A., Agosti, D., Austin, A.D., Balhoff, J.P., Borowiec, M.L., Brady, S.G., Broad, G.R., Brothers, D.J., Burks, R.A., Buffington, M.L., Campbel, H.M., Dew, K.J., Ernst, A.F., Fernández-Triana, J.L., Gates, M.W., Gibson, G.A.P., Jennings, J.T., Johnson, N.F., Karlsson, D., Kawada, R., Krogmann, L., Kula, R.R., Mullins, P.L., Ohl, M., Rasmussen, C., Ronquist, F., Schulmeister, S., Sharkey, M.J., Talamas, E., Tucker, E., Vilhelmsen, L., Ward, P.S., Wharton, R.A. & Deans A.R. (2012) A hymenopterists’ guide to the Hymenoptera Anatomy Ontology: utility, clarification, and future directions. Journal of Hymenoptera Research, 27, 67–88. DOI:
  44. Sottile, S., Cerasa, G., Massa, B. & Lo Verde, G. (2022) Andricus cydoniae Giraud, 1859 Junior Synonym of Cynips conifica Hartig, 1843, as Experimentally Demonstrated (Hymenoptera: Cynipidae: Cynipini). Insects, 13 (2), 200.
  45. DOI:
  46. 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.
  47. DOI:
  48. 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. DOI:
  49. Stone, G.N., Schönrogge, K., Atkinson, R.J., Bellido, D. & Pujade-Villar, J. (2002) The population biology of oak gall wasps (Hymenoptera: Cynipidae). Annual Review of Entomology, 47, 633–668. DOI:
  50. Tavakoli, M., Melika, G., Sadeghi, S.E., Pénzes, Z., Assareh, M.A., Atkinson, R.S., Bechtold, M., Miko, I., Zargaran, M.R., Aligolizade, D., Barimani, H., Bihari, P., Fülöp, D., Somogyi, K., Challis, R., Preuss, S., Nicholls, J. & Stone, G.N. (2008) New species of oak gallwasps from Iran (Hymenoptera: Cynipidae: Cynipini). Zootaxa, 1699 (1), 1–64. DOI:
  51. Walker, P. (2001) Andricus aries does have a sexual generation on Turkey oak. Cecidology, 16, 94–95. DOI:
  52. Walker, P. (2002) Two new records for cynipid oak galls (Cynipidae: Hymenoptera) in Britain. Cecidology, 17, 64–67.
  53. Yoder, M.J., Mikó, I., Seltmann, K.C., Bertone, M.A. & Deans, A.R. (2010) A gross anatomy ontology for Hymenoptera. PLoS ONE, 5 (12), e15991. DOI: