Cynipini, Billberg, 1820

Buffington, Matthew L., Forshage, Mattias, Liljeblad, Johan, Tang, Chang-Ti & Noort, Simon van, 2020, World Cynipoidea (Hymenoptera): A Key to Higher- Level Groups, Insect Systematics and Diversity 4 (4), No. 1, pp. 1-69 : 40-42

Cynipini

Figs. 199–204

The majority of cynipid species belong to this single tribe. Members of the tribe are gall inducers on species of Quercus , Castanea , Castanopsis , Chrysolepis , and Lithocarpus (Fagaceae) , by far the largest number on oaks. Many more species and genera are expected to be discovered and described from Asia where Fagaceae generic diversity is relatively high and little work has been done on the Cynipini fauna until recently (e.g., Tang et al. 2009; Ide et al. 2010; Tang et al. 2011; Melika et al. 2011; Ide et al. 2012; 2013; Melika et al. 2013b; Ide and Abe 2015; Tang et al. 2016). New genera and new species have also been continuously discovered in the Nearctic and Neotropics (e.g., Medianero and Nieves-Aldrey 2011; Medianero et al. 2011; Nieves-Aldrey et al. 2012; Pujade- Villar et al. 2012a, b; Pujade-Villar et al. 2013; Medianero and Nieves-Aldrey 2013), highlighting the fact that the taxonomy of Cynipini is still far from complete. Taxonomy of Cynipini has been controversial, including several species previously classified in Andricus and Callirhytis having been moved between these two genera ( Burks 1979), and several Nearctic Callirhytis species transferred to newly erected genera Kinseyella ( Pujade-Villar et al. 2010) and Zapatella ( Pujade-Villar et al. 2012b) and Melikaiella (Pujade- Villar et al. 2014). Two Palaearctic genera, Belizinella Kovalev and Ussuraspis Kovalev , were synonymized with Trigonaspis ( Melika and Abrahamson 2002) , but then reestablished as valid genera ( Melika 2012 ). Three Nearctic genera, Dros Kinsey , Erythres Kinsey , and Femuros Kinsey , were synonymized with Andricus ( Melika and Abrahamson 2002) but then reestablished as valid genera (Pujade- Villar and Melika 2014 , Pujade-Villar and Ferrer-Suay 2015, Pujade- Villar et al. 2017). Lastly, Sphaeroteras , synonymized with Biorhiza by Melika and Abrahamson (2002), was reestablished as a valid genus ( Pujade-Villar et al. 2018).

Biology. The life cycle of Cynipini involves cyclical parthenogenesis (heterogony), with a sexual generation where males and females mate to produce an asexual generation of only females, that reproduce parthenogenetically and gives rise to a new bisexual generation. The two generations differ in terms of the gall phenology, galling positions, gall structure, and adult morphology and size. These biological distinctions have been led the two generations of the same species classified as different species and even as different genera in the past. Two Palaearctic genera, Andricus and Callirhytis , are host alternators (heteroecy) that their life cycles alternate between two host-plant subgroups, section Cerris and section Quercus sensu stricto in the genus Quercus . Although in some species, the two alternating generations have been associated, for many species they have not yet been matched. Field observation is a firm approach to associate the two generations, however, DNA barcoding is another useful tool to pair the two generations (e.g., Ács et al. 2007, Melika et al. 2013 a, Nicholls et al. 2018). There are only three exceptions in Cynipini known to reproduce purely parthenogenetically. The first case is the global pest, the chestnut gallwasp Dryocosmus kuriphilus Yasumatsu ( Aebi et al. 2006) . This species has one generation per year and has been introduced into Europe, North America, and some countries in Asia outside its native region in China. The other two species, Andricus targionii Kieffer and A. pseudoflos (Monzen) , from Japan, Korea, China and Far East of Russia, are derived from their cyclically parthenogenetic ancestors A. mukaigawae (Mukaigawa) and A. kashiwaphilus Abe through the deletion of sexual generation (Abe 2007). Oak gallwasps have been very conservative on their host-plant choices, and host switches were extremely rare in the evolution of oak gallwasps ( Stone et al. 2009).

Distribution. Holarctic, Neotropical, and Oriental Regions.

Relevant literature. Nieves-Aldrey (2001) reviews the Iberian fauna. Melika and Abrahamson (2002) made a world revision of Cynipini on the basis of Weld’s Cynipoidea monograph ( Weld 1952). Stone et al. (2002) and Csóka et al. (2005) reviewed the general biology, ecology, and evolution of Cynipini , and Abe et al. (2007) reviewed species richness, host-plant diversity, and background on the hypotheses of geographic origin of Cynipini . The Western Palaearctic fauna of Cynipini is reviewed in Melika (2006) on the basis of the Ukrainian fauna. Melika et al. (2010) made a taxonomic revision of species, particularly those that gall section Cerris of Quercus in the Palaearctic and Oriental region. The most recent knowledge of Cynipini in Eastern Palaearctic and Oriental region is reviewed in Pénzes et al. (2018)

Classification

Cynipini Billberg, 1820

Acraspis Mayr, 1881 ; 20 species NA

Amphibolips Reinhard, 1865 ; 53 species NA, NT

Andricus Hartig, 1840 ; ca 375 species but taxonomy is uncertain and far from stable with many more or less dubious taxa NA, PA, OR, NT

Aphelonyx Mayr, 1881 ; 3 species PA

Atrusca Kinsey, 1930 ; 59 species but taxonomy is uncertain NA, NT

Barucynips Medianero and Nieves-Aldrey, 2013 ; 1 species NT Bassettia Ashmead, 1887 ; 9 species NA

Belizinella Kovalev, 1965 ; 2 species ePA

Belonocnema Mayr, 1881 ; 2 species NA

Biorhiza Westwood, 1840 ; 6 species PA, NA

Callirhytis Förster, 1869 ; 80 species but taxonomy is uncertain and far from stable NA, PA

Cerroneuroterus Melika and Pujade-Villar, 2009 ; 9 species PA, OR

Chilaspis Mayr, 1881 ; 2 species wPA

Coffeikokkos Pujade-Villar and Melika, 2012 ; 2 species NT Cyclocynips Melika, Tang and Sinclair, 2013 ; 2 species OR Cycloneuroterus Melika and Tang, 2011 ; 17 species OR, ePA Cynips Linnaeus, 1758 ; ca 75 species but taxonomy is uncertain and far from stable with many more or less dubious taxa NA, PA, NT

Disholcaspis Dalla Torre and Kieffer, 1910 ; 55 species NA, NT Dros Kinsey, 1937 ; 5 species NA

Dryocosmus Giraud, 1859 ; 50 species but diagnostic characters need to be revised and new genera will be erected; PA, NA, OR Erythres Kinsey, 1937 ; 2 species NA

Eumayria Ashmead, 1887 ; 6 species NA

Eumayriella Melika and Abrahamson, 1997 ; 2 species NA Femuros Kinsey, 1937 ; 7 species NA

Heteroecus Kinsey, 1922 ; 12 species NA

Holocynips Kieffer, 1910 ; 5 species NA

Kinseyella Pujade-Villar and Melika, 2010 ; 2 species NA Kokkocynips Pujade-Villar and Melika, 2013 ; 1 species NT Latuspina Monzen, 1954 ; 9 species ePA, OR

Loxaulus Mayr, 1881 ; 17 species NA, NT

Melikaiella Pujade-Villar, 2014 ; 14 species NA

Neuroterus Hartig, 1840 ; 79 species but taxonomy is uncertain and far from stable including more or less dubious taxa; PA, NA, NT

Odontocynips Kieffer, 1910 ; 3 species NA, NT

Philonix Fitch, 1859 ; 7 species NA

Phylloteras Ashmead, 1897 ; 9 species NA

Plagiotrochus Mayr, 1881 ; 22 species PA, OR Protobalandricus Melika, Nicholls and Stone, 2018 ; 1 species NA

Pseudoneuroterus Kinsey, 1923 ; 4 species wPA

Sphaeroteras Ashmead, 1897 ; recently resurrected with 5 species; NA, NT

Striatoandricus Pujade-Villar, 2020 ; 6 speceis NA

Trichagalma Mayr, 1907 ; 3 species ePA, OR

Trigonaspis Hartig, 1840 ; 22 species PA, NA

Ussuraspis Kovalev, 1965 ; 1 species ePA

Zapatella Pujade-Villar and Melika, 2012 ; 13 species NA, NT Zopheroteras Ashmead, 1897 ; 6 species NA