A new genus of oak gallwasps, Zapatella Pujade-Villar & Melika, gen. n., with a description... 75 A new genus of oak gallwasps, Zapatella Pujade-Villar & Melika, gen. n., with a description of two new species from the Neotropics (Hymenoptera, Cynipidae, Cynipini) Juli Pujade-Villar1,†, Paul Hanson2,‡, Claudia A. Medina3,§, Miguel Torres3,|, George Melika4,¶ 1 Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Animal, Avda. Diagonal 645, 08028-Barcelona, Spain 2 Universidad de Costa Rica. Escuela de Biologia. San Pedro, Costa Rica 3 Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Claustro de San Agustín, Villa de Leyva, Colombia 4 Pest Diagnostic Laboratory, Plant Protection & Soil Conservation Directorate of County Vas, Ambrozy setany 2, Tanakajd 9762, Hungary † urn:lsid:zoobank.org:author:94C497E0-C6A1-48BD-819D-FE5A8036BECD ‡ urn:lsid:zoobank.org:author:0826E20B-DFB4-48D5-A80E-6315C1C184F6 § urn:lsid:zoobank.org:author:A4195A74-EBF5-452D-B3E9-AF7E567AD741 | urn:lsid:zoobank.org:author:CEC17E6B-9D4C-47A7-BF81-D0E87DACA7A7 ¶ urn:lsid:zoobank.org:author:A6FEF774-3A77-4058-94E5-E670D880ED69 Corresponding author: George Melika (melikageorge@gmail.com) Academic editor: M. Buffington  |  Received 27 February 2012  |  Accepted 11 June 2012  |  Published 24 July 2012 urn:lsid:zoobank.org:pub:6719AA7E-3C2B-41AE-9DF5-EBCCB0001345 Citation: Pujade-Villar J, Hanson P, Medina CA, Torres M, Melika G (2012) A new genus of oak gallwasps, Zapatella Pujade-Villar & Melika, gen. n., with a description of two new species from the Neotropics (Hymenoptera, Cynipidae, Cynipini). ZooKeys 210: 75–104. doi: 10.3897/zookeys.210.3014 Abstract A new genus of cynipid oak gallwasp, Zapatella Pujade-Villar & Melika, gen. n. (Hymenoptera: Cyn- ipidae: Cynipini), with two new species, Zapatella grahami Pujade-Villar & Melika, sp. n. and Z. nieve- saldreyi Melika & Pujade-Villar, sp. n., is described from the Neotropics. Zapatella grahami, known only from the sexual generation, induces galls in acorns of Quercus costaricensis and is currently known only from Costa Rica. Zapatella nievesaldreyi, known only from the asexual generation, induces inconspicuous galls in twigs of Q. humboldtii, and is known only from Colombia. Diagnostic characters for both new species are given in detail. Five Nearctic species are transferred from Callirhytis to Zapatella: Z. cryptica (Weld), comb. n., Z. herberti (Weld), comb. n., Z. oblata (Weld), comb. n., Z. quercusmedullae (Ash- ZooKeys 210: 75–104 (2012) doi: 10.3897/zookeys.210.3014 www.zookeys.org Copyright Juli Pujade-Villar et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. ReseARCH ARtiCle Launched to accelerate biodiversity research A peer-reviewed open-access journal

Juli Pujade-Villar et al. / ZooKeys 210: 75–104 (2012) 76 mead), comb. n., Z. quercusphellos (Osten Sacken), comb. n. (= Z. quercussimilis (Bassett), syn. n.). A key based on adults for the species belonging to Zapatella is also given. Generic limits and morphological characteristics of Zapatella and closely related genera are discussed. Keywords Cynipini, Zapatella, Callirhytis, Colombia, Costa Rica, taxonomy, morphology, distribution, biology introduction The cynipid gallwasp fauna (Hymenoptera, Cynipidae) of the Neotropical region is very poorly known. Recently it was updated to include 6 tribes, 21 genera and 45 species, of which 41 are native and 4 have been introduced into the region the native fauna includes 17 described species of oak gallwasps and 15 associated in- quiline species (Medianero and Nieves-Aldrey 2011b). The Neotropics, particularly southern Mexico, harbours the greatest diversity of oak species in the New World. Some species are widespread from Mexico to Costa Rica and Panama. At least one clade of red oaks (Section Lobatae of Quercus subg. Quercus L.) is common in Cen- tral America and Colombia (Govaerts and Frodin 1998, Nixon 2006). In Mexico, gallwasps, gallwasps were reported from 11 different red oak species (Pujade-Villar et al. 2009 Melika et al. 2009a, 2011a Pujade-Villar et al. 2011), and on all oaks 157 gallwasp species on 33 oak species are mentioned (Pujade-Villar et al. 2009). A large diversity of oak gallwasps is also known from Panama, where 65 different cynipid galls were found on oaks, and from 45 of these galls, adult gallwasps were reared: Andricus Hartig (12 species), Neuroterus Hartig (9), Dryocosmus Giraud (7), Cynips L. (4), Amphibolips Reinhard (3), Disholcaspis Dalla Torre & Kieffer, Loxau- lus Mayr and Odontocynips Kieffer (with two species each), Callirhytis Förster and Bassettia Ashmead (with one species each) (Medianero and Nieves-Aldrey 2011b). Ten of these species have been described (Medianero and Nieves-Aldrey 2010a, b, 2011a, Medianero et al. 2011a, b). Thus far, only four oak gallwasp species have been described from Costa Rica: Odontocynips hansoni (Pujade-Villar 2009), Andri- cus costaricensis (Melika et al. 2009b), Disholcaspis costaricensis (Melika et al. 2011b) and Coffeikkokos copeyensis (Pujade-Villar et al. 2012) however, there might be more than 30 species in total (Fergusson 1995, Pujade-Villar and Hanson 2006). Another six species are known from Guatemala (Cameron 1883, Kinsey 1936, Weld 1952) and one species from the southern part of Mexico (Kinsey 1937). The evaluation of the Neotropic gallwasp fauna cannot be done without a thor- ough examination of the Nearctic species, especially in the case of establishing new gallwasp genera. The current morphology-based taxonomy of the Nearctic Cynipi- ni, with the last review of genera by Melika and Abrahamson (2002), needs a major revision at both the genus and species levels. Sets of character states used to identify genera/species are often not appreciable for taxonomic purposes in many genera

A new genus of oak gallwasps, Zapatella Pujade-Villar & Melika, gen. n., with a description... 77 plesiomorphic characters were used instead of synapomorphies or autapomorphies, and thus the majority of current Cynipini genera in the Nearctic are polyphyletic, instead of being monophyletic. The new data recently obtained on the phylogeny, phylogeography, evolutionary conservatism of host shifts were not considered in the previous reviews and revisions (Liljeblad et al. 2008, Stone et al. 2009). The validity of some Nearctic species of Callirhytis and there taxonomic position are discussed. Material and methods Adult gallwasps of an undescribed species were reared from acorn galls collected on Quercus costaricensis by the second author (PH) in Costa Rica specimens belonging to yet another species were reared from galls collected on Q. humboldtii by the first author (JPV) together with Claudia A. Medina and Miguel Torres in Colombia. We follow the current terminology of morphological structures (Liljeblad and Ronquist 1998, Melika 2006). Abbreviations for forewing venation follow Ronquist and Nordlander (1989), cuticular surface terminology follows that of Harris (1979). Measurements and abbreviations used here include: F1–F12, 1st and subsequent flag- ellomeres POL (post-ocellar line) is the distance between the inner margins of the posterior ocelli OOL (ocellar-ocular line) is the distance from the outer edge of a posterior ocellus to the inner margin of the compound eye LOL (lateral ocellar line), the distance between lateral and frontal ocelli. The width of the forewing radial cell is measured from the margin of the wing to the Rs vein. Digital images of wasp anatomy were produced with a digital Nikon Coolpix 4500 camera attached to a Leica DMLB compound microscope, followed by processing in CombineZP (Alan Hadley) and Adobe Photoshop 6.0 by the last author (GM). The SEM pictures were taken with a Stereoscan Leica-360 by Palmira Ros-Farré (Barcelona University, Spain) at a low voltage (15KV) and with gold coating the forewing of Z. nievesaldreyi was taken by JPV with a digital camera Cannon SX-210-IS, attached di- rectly to the ocular of a stereomicroscope. Gall images of Z. grahami were taken by P. Hanson galls of Z. nievesaldreyi by the fourth author (M T). The type material is deposited in the following institutions: UB University of Barcelona, Spain (J. Pujade-Villar) PDL Pest Diagnostic Laboratory (the former Systematic Parasitoid Laboratory, SPL), Tanakajd, Hungary (G. Melika) MZUCR Museum of Zoology, University of Costa Rica, San Pedro Costa Rica (Paul Hanson) IAvH Instituto Alexander von Humboldt, Villa de Leyva, Colombia (Claudia Medina).

Juli Pujade-Villar et al. / ZooKeys 210: 75–104 (2012) 78 Results Zapatella Pujade-Villar & Melika, gen. n. urn:lsid:zoobank.org:act:D093C259-5DB1-43AE-A999-0AD53B8F4EA4 http://species-id.net/wiki/Zapatella Figures 1–62 Type species. Zapatella grahami Pujade-Villar & Melika, sp. n. by present designation. Diagnosis. Partially resembles Callirhytis, Bassettia and Plagiotrochus. However, in Zapatella, the malar sulcus is absent mesosoma strongly arched, short, as long as high in lateral view mesoscutum with numerous fine short, interrupted transverse striae with numerous longitudinal anastomosis connecting transverse striae and together forming a net-like, delicately reticulate, irregular sculpture the pronotum laterally delicately reticulate the metascutellum rugoso-reticulate the metanotal trough and the lateral area of the propodeum with dense white setae. In Callirhytis a distinct malar sulcus is present the mesosoma less arched, always at least slightly longer than high in lateral view the transversely orientated rugae on the mesoscutum are much stronger with much fewer anastomoses between them the pronotum with distinct strong rugae laterally the metascutellum rugose, never reticulate the metanotal trough and the lat- eral area of the propodeum without or with very few setae. In Bassettia the mesosoma is strongly compressed dorsolaterally, distinctly longer than broad the head always more massive from above and nearly rounded in anterior view, broader than the mesosoma. In Plagiotrochus the sculpture of the mesopleuron, the shape of propodeal carinae and the length of the prominent part of the ventral spine of the hypopygium are quite different. The most striking characters that differentiates Zapatella from the above- mentioned genera are the long prominent part of the ventral spine of the hypopygium, which is 6.0–8.5 times longer than broad hind coxae with dense white setae on the dorsoposterior surface, while in the other mentioned genera the prominent part of the ventral spine of the hypopygium is very short, at most 2–3 times longer than broad, and hind coxae without dense setae. For more details see also the Discussion. Description. Body, including antennae and legs, predominantly chestnut brown in some species head partially, mesoscutellum and stripes on mesoscutum dark brown to black. Head 1.3–1.5 times as broad as high in anterior view, massive from above and slightly broader than mesosoma. Gena broadened behind eye, as broad as transverse di- ameter of eye malar sulcus absent. Antenna with 11 flagellomeres in female, 13 in male. Mesosoma strongly arched, short, as long as high in lateral view. Pronotum deli- cately reticulate laterally mesoscutum with numerous fine interrupted short trans- verse striae with numerous longitudinal anastomosis connecting transverse striae and together forming a net-like, delicately reticulate, irregular sculpture. Notauli com- plete (only in Z. herberti) or incomplete, extending to 1/2–2/3 length of mesoscu- tum, converging, deep and broad posteriorly [in some species, on first view, notauli seem to be complete however, these are just darker lines, not impressed notauli, e.g. Z. quercusmedullae]. Anterior parallel lines extending to 1/2 length of mesoscutum

A new genus of oak gallwasps, Zapatella Pujade-Villar & Melika, gen. n., with a description... 79 parapsidal lines distinct and broad, starting from posterior margin and extending to 1/2 length of mesoscutum median mesoscutal line present or absent. Mesoscutellum 0.5 times as long as mesoscutum, as long as broad, not or only slightly overhanging metanotum, center of disk reticulate, sides and posterior 1/3–2/3 dull rugose scutel- lar foveae present, indistinctly delimited posteriorly. Mesopleuron uniformly delicate- ly reticulate, smooth and shiny basally. Metascutellum rugoso-reticulate metanotal trough and lateral propodeal area with dense setae. Central propodeal area delimited Figures 1–8. Zapatella grahami 1 head, female (anterior view) 2 head, female (dorsal view) 3 head, female (posterior view) 4 head, male (anterior view) 5 head, male (dorsal view) 6 antenna, female 7 an- tenna, male 8 hind leg, female (arrow indicates the dense white setae on dorsoposterior surface of coxa).