Synergus Hartig species group (Hymenoptera: Cynipidae: Synergini) with partially smooth mesopleurae from the New World Author Lobato-Vila, Irene 0000-0001-5965-9975 Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals, Avda. Diagonal 645, 08028 Barcelona, Spain Author Equihua-Martínez, Armando 0000-0003-4392-3924 Instituto de Fitosanidad, Colegio de Postgraduados, 56230 Montecillo, Texcoco, Estado de México, México & equihuaa @ colpos. mx; https: // orcid. org / 0000 - 0003 - 4392 - 3924 equihuaa@colpos.mx Author Estrada-Venegas, Edith G. 0000-0003-1347-6369 Instituto de Fitosanidad, Colegio de Postgraduados, 56230 Montecillo, Texcoco, Estado de México, México & estradae @ colpos. mx; https: // orcid. org / 0000 - 0003 - 1347 - 6369 estradae@colpos.mx Author Cibrián-Tovar, David 0000-0001-6788-0178 División de Ciencias Forestales, Universidad Autónoma Chapingo, Km 38.5 Carretera México-Texcoco, Chapingo, Estado de México, México & dcibrian 48 @ gmail. com; https: // orcid. org / 0000 - 0001 - 6788 - 0178 dcibrian48@gmail.com Author Barrera-Ruíz, Uriel M. 0000-0002-2451-807X División de Ciencias Forestales, Universidad Autónoma Chapingo, Km 38.5 Carretera México-Texcoco, Chapingo, Estado de México, México & umbr 757 @ gmail. com; https: // orcid. org / 0000 - 0002 - 2451 - 807 X umbr757@gmail.com Author Pujade-Villar, Juli 0000-0001-7798-2717 Universitat de Barcelona, Facultat de Biologia, Departament de Biologia Evolutiva, Ecologia i Ciències Ambientals, Avda. Diagonal 645, 08028 Barcelona, Spain & jpujade @ ub. edu; https: // orcid. org / 0000 - 0001 - 7798 - 2717 jpujade@ub.edu text Zootaxa 2020 2020-08-04 4822 1 1 38 journal article 8791 10.11646/zootaxa.4822.1.1 2bbc92ef-c6a7-4e7a-8da8-5f542394fda1 1175-5326 4450519 668A9E94-DB24-4AD0-B5D7-1A5D3273FC9F Synergus agrifoliae Ashmead, 1896 ( Figure 1 ) Synergus agrifoliae Ashmead, 1896 . Trans. Am. Entomol. Soc. 23: 189. Type material: USNM. Synergus maculatus Fullaway, 1911 . Ann. Entomol. Soc. Am. 4: 371 [not Synergus maculatus Tavares, 1920 ]. Type material: USNM (see remarks). Synonymized by Weld (1952: 337). Synergus obscurus McCracken & Egbert, 1922 . Stanford Univ., Pubs., Univ. Ser. Biol. Sci. 3 (1): 57. Type material: CAS. Synonymized by Weld (1952: 337). Type material of Synergus agrifoliae Ashmead, 1896 (examined) . LECTOTYPE ♀ with the following labels: ‘ Los Angeles Co. CAL .’ (white label) / ‘125’ (white label, handwritten) / ‘ Through C. V . Riley’ (white label) / ‘ Type No. 3289 U.S. N.M’ (red label) / ‘ Synergus agrifoliae Ashm. ’ (white label, handwritten) / ‘USNMENT 00779886’ (white label, QR code) / ‘ Lectotype ♀ Synergus agrifoliae Ashmead, 1896 , IL-V desig-2017’ (red label). PARALECTOTYPES ( 4♂ & 4♀ ) with the following labels: ‘125’ (white label, handwritten) / ‘ Los Angeles Co. CAL .’ (white label) / ‘ Through C. V . Riley’ (white label) / ‘ Type No. 3289 U.S. N.M’ (red label) / ‘ Synergus agrifoliae Ashm. ’ (white label, handwritten) / ‘ Paralectotype Synergus agrifoliae Ashmead, 1896 , IL-V desig-2017’ (red label) ( 4♂ and 2♀ ); ‘ Los Angeles Co. CAL .’ (white label) / ‘ Through C. V . Riley’ (white label) / ‘ Cynipid on Quercus agrifolia , 125’ (white label, handwritten) / ‘ Type No. 3289 U.S. N.M’ (red label) / ‘ Paralectotype ♀ Synergus agrifoliae Ashmead, 1896 , IL-V desig-2017’ (red label) ( 2♀ ). Type material of Synergus maculatus Fullaway, 1911 (examined) . LECTOTYPE ♂ with the following labels: ‘Stanford Univ. Calif. R . W. Patterson’ (white label, handwritten) / ‘cotype’ (pink label) / ‘ Synergus maculatus Full. ’ (white label, handwritten) (x2) / ‘ Lectotype ♂ Synergus maculatus Fullaway, 1911 , IL-V desig-2019’ (red label) / ‘ Synergus agrifoliae Ashmead, 1896 , IL-V det. 2019’ (white label). Type material of Synergus obscurus McCracken & Egbert, 1922 (examined) . LECTOTYPE ♀ with the following labels: ‘L.S.J.U., Lot. 554, Sub. 67’ (white label) / ‘Type’ (red label) / ‘ Synergus obscurus McC & Eg’ (white label with a red frame) / ‘ California Academy of Sciences, Type No. 5805’ (white label) / ‘ Lectotype ♀ Synergus obscurus McCracken & Egbert, 1922 , IL-V desig-2019’ (red label) / ‘ Synergus agrifoliae Ashmead, 1896 , IL-V det. 2019’ (white label). Additional material ( 9♂ & 10♀ ). Non-type material deposited in USNM (examined) with the following labels: ‘Santa Cruz, Calif., Mar.’ (white label, handwritten) / ‘Ex. 1726’ (white label, handwritten) / ‘ Q. agrifolia ’ (white label) / ‘ Synergus obscurus McC + Egb.’ (white label, handwritten) ( 1♂ & 1♀ ); ‘Palo Alto, Cal., 1915’ (white label) / ‘1726’ (white label, handwritten) / ‘ Synergus agrifoliae Ashm. , cwt. Weld 1923’ (white label, handwritten) ( 1♀ ); ‘Pasadena, Cal.’ (white label) / Èx. 1727’ (white label, handwritten) ( 1♀ ); ‘StaCruz, Mts. Cal.’ (white label) / ‘Through C. V . Riley’ (white label) / ‘ Synergus agrifoliae Ashm. , det. Weld 1942’ (white label, handwritten) ( 1♀ ); ‘ Synergus obscurus McC. + Eg.’ (white label, handwritten) / ‘On Q. agrifolia , Stan Univ., Cal. ’ (white label, handwritten) / ‘3B, 1B, Mar. 14, 33’ (white label, handwritten) / ‘ Synergus obscurus McC+ Egb.’ (white label, handwritten) ( 1♂ ); ‘PcificGrove, Cal., Jan 2-18’ (white label, handwritten) ( 1♂ ); ‘SantaBarbara, Cal.’ (white label) / ‘1726’ (white label, handwritten) / ‘ Synergus maculatus Full. , c.w.t. Weld 1923’ (white label, handwritten) ( 1♂ ); ‘Orange, Calif.’ (white label, handwritten) / ‘ Q. agrifolia ’ (white label) / ‘1726’ (white label, handwritten) ( 2♂ & 1♀ ); ‘Hopk. U.S. 15640b’ (white label) / ‘Reared June 26.22 (white label) / ‘ Quercus garryana ’ (white label) / ‘Fort Jones, Calif.’ (white label) / ‘L. H. Weld Collector’ (white label) ( 3♂ & 5♀ ). Diagnosis . Synergus agrifoliae is morphologically close to S. confertus McCracken & Egbert, 1922 and S. walshii Gillette, 1896 (see below), but differs from these species mainly by having the mesoscutum finely coriaceous to reticulated (coriaceous or imbricated, with weak discontinuous transversal elements in the above-mentioned species), mesopleurae basally and medially with spaced striae (with dense striae in in the above-mentioned species) and the body mainly brownish yellow to yellow (mainly black, dark brown or rufous in the above-mentioned species) (see the descriptions of both species and the identification key). TABLE 1. List of the Synergus with partially smooth mesopleurae from the New World, with information regarding their host gall species, host oak species, the host oak’s section* and their distribution. E = East; W = West.

Synergus species	Gall(s)
S. agrifoliae Ashmead	Neuroterus spp.	Q. agrifolia	Lobatae	USA (W)
(= S. maculatus Fullaway , = S. obscurus McCracken & Egbert )	Unknown gall	Q. garryana	Quercus
S. aurofacies Lobato-Vila & Pujade-Villar , sp. nov.	Femuros lusum	Q. laeta	Quercus	Mexico
S. campanula Osten-Sacken	Disholcaspis quercusglobulus	White oaks	Quercus	USA (E)
Disholcaspis quercusmamma	Q. bicolor	Quercus
Trigonaspis ? quercusforticorne ‘ incertae sedis ’	Quercus sp.
S. castanopsidis (Beutenmüller)	Dryocosmus castanopsidis	Chrysolepis chrysophylla		USA (W)
S. cibriani Pujade-Villar & Lobato-Vila	Undetermined spherical galls on leaves	Q. glabrescens	Quercus	Mexico
Disholcaspis spp.	Quercus sp.
S. citriformis (Ashmead) (= S. elegans Nieves-Aldrey & Medianero , = S. niger Fullaway )	Mainly Amphibolips , also woolly galls of Andricus , Stria- toandricus and Cynips , other galls of Andricus , Atrusca , Cynips , Disholcaspis , Dros and Loxaulus	Oaks of both Lobatae and Quercus sections		Mexico Panama USA (E and W)
S. confertus McCracken & Egbert	Andricus confertus	Q. lobata	Quercus	USA (W)
S. flavens McCracken & Egbert (= S. variegatus McCracken & Egbert syn. nov. )	Heteroecus flavens Heteroecus pacificus	Q. vaccinifolia	Protobalanus	USA (W)
Trichoteras coquilletti	Q. chrysolepis
S. gilletti Pujade-Villar & Lobato-Vila	Andricus sp. (woolly galls)	Q. laeta	Quercus	Mexico
Atrusca sp.	Q. rugosa Q. laeta
S. laeviventris (Osten-Sacken)	Amphibolips confluens	Q. rubra	Lobatae	USA (E and W)
Unnamed gall	Q. falcata
Amphibolips quercusspongifica Atrusca quercuscentricola Disholcaspis rubens Disholcaspis quercusglobulus	Quercus sp.
S. laticephalus Nieves-Aldrey & Medianero	Undetermined spherical galls on twigs	Q. salicifolia	Lobatae	Panama

......continued on the next page TABLE 1. (Continued)

Synergus species	Gall(s)
S. longimalaris Pujade-Villar & Lobato-Vila	Amphibolips sp.	Q. conzattii	Lobatae	Mexico
Amphibolips dampfi Amphibolips sp.	Q. affinis
Andricus cylindratum	Q. glabrescens	Quercus
Andricus group tecturnarum Amphibolips sp.	Quercus sp.
S. longiscapus Pujade-Villar & Lobato-Vila	Mainly woolly galls of Andricus , other galls of Andricus , Cynips , Disholcapsis , Neuroterus eugeros , Loxaulus hyali- nus and some undetermined small spherical galls on leaves	Oaks of both Lobatae and Quercus sections		Mexico
S. mendax Walsh	Callirhytis quercuspunctata	Q. velutina	Lobatae	USA (E)
S. pacificus McCracken & Egbert	Heteroecus pacificus	Q. chrysolepis	Protobalanus	USA (W)
(= S. profusus McCracken & Egbert syn. nov. )	Disholcaspis canescens	Q. douglasii	Quercus
S. pomiformis (Ashmead)	Callirhytis quercuspomiformis	Quercus sp.	Lobatae	USA (W)
(= S. flavus Kieffer syn. nov. , = S. varicolor Fullaway syn. nov. )	Amphibolips sp.	Q. agrifolia	Lobatae	Mexico
S. punctatus Gillette	Disholcaspis rubens Disholcaspis eldoradensis Biorhiza eburnea ‘ incertae sedis ’	Oaks of Quercus section		USA (W)
S. stelluli Burnett	Andricus stellulus	Q. dumosa	Quercus	USA (W)
S. striatifrons Pujade-Villar & Lobato-Vila	Amphibolips hidalgoensis	Q. crassipes	Lobatae	Mexico
Amphibolips zacatequensis	Q. eduardi
S. succinipedis (Ashmead)	Disholcaspis quercusvirens	Q. virginiana	Virentes	USA (E)
S. walshii Gillette	Andricus quercusflocci	White oaks	Quercus	USA (E)

* Quercus sections have been assigned following http://oaks.of.the.world.free.fr/liste.htm (Last entry date: May 2020) Redescription. FEMALE. Length . Body length 2.0– 2.4 mm (n = 5). Color ( Fig. 1a, b ). Brownish yellow to yellow. Head yellow, lower face and genae yellow, frons and vertex usually yellow, with some black near ocelli or even with a conspicuous black spot in some specimens; sometimes, frons, vertex and occiput brownish. Antennae yellowish brown to testaceous, the tip slightly darker. Mesosoma mostly yellow to yellowish brown; mesoscutum usually yellow, somewhat dark anteriorly in some specimens; scutellum anteriorly sometimes black; space between coxae black in some specimens; mesopleurae usually yellow, brown to dark brown in some specimens, sometimes only black basally; propodeum usually dark, rarely totally yellow; tegulae yellow. Metasoma yellowish orange, sometimes dorsally darker. Legs mostly light yellow. Wings hyaline, veins pale yellow. Head . In frontal view, almost 1.3 times as wide as high, genae not expanded behind compound eyes. Face faintly pubescent, lower face with striae radiating from clypeus. Clypeus indistinct, ventral margin slightly projected over mandibles. Malar space about 0.7 times as long as height of compound eye. Anterior tentorial pits visible; pleurostomal and epistomal sulcus absent. Transfacial line about 1.2 times as long as height of compound eye. Toruli situated slightly under mid-height of compound eye; distance between torulus and compound eye shorter than diameter of torulus; distance between toruli shorter than diameter of toruli. Frons coriaceous, without punctures; frontal carinae very short and narrow, not reaching lateral ocelli. Head in dorsal view is about 2.1 times as wide as long. Vertex coriaceous, without punctures. POL: OOL: LOL = 12.5: 6: 6.5 and diameter of lateral ocelli, 4. Occiput coriaceous, without punctures. Antennae ( Fig. 1a ). 14-segmented; filiform, not broadened apically; pubescence dense and short. Scape plus pedicel about as long as F1; pedicel 1.4 times as long as wide; F1 about 1.3 times as long as F2, F2 and F3 subequal, the following segments progressively shorter. Last flagellar segment about 2.7 times as long as wide and almost 2.0 times as long as F11. Mesosoma . Slightly longer than high in lateral view, including nucha, with short and not dense pubescence ( Fig. 1a ). Ratio of length of pronotum medially/laterally: 0.4. Pronotal plate indistinct ( Fig. 1b ). Lateral pronotum ( Fig. 1a ) finely coriaceous; lateral margins of pronotum rounded, without lateral carina. Mesoscutum ( Fig. 1b ) about 1.2 times as wide as long, finely coriaceous to reticulated; anterior grooves inconspicuous. Notauli incomplete, narrow, not well impressed. Median groove absent. Parapsidal grooves weakly impressed, reaching tegulae. Scutellum ( Fig. 1b ) rounded, about as long as wide, weakly wrinkled; circumscutellar carina weak, obscured by wrinkles; scutellar foveae small, ovate, shallow, smooth, not well defined posteriorly and separated by a narrow carina. Mesopleurae ( Fig. 1a ) basally and medially with spaced striae, speculum smooth; little pubescent basally. Metapleural sulcus reaching 3/4 of mesopleural height. Propodeum pubescent and weakly sculptured; propodeal carinae straight and parallel. Nucha sulcated dorsally and laterally. Legs . Tarsal claws with a small basal tooth. Wings . Fore wings pubescent with moderately long marginal setae, as long as mesosoma plus metasoma ( Fig. 1a ). Radial cell closed and 2.5 times as long as wide; areolet inconspicuous. Rs+M inconspicuous, not reaching the basal vein. Basal cell with sparsely spaced setae. Metasoma . About as long as head plus mesosoma, almost as long as high in lateral view ( Fig. 1a ). First metasomal segment sulcated dorsally and laterally. Syntergite smooth, anterolateral pubescence composed of a few setae and with a posterodorsal patch of micropunctures somewhat extended laterally (not reaching the half of the syntergite height); not incised dorsodistally, slightly pointed. Hypopygial spine about as long as wide and with a few lateral setae; without apical setae. MALE. Similar to female, except for the following morphological traits: body length 1.5 mm (n = 4). Antennae 15-segmented; F1 slightly curved, medially incised, long, slightly expanded basally and almost straight apically. Metasoma shorter than head plus metasoma. Distribution . USA . Different localities from the state of California ( Ashmead 1896 ; Fullaway 1911 ; McCracken & Egbert 1922 ; and in this work, see the additional material examined). Biology . Synergus agrifoliae was described from specimens reared from a gall ‘not unlike Neuroterus saltatorius (Edwards, 1874) occurring on Quercus agrifolia Née ( Lobatae section)’ ( Ashmead 1896: 189 ). The galls from which the synonyms of S. agrifoliae ( S. maculatus and S. obscurus ) were reared are described by both Fullaway (1911: 371) and McCracken & Egbert (1922: 58) as small round brown galls 2.0 mm in diameter located on upper sides of leaves of Q. agrifolia . Synergus agrifoliae was also reared from galls on Q. garryana Dougl. ex Hook. ( Quercus section), according to the labels of some of the non-type specimens found in USNM (see details under the additional material examined). Remarks . Synergus agrifoliae was originally described from 13 specimens ( Ashmead 1896: 189 ). We located and examined 4♂ and 5♀ belonging to the type series deposited in USNM. As it is not stated in the original description, a lectotype has been designated. Synergus maculatus was described from an unstated number of specimens ( Fullaway 1911: 371 ). We located and examined a single male deposited in USNM; however, based on Fullaway’s description there could be more specimens belonging to the type series. According to Fullaway (1911: 331–332) , the type material of S. maculatus was deposited in the Stanford collection (later absorbed by CAS), but there are not specimens belonging to the type series in this institution (R. Zuparko pers. comm .). As a result, and since the descriptor did not designate a holotype , we have designated the male found in USNM as the lectotype of S. maculatus . Synergus obscurus was described from 4♀ ( McCracken & Egbert 1922: 58 ). We located and examined a single female deposited in CAS. The authors do not specify which female is the holotype , so we designated the single located female as the lectotype of S. obscurus . According to the original description of S. agrifoliae ( Ashmead 1896: 189 ) , females have 13-segmented antennae; however, all the examined females of the type series have 14-segmented antennae (which matches with the descriptions of S. maculatus and S. obscurus ).